Residue Analysis of Stone Artifacts from the Wenquan Paleolithic Site Reveals Early Exploitation of Wild Cereals and Food Processing Functions ofAcheulean Tools in Northern China (70,000–40,000 BP)

Residue Analysis of Stone Artifacts from the Wenquan Paleolithic Site Reveals Early Exploitation of Wild Cereals and Food Processing Functions ofAcheulean Tools in Northern China (70,000–40,000 BP) | 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 Residue Analysis of Stone Artifacts from the Wenquan Paleolithic Site Reveals Early Exploitation of Wild Cereals and Food Processing Functions ofAcheulean Tools in Northern China (70,000–40,000 BP) Yuqian Yan, Wentai Lou, Jingyi Li, Shenglan Peng, Zizhen Zhao, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9007914/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 Wenquan Site is located in Wenquan Town, Ruzhou City, Pingdingshan City, Henan Province, China. It is a Paleolithic site dating from approximately 70,000 to 40,000 years ago, and the unearthed stone tools exhibit distinct Acheulean technical features, including handaxes, cleavers, and scrapers. This study conducted residue analysis on 11 typical artifacts from the Wenquan Site to investigate plant food resource utilization by its ancient inhabitants and the functions of stone tools. Results indicate that starch grains were found on all 11 stone tools (including flakes, large flakes, handaxes, picks, cleavers, scrapers, heavy-duty scrapers, and denticulate tools). These starch grains show a high degree of similarity with those from grasses and rhizomatous plants, such as species from the Triticeae, Paniceae, Coix , Trichosanthes , Setaria , Agropyron , Dioscorea , and Pueraria tribes or genera. The Triticeae starch grains discovered on stone tools from the Wenquan site represent the earliest recorded evidence of Triticeae starch grains in northern China. This indicates that ancient humans at the Wenquan site had already begun to collect and consume grass seeds, providing important physical evidence for the study of the exploitation and utilization of wild cereals prior to their domestication. Moreover, a higher concentration of starch grains from Poaceae and rhizomatous plants was discovered on handaxes, picks, and cleavers, clearly indicating that ancient humans at the Wenquan site between 70,000 and 40,000 years ago used these large-sized tools to process food resources. This finding provides new directions and insights for discussing the functions of large Acheulean tools. Wenquan Paleolithic Site Starch grain analysis Plant utilization Stone tool function Figures Figure 1 Figure 2 Figure 3 Figure 4 1. Introduction During the Paleolithic, ancient populations sustained themselves primarily through gathering and hunting, with plants serving as one of their crucial food sources. Starch, a long-chain compound formed by the linkage of glucose molecules, is widely present in the form of starch granules within the cells of plant organs such as roots, stems, and seeds [ 1 ]. Due to the taxonomic significance of starch granules in botany, they are frequently used in archaeological research to identify ancient plant remains rich in starch [ 2 ]. During the collection and processing of plants, starch granules readily adhere to the surfaces of tools such as stone artifacts, where they can be preserved for extended periods. Through the systematic extraction and identification of these residual starch grains on tool surfaces, archaeologists can infer the functions of the related tools, thereby revealing how early humans exploited plant resources. As one of the key methods in archaeological residue studies, starch grain analysis has been widely applied to prehistoric sites to explore ancient populations’ utilization of plant food resources and the functions of stone tools. Regarding tool function, starch grains from cereals and other gathered plants were identified on stone tools from the Oldenburg LA 77 site in northern Germany(3270–2920 BC), indicating that its inhabitants used stone tools to process a highly diverse of plant foods [ 3 ]. Ma et al. analysed a large number of starch grains and phytoliths recovered from the surfaces of four stone knives unearthed at the Lajia site, demonstrating that one of their functions was crop harvesting [ 4 ]. Regarding plant food resources, starch grains from Triticeae, Fabaceae, and Fagaceae were identified in four Neolithic dental calculus samples from Dehesilla Cave in Spain, providing new insights into dietary patterns at the site [ 5 ]. In China, starch grains from Panicoideae, Triticeae, Job’s tears ( Coix spp.), and underground storage organs were discovered on 10 amphora sherds from the Liujiazhai site, providing evidence for investigating the subsistence economy of migrants in northwestern Sichuan[ 6 ]. Several pottery fu pots unearthed from the early strata of the Hemudu site contained black foodcrusts; residue analysis of one sample revealed starch grains from rice ( Oryza sp.), Triticeae tribe, oak acorn ( Quercus sp.), Papilionoideae beans, and lily bulbs ( Lilium sp.), reflecting the Hemudu people’s exploitation of diverse plant food resources [ 7 ]. Currently, while starch grain analysis has demonstrated significant potential in studying and discussing the plant food resources of prehistoric populations and the functions of artifacts, its application in Paleolithic sites remains relatively limited. In southern Italy and Sicily, starch grain analysis of dental calculus from unearthed human remains revealed starch grains from the Triticeae tribe, Paniceae tribe, Andropogoneae tribe, and Sambucus, providing evidence for the study of plant foods in the Late Paleolithic of this region [ 8 ]. Starch grains from various plants, including Arundo donax (cane), Typha sp., and species in the Triticeae tribe, were extracted from ground stones of the Paleolithic period in the Pontic steppe, revealing the processing behaviors of the past populations toward starchy plants[ 9 ]. In China, starch grain analysis was conducted on some specimens from the Zhanghuokou site. Combined with experimental results and specimen morphology, the analysis indicated that Luonan handaxes might have been used for digging underground roots and tubers [ 10 ]. Starch grains extracted from three grinding tools at the Shizitan site in the middle reaches of the Yellow River (23,000–19,500 BP) revealed that ancient humans intensified the use of Paniceae and tuber plants more than ten thousand years earlier than the Neolithic period [ 11 ]. The Wenquan Site is a large-scale, open-air Paleolithic site where a significant quantity of stone artifacts and animal fossils have been discovered. The stone artifacts include cores, flakes, hammerstones, choppers, scrapers, points, handaxes, picks, cleavers, spheres, and other types. The overall lithic assemblage exhibits characteristics of both a simple core-and-flake lithic industry and tool types commonly associated with the Acheulean tradition, such as handaxes and cleavers, dating to the Late Pleistocene. This study aims to conduct starch grain analysis by extracting residues from stone tools at the Wenquan site to explore how ancient humans utilized these tools 70,000 to 40,000 years ago, as well as the types of plant food resources they processed and utilized with them. 2. Materials and methods 2.1 Sites and Stone Tools The Wenquan Site is located in Wenquan Town, Ruzhou City, Pingdingshan City, Henan Province, China, at geographic coordinates 34°12′16.9″N, 112°37′57.3″E, and an altitude of 236 meters (Fig. 1 ). Since 2020, the Henan Provincial Institute of Cultural Heritage and Archaeology has conducted three archaeological excavations at the site, yielding a substantial assemblage of stone artifacts. These include handaxes, cleavers, picks, large knives, heavy-duty scrapers, and choppers, which date to between 70,000 and 40,000 years ago [ 12 ]. The research materials consist of 11 stone artifacts recovered from the Wenquan Site during the 2022 and 2023 excavations. These include flake tools, Acheulean implements, scrapers, and a denticulate(Fig. 2 ). The flake assemblage comprises four specimens: Flake (23wq:1480), Large Flake 1 (23wq:563), Large Flake 2 (23wq:1032), and Large Flake 3 (23wq:1344). The Acheulean tools are represented by three pieces: a Handaxe (22wq:160), a Pick (23wq:908), and a Cleaver (22wq:222). The scraper category includes Heavy-duty Scraper 1 (23wq:1138), Heavy-duty Scraper 2 (23wq:1341), and Scraper (23wq:965). Finally, the denticulate is represented by a single specimen: Denticulate (23wq:879). Detailed classification and descriptions of these stone artifacts are provided in Supplementary Material 1. 2.2 Sample collection This study followed the residue extraction and laboratory processing methods established in published studies [ 13 – 15 ]. In the processes of sample collection and pre-treatment, we consistently maintained a contamination-free environment for both the stone artifacts and the laboratory to prevent sample contamination. The main measures we took included: Sample preservation phase: From excavation to storage in the repository, the stone artifact samples were kept in contamination-isolated sample bags at all times. Laboratory sampling phase: During the entire residue extraction process from stone tool surfaces, a disposable toothbrush head was used for each individual sample at every step, and powder-free gloves were worn to prevent cross-contamination. Experimental processing phase: To prevent contamination during residue handling, laboratory personnel wore laboratory coats, masks, and powder-free gloves throughout the entire process. Contamination control phase: To account for potential airborne starch grain contamination in the sampling environment which could interfere with experimental results, Purified water was added to 10 cm Petri dishes, which were then exposed to ambient air for three days, serving as contamination controls. Sample Collection: A zoned approach was adopted, with sampling limited to one-half of the working edge surface area of each stone tool. The residue sampling procedure consisted of three steps:1. Collecting soil samples adhering to the tool surface by scraping with clean implements. 2.Cleaning the tool surfaces with purified water and a toothbrush to remove potential contaminants. 3.Repeatedly agitating and scrubbing the working edge surfaces with purified water using an ultrasonic toothbrush to collect ultrasonic samples. Some stone artifacts lacked adhering soil samples, so only ultrasonic samples were extracted from them. In total, 11 stone artifacts were selected for surface residue sampling in this study, yielding 19 residue samples (Table 1 ). Laboratory Processing of Samples: Pretreatment and slide preparation of the samples were conducted at the Henan Provincial International Joint Laboratory of Modern Human Origins. Three of these samples were processed at the Laboratory of Archaeometry at Xiamen University. The laboratory pretreatment and slide preparation procedures followed established protocols [ 16 , 17 ] and were as follows: 1) Add deionized water to the tube, balance the centrifuge tubes, and centrifuge at 2000 rpm for 5 min. Discard the supernatant to concentrate the sample. 2) Add a 5% (NaPO 3 ) 6 solution and place the tube on a shaker for 2 h to disperse sample particles. Then centrifuge at 5000 rpm for 5 min and discard the supernatant. 3) Add a heavy-liquid solution of sodium polytungstate (density 2.4 g/cm 3 ). Agitate using a vortex mixer for 10 seconds, then centrifuge at 2000 rpm for 5 min. Using a micropipette, carefully extract the floating fraction (containing lighter plant remains such as starch grains) and transfer it to a new centrifuge tube. 4) Add deionized water, centrifuge at 5000 rpm for 5 min, and discard the supernatant. The resulting sediment contains the plant remains fraction. Repeat this step three times to thoroughly remove residual sodium polytungstate heavy liquid. 5) Extract a portion of the suspension containing plant remains and place a drop in the center of a clean glass microscope slide. Add a drop of 50% glycerol solution, cover with a coverslip, and seal the edges with nail polish. Allow the slide to dry before microscopic examination. Table 1 Information for Stone Tool Samples studied from the Wenquan Site Specimen ID Lab ID Artifact Type Sample Description Sampling Location 23wq:1138 S1 Heavy-duty scraper Surface Soil Blade edge 23wq:1138 S2 Heavy-duty scraper Ultrasonicated Blade edge 23wq:1344 S3 Large flake Surface Soil Blade edge 23wq:1344 S4 Large flake Ultrasonicated Blade edge 23wq:1032 S5 Large flake Surface Soil Blade edge 23wq:1032 S6 Large flake Ultrasonicated Blade edge 23wq:1341 S7 Heavy-duty scraper Surface Soil Blade edge 23wq:1341 S8 Heavy-duty scraper Ultrasonicated Blade edge 23wq:563 S9 Large flake Surface Soil Blade edge 23wq:563 S10 Large flake Ultrasonicated Blade edge 22wq:160 S11 Handaxe-like Surface Soil Blade edge 22wq:160 S12 Handaxe-like Ultrasonicated Blade edge 23wq:908 S13 Pick Surface Soil Blade edge 23wq:908 S14 Pick Ultrasonicated Blade edge 22wq:222 S15 Cleaver Surface Soil Blade edge 22wq:222 S16 Cleaver Ultrasonicated Blade edge 23wq:965 S17 Scraper Ultrasonicated Blade edge 23wq:879 S18 denticulate Ultrasonicated Blade edge 23wq:1480 S19 Flake Ultrasonicated Blade edge 2.3 Sample Microscopic Examination A total of 107 slides were prepared from the 21 samples in this study. Microscopic examination was conducted using a Sunny Optical biological microscope (model EX31) at the Henan Provincial International Joint Laboratory on Origins of Modern Humans in East Asia and a Zeiss Scope A1 biological microscope at the Laboratory of Archaeometry at the Xiamen University. Photomicrography and data measurement were performed using imaging systems and measurement software integrated with the respective microscopes. Identification of residues, particularly starch grains, followed published research on Chinese starch grain morphology[ 18 – 20 ]. Starch grains were identified mainly according to their morphological characteristics, including size, shape, position of the hilum, presence of fissures and configuration, and extinction cross features under polarized light[ 21 ]. 3. Results and analysis 3.1 Contamination assessment A total of 205 starch grains were identified across the 19 samples obtained from 11 stone tools from the Wenquan site. Among these, 32 grains were found in the surface soil samples, and 173 grains were found in the ultrasonic samples. Furthermore, no starch grains were detected in the control samples, indicating that the sampling and slide preparation processes were free from contamination, and thus the starch grains discovered in this study are of ancient origin. The small quantity of starch grains found in the surface soil samples reflects their presence in the burial environment. However, their number is substantially lower than that found in the ultrasonic samples, demonstrating that the starch grains in the ultrasonic samples originate from residues preserved on the artifacts themselves, rather than from post-depositional contamination in the burial context. Consequently, the starch grains identified in the ultrasonic samples serve as the primary subjects for analysis in this study, while those from the surface soil samples serve as a comparativce reference. The detailed classification and comparison of starch grains from the surface soil samples and ultrasonic samples can be found in Supplementary material 2. 3.2 Identification and statistical analysis Among the starch grains discovered in this study, 159 grains were identifiable, accounting for 77.6% of the total starch grains found. Based on morphological characteristics, these were classified into eight types (Fig. 3 ). The remaining 46 grains could not be identified due to morphological damage or lack of diagnostic features (Table 2 ,Table 3 ). To enable accurate quantitative analysis, this study employed three quantitative indicators: the absolute count of starch grains, percentage, and occurrence frequency. The absolute count refers to the total number of starch grains counted for each taxon. The percentage is the proportion of the absolute count of a specific starch grain type relative to the total absolute count of all starch grains. The occurrence frequency was calculated using the formula: (Number of artifacts bearing a specific starch grain type / Total number of artifacts) × 100%. Table 2 Statistical Analysis of Starch Grains from Ultrasonicated Residues at the Wenquan Site Category Triticeae Paniceae Coix Trichosanthes Setaria Agropyron Dioscorea Pueraria Unidentified Total Damaged Grains Starch Morphotype Ⅰ Ⅱ Ⅲ Ⅳ Ⅴ Ⅵ Ⅶ Ⅷ - - - S2 - - 1 - - - - - - 1 - S4 16 12 13 4 1 8 - 2 16 72 16 S6 1 3 1 - 2 - - - 3 10 - S8 7 4 2 2 8 - 1 - - 24 - S10 - - - 1 - - - - - 1 - S12 - - 1 1 1 1 - - 1 5 - S14 - 7 - - - - - - - 7 1 S16 2 1 1 - - - 1 - 7 12 7 S17 2 1 1 - 7 1 1 - 10 23 5 S18 2 - - 2 1 - - - - 5 - S19 5 1 1 - - 1 - - 5 13 7 Total 35 29 21 10 20 11 3 2 42 173 36 Percentage (%) 20.23 16.76 12.14 5.78 11.56 6.36 1.73 1.16 24.3 - 20.81 Frequency (%) 63 63 72 45 54 36 27 0.9 54 - 45 Table 3 Statistical Analysis of Starch Grains from Surface Soil Residues at the Wenquan Site Category Triticeae Paniceae Coix Trichosanthes Setaria Agropyron Dioscorea Pueraria Unidentified Total Damaged Grains Starch Morphotype Ⅰ Ⅱ Ⅲ Ⅳ Ⅴ Ⅵ Ⅶ Ⅷ - - - S1 1 - 1 - 1 - 1 - - 4 - S3 - 3 1 - 2 - - - 4 10 3 S5 - - - - - - - - 2 2 1 S7 - 1 1 - - - - - 1 3 - S9 1 - - 1 - - - - - 2 - S11 1 - - 1 - - 1 - - 3 - S13 - - - - 1 1 - - 2 4 - S15 - - - 3 - - 1 - - 4 - Total 3 4 3 5 4 1 3 0 9 32 4 Percentage (%) 9.38 12.5 9.38 15.63 12.5 3.13 9.38 0 28.13 - 12.5 Frequency(%) 37 25 37 37 37 12 37 0 50 - 25 Type I starch grains totaled 38, with 35 recovered from ultrasonic samples and 3 from surface soil samples. Type I starch grains are generally large, with a size range of 13.79–29.61 µm and a mean size of 21.52 µm. The shapes are subcircular or oval, with the hilum located centrally, and faint lamellae are visible. The extinction cross arms appear as "+" or "X" shapes. Based on these morphological characteristics and comparisons with modern reference collections, Type Ⅰgrains are identified as originating from the tribe Triticeae. Type II starch grains totaled 33, with 29 from ultrasonic samples and 4 from surface soil samples. Type II starch grains are relatively small, with a size range of 9.78–20.63 µm and a mean size of 14.50 µm. The shapes are subcircular or polygonal, with a visible central hilum; some grains exhibit "V" or "Y" shaped fissures. The extinction cross arms are perpendicular, forming a "+" shape. These morphological features resemble starch grains from seeds of the tribe Paniceae [ 22 ]. Additionally, these starch grains could potentially originate from the stems or leaves of the subfamily Panicoideae plants [ 23 ]. Type III starch grains totaled 24, with 21 from ultrasonic samples and 3 from surface soil samples. Type III starch grains are predominantly single grains, with a size range of 11.30–27.12 µm and a mean size of 18.33 µm. Shapes are subcircular or oval, sometimes with rounded rectangular forms. The hilum is central, and fissures are "V", "Y", or "—" shaped. Under polarized light, they exhibit strong extinction with mutually perpendicular extinction arms that display a Z-shaped bend near the edges. Their morphological characteristics resemble starch grains from Job's tears [ 2 , 24 , 25 ], hence they are identified as starch grains from the Coix . Type IV starch grains totaled 15, with 10 from ultrasonic samples and 5 from surface soil samples. These grains have a size range of 9.64–24.77 µm (mean: 14.81 µm) and exhibit diverse shapes, including subcircular, oval, bell-shaped, and polygonal forms. The hilum is eccentric, lamellae are mostly invisible, and the extinction cross arms are bent or exhibit a certain curvature [ 11 ]. Based on morphological characteristics, Type IV grains are identified as originating from Trichosanthes (snake gourd). Type V starch grains totaled 24, with 20 from ultrasonic samples and 4 from surface soil samples. Type V starch grains are small, with a size range of 6.02–10.99 µm and a mean size of 8.25 µm. Their morphology is similar to Type II but smaller in size, being subcircular to oval in shape with a mostly visible, central hilum. The extinction cross arms are perpendicular, appearing as a "+" shape. Based on morphological characteristics, this type is identified as starch grains from Setaria . Type VI starch grains totaled 12, with 11 from ultrasonic samples and 1 from surface soil samples. These grains are relatively large, with a size range of 14.39–34.45 µm and a mean size of 26.02 µm. They are predominantly single grains, subcircular in shape, with a central hilum; some grains visible growth rings. The extinction cross is diffuse and appears as an "X" shape [ 2 ]. Based on these morphological characteristics, Type VI grains are identified as originating from Agropyron . Type VII starch grains totaled 6, with 3 from ultrasonic samples and 3 from surface soil samples. Type VII starch grains are large, with a size range of 18.80–49.99 µm and a mean size of 28.34 µm. Shapes are mostly oval or rounded triangular, with a few irregular forms; the hilum is eccentric, often located near one end. Under polarized light, extinction is strong, and the extinction cross arms are non-perpendicular and bent. Based on these characteristics, Type VII is identified as starch grains from Dioscorea . Type VIII starch grains totaled 2, both from ultrasonic samples. Type VIII starch grains have a size range of 21.21–22.69 µm and a mean size of 21.96 µm. The morphology is polygonal, with a visible central hilum; one grain exhibits a "—" shaped fissure at the hilum, and the extinction cross arms are bent. The morphological characteristics of Type VIII starch grains resemble those from tubers of the genus Pueraria and support their identification as such [ 26 ]. A total of 40 damaged starch grains were observed across the residue samples, with 36 recovered from ultrasonic samples and 4 from surface soil samples. The damaged starch grains in the ultrasonic samples account for 20.81% of the total ultrasonic sample grains, with a frequency of occurrence of 45%. These damaged starch grains exhibit features such as breakage, rough edges, and central depressions, which are consistent with mechanical damage caused by grinding [ 27 ]. This pattern suggests that grinding processing was likely applied to the processed materials(Fig. 4 ). 4. Discussions 4.1 Utilization of plant resources by past population at the Wenquan site A total of 173 starch grains, belonging to eight taxonomic groups, were identified across the 19 samples obtained from the 11 stone artifacts examined at the Wenquan Site. These groups include Triticeae, Paniceae, Coix, Trichosanthes, Setaria, Agropyron, Dioscorea, and Pueraria. Among these, starch grains from grasses were the most abundant, including those from the Panicoideae (Paniceae, Coix, Setaria) (70 grains) and the Triticeae (Triticeae, Agropyron (within Triticeae)) (46 grains), totaling 116 grains. This accounts for approximately 67% of the identifiable starch grains, indicating that grasses constituted a significant proportion of the diet of the ancient inhabitants at the Wenquan site. Furthermore, 15 starch grains were identified from rhizomatous plants (Trichosanthes, Dioscorea, and Pueraria), accounting for about 8.7% of the identifiable starch grains. Current research indicates that grasses and rhizomatous plants have been widely discovered at Paleolithic sites across China, highlighting their importance as key food resources for ancient human populations during the Chinese Paleolithic period. In northern China, the earliest evidence for the collection and consumption of wild grasses by prehistoric humans dates back to the Shizitan Site Complex in Jixian County, Shanxi Province. Residue analysis of stone tools from Shizitan Locality 29 (ca. 28,000–13,000 cal. BP), Locality 14 (ca. 23,000–19,500 cal. BP), and Locality 9 (ca. 13,800–8,500 cal. BP) revealed starch grains from wild grass seeds, primarily from tribes Triticeae and Paniceae (subfamilies Pooideae and Panicoideae) [ 28 – 30 , 16 ]. Starch grains from Triticeae and Panicoideae grasses have also been found on stone artifacts excavated from the Gezishan site in Qingtongxia, Ningxia. This indicates that around 12,000–8,000 years ago, the ancient inhabitants of Gezishan had already begun exploiting wild grass seeds, with grasses dominating their plant resource utilization [ 31 ]. In southern China, large numbers of Triticeae and Paniceae starch grains were found on shell tools unearthed from the Xianren Cave and Diaotonghuan sites in Wannian County, Jiangxi Province. These findings demonstrate that as early as approximately 20,000 years ago, the inhabitants of the these sites were gathering and consuming grass plants, including those from the Triticeae and Paniceae[ 32 ]. The discovery of Triticeae starch grains in dental calculus from hominins at the Fuyan Cave in Daoxian, Hunan, extends the earliest evidence for the consumption of wild grass seeds to approximately 80,000 years ago[ 33 ]. Trichosanthes, Dioscorea, and Pueraria all produce edible roots and tubers. Their underground storage organs are rich in starch, providing a high caloric return with relatively low foraging effort, making them prime targets for prehistoric collection and utilization. At the Gantangqing site in Jiangchuan, Yunnan, underground corms and rhizomes of aquatic plants were discovered. Combined with the presence of wooden tools exhibiting clear use-wear traces at the site (approximately 361,000–250,000 years ago), researchers propose that the hominins at Gantangqing used fabricated tools of selected wood to exploit underground tubers, rhizomes, or corms[ 34 ]. On handaxe unearthed from the Zhanghuokou site in Luonan, Shaanxi (approximately 250,000–100,000 years ago), definitive starch grains of rhizomatous plants have been discovered. This finding also provides evidence for investigating the utilization of rhizomatous plants by early hominin [ 10 ]. Starch grain analysis of stone artifacts from Locality 2 of the Shuidonggou site in Lingwu, Ningxia, indicates that during the MIS 3, wild grass seeds, underground storage organs, and wild Fabaceae were all incorporated into the subsistence strategies of the hominins at the site[ 17 ]. Similarly, starch grains of rhizomatous plants have been found on stone artifacts unearthed from the Shizitan site in Jixian County, Shanxi (ca. 28,000–8,500 cal. BP) [ 28 – 30 , 16 ]. Starch grain analysis of soil samples from the Paleolithic strata (approximately 17,000–13,000 years ago) at the Qihe Cave site in Zhangping, Fujian, has identified starch grains from plants of Pueraria , suggesting that Pueraria served as a major source of starchy food for the inhabitants of the site during the late Pleistocene [ 26 ]. Our study demonstrates that in northern China, Triticeae plants had a long history of exploitation and utilization prior to their domestication. As early as 70,000–40,000 years ago, ancient hominins at the Wenquan site had already begun collecting and consuming wild grass seeds and rhizomatous plants. The gathering and consumption of grasses constituted a significant component of the subsistence economy of the inhabitants of the Wenquan site. 4.2 Functional study of stone tools at the Wenquan site The lithic artifacts from the Wenquan site are predominantly made of quartzite and andesite, with tool production achieved primarily through hard-hammer percussion. Typological analysis reveals the presence of small scrapers and simple core-chopper tools at the Wenquan site, alongside large cutting tools such as handaxes, cleavers, picks, and large knives, representing characteristic of Acheulean-type lithics[ 12 ].In this study, 11 stone tools were selected for the extraction and analysis of surface residues, including flakes (both regular and large flakes), Acheulean large tools (handaxes, picks, and cleavers), scrapers (both regular and heavy-duty scrapers), and denticulates. Based on the quantity and identification results of the starch grains, we conducted a preliminary discussion on how the early inhabitants of the Wenquan site used these stone tools. Based on the absolute counts, morphological types, and presence of damaged starch grains found on the sampled stone tools, all 11 tools were likely used to collect or process at least one or multiple plant types (Table 4 ). For the flakes, the assemblage includes one regular flake and three giant flakes. On flake (23wq:1480) and giant flake 2 (23wq:1032), starch grains from grass plants were identified. Giant flake 1 (23wq:563) yielded starch grains from rhizome plants. Giant flake 3 (23wq:1344) was found contain starch grains from both grass and rhizome plants. Notably, this stone tool yielded the highest quantity and most diverse types of starch grains, with 72 grains identified from the ultrasonicated sample, belonging to six distinct types. These results suggest that the Wenquan inhabitants did not perform secondary modification to the flakes after initial production. Instead, they used them directly for harvesting grass plants and excavating rhizomatous plants. Regarding the Acheulean large tools, these include handaxes, picks, and cleavers. Starch grains from both grass plants and rhizomatous plants were identified on handaxe 22wq:160 and cleaver 22wq:222. On pick 23wq:908, only starch grains from grass plants were found. The results of the starch grain analysis indicate that the ancient humans of the Wenquan Site likely used these Acheulean large tools (handaxes, picks, and cleavers) to process and handle both grass plants and rhizomatous plants. Guan Ying et al. [ 10 ] previously reported starch grains from rhizomatous plants on handaxes from the Zhanghuokou site in Luonan, Shaanxi Province, and proposed that such tools may have been used for digging the underground storage organs of plants. Our findings align with and extend this interpretation: large tools from the Acheulean cultural tradition at Wenquan could have served for cutting and harvesting starch-rich grasses and rhizomes. These results further demonstrate the multifunctional nature of large Paleolithic tools such as handaxes and picks. They were not limited to woodworking but were also employed for exploiting and processing starch-containing grasses and rhizomes. Regarding the scrapers, the assemblage includes both regular scrapers and heavy-duty scrapers. A relatively high quantity of starch grains was recovered from two scrapers: regular scraper 23wq:965 and heavy-duty scraper 23wq:1341. These included grains from Triticeae, Paniceae, Coix , and others taxa. Overall, the highest quantities of starch grains among all sampled tools were found on the scrapers. On Heavy-duty Scraper 1 (23wq:1138), starch grains from grass plants were identified. In contrast, Heavy-duty Scraper 2 (23wq:1341) and regular scraper (23wq:965) yielded starch from both grass plants and rhizomatous plants, with comparatively high counts on these two artifacts. The results suggest that scrapers at the Wenquan Site were likely used for collecting and processing wild grass seeds and rhizomatous plants. Regarding the denticulate, only one denticulate(23wq:879) was sampled. The starch grains recovered from it resemble those from Triticeae, Trichosanthes, and Setaria plants. This suggests that the denticulate may have been used for handling and processing both grass plants and rhizomatous plants. Table 4 Statistical Analysis of Starch Grains from Stone Tool Residues at the Wenquan Site Tool Type Artifact ID Sample ID Sample Description Identified Starch Taxa Flakes Flake 23wq:1480 S19 Ultrasonicated Triticeae、Paniceae、Coix、Agropyron Large flake 1 23wq:563 S10 Ultrasonicated Trichosanthes Large flake 2 23wq:1032 S6 Ultrasonicated Triticeae、Paniceae、Coix、Setaria Large flake 3 23wq:1344 S4 Ultrasonicated Triticeae、Paniceae、Coix、Trichosanthes、Setaria、Agropyron、Pueraria Acheulean large tools handaxe 22wq:160 S12 Ultrasonicated Coix、Trichosanthes、Setaria、Agropyron Pick 23wq:908 S14 Ultrasonicated Paniceae Cleaver 22wq:222 S16 Ultrasonicated Triticeae、Paniceae、Coix、Dioscorea Scrapers Heavy-duty scraper 1 23wq:1138 S2 Ultrasonicated Coix Heavy-duty scraper 2 23wq:1341 S8 Ultrasonicated Triticeae、Paniceae、Coix、Trichosanthes、Setaria、Dioscorea Scraper 23wq:965 S17 Ultrasonicated Triticeae、Paniceae、Coix、Setaria、Agropyron、Dioscorea Denticulate Denticulate 23wq:879 S18 Ultrasonicated Triticeae、Trichosanthes、Setaria 5. Conclusions This study conducted residue extraction and analysis on 11 stone tool from the Wenquan site in Ruzhou, Henan Province. Starch grains assignable to eight plant types were identified, including those from the tribes Triticeae and Paniceae, as well as genera such as Coix , Trichosanthes , Seteria, Agropyron , Dioscorea , and Pueraria . These findings indicate that wild grass seeds and root/tuber plants (rhizomatous plants) were important plant food resources for the local Paleolithic inhabitants. The inhabitants of the Wenquan Site used stone tools to collect and process wild grass seeds and rhizomatous plants available in their environment. This indicates that, early in the Paleolithic period, humans in the Central Plains region consciously incorporated wild grains into their diet. Furthermore, the diverse starch grain types recovered from the surfaces of various stone tools, including scrapers, handaxes and picks provide new evidence and insights into the functional roles of these stone tools at the Wenquan site. In particular, the presence of grass starch grains on Acheulean large tools, such as handaxes and picks, provides fresh perspectives on the use patterns and versatility of such tools in early Paleolithic subsistence strategies. Declarations Acknowledgements This study was supported by the Henan Cultural Relics Protection Research Project (Grant No. 25HNWWJ—KJ01), the Henan Provincial Department of Finance Basic Scientific Research Operating Funds Project (2025HKYJKY01), the Archaeological Talent Promotion Program of China (Grant No. 2025-191), the Chongqing Municipal Education Commission Science and Technology Youth Project (KJQN202500536), the Henan Social Science Project “Research on Late Pleistocene Lithic Technology in the Central Plains Based on the Ruzhou Wenquan Site”, and the Henan Provincial Institute of Cultural Heritage and Archaeology Special Conservation Project “Archaeological Survey and Exploration of the Ruzhou Wenquan Paleolithic Site”. Author contributions Yuqian Yan, Wentai Lou, Jingyi Li, and Wei Ge conducted the experiments and data analysis. Yuqian Yan wrote the manuscript. Wentai Lou, Qingpo Zhao, Huanhuan Ma, and Wei Ge revised the paper. Shenglan Peng, Zizhen Zhao, Jia Huang, and Jun Chen participated in the research and were responsible for the organization of lithic materials and the compilation of supplementary data. All authors reviewed and approved the final manuscript. Competing interests The authors declare no competing interests. Data availability The authors confirm that the data supporting the findings of the study are available within the article and supplementary materials. References Yang X,Lü H,Xia Z.The application of plant starch grain analysis in archaeology.Archaeology and Cultural Relics,2006,(03): 87-91. Ge W.The application of starch grain analysis in archaeology. University of Science and Technology of China, 2010. An J P, Chevalier A, Brozio J P, Müller J, Kirleis W. 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Harvesting and processing wild cereals in the Upper Palaeolithic Yellow River Valley, China, Antiquity. 2018. 92(363), 603-619. https://doi.org/10.15184/aqy.2018.36 Liu L, Bestel S, Shi J, Song Y, Chen X. Paleolithic human exploitation of plant foods during the last glacial maximum in North China, Proceedings of the National Academy of Sciences, 2013, 110(14), 5380-5385. https://doi.org/10.1073/pnas.1217864110 Song Y, Shi J, Liu L. The origins of millet agriculture in north China from the evidences on the grinding stones at locality S9 of Shizitan Paleolithic site in Shanxi province.Agricultural history of China, 2013, 32(03): 3-8+101. Tian C. An analysis of plant residues at the Pigeon Mountain Site in Ningxia. University of Chinese Academy of Sciences ,2018. Wang Z, Ma Z, Yang X, Zhang C, Zhou G, Fan C, Ge Q. Starch residues from shell tools from sites of Xianrendong and Diaotonghuan and its implications for paleoclimate. Quaternary Sciences, 2012, 32(02): 256-262. 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(A) Location map of Henan Province in China; (B) Elevation map of the Pingdingshan area; (C) Aerial view of the Wenquan site\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/55b8dbdd2b4d021fdcdf89a8.png"},{"id":104808393,"identity":"81f7e303-d9b1-42ca-8665-9ba3221cc04e","added_by":"auto","created_at":"2026-03-17 12:37:04","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":621000,"visible":true,"origin":"","legend":"\u003cp\u003eResidue-sampled stone tools from the Wenquan Site (red circles indicate sampling locations)\u003c/p\u003e\n\u003cp\u003e(1.Handaxe(22wq：160);2.Cleaver(22wq：222);3. Pick(23wq：908);4. Large flake 2(23wq：1032);5. Large flake 1(23wq：563);6. Heavy-duty scraper 1(23wq：1138);7. Heavy-duty scraper 2(23wq：1341);8. Large flake 3(23wq：1344);9. Flake(23wq：1480);10. Scraper(23wq：965);11. Denticulate(23wq：879))\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/21038cbf9beaf5a3bf61ab3f.png"},{"id":104460675,"identity":"9fd93eda-b710-4424-a328-43764d024bab","added_by":"auto","created_at":"2026-03-12 03:58:20","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":791842,"visible":true,"origin":"","legend":"\u003cp\u003eStarch grain types recovered from residues at the Wenquuan site（a.Triticeae；b-c.Paniceae；d-e.Coix；f.Trichosanthes；g.Setaria；h.Agropyron；i.Dioscorea；j.Pueraria）\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/f93398131b6d8d8bbe9308b5.png"},{"id":104460678,"identity":"426a94f9-2080-49ac-ab21-967c2043a193","added_by":"auto","created_at":"2026-03-12 03:58:20","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":506383,"visible":true,"origin":"","legend":"\u003cp\u003eDamaged starch grains recovered from residues at the Wenquan site（a.Central indentation；b.Fragmentation；c.Rough and ragged periphery）\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/984da82bf5eb9419077b1f05.png"},{"id":104809138,"identity":"0778048d-058a-4bd2-b6c4-e77539d08ec6","added_by":"auto","created_at":"2026-03-17 12:47:47","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3977284,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/192af64e-3691-47ac-8fb8-c4530829ce78.pdf"},{"id":104780492,"identity":"8f7e32e9-a976-4cf2-bd2a-83e24e9c1886","added_by":"auto","created_at":"2026-03-17 07:53:09","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":15697,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial1.docx","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/c796732efcf05130453aaf6d.docx"},{"id":104780444,"identity":"691f15d9-e5ea-4cf9-b419-2a069e62ec4a","added_by":"auto","created_at":"2026-03-17 07:52:59","extension":"xlsx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":12507,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial2.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-9007914/v1/d9cbc5c78c44f98381a568af.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Residue Analysis of Stone Artifacts from the Wenquan Paleolithic Site Reveals Early Exploitation of Wild Cereals and Food Processing Functions ofAcheulean Tools in Northern China (70,000–40,000 BP)","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eDuring the Paleolithic, ancient populations sustained themselves primarily through gathering and hunting, with plants serving as one of their crucial food sources. Starch, a long-chain compound formed by the linkage of glucose molecules, is widely present in the form of starch granules within the cells of plant organs such as roots, stems, and seeds [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Due to the taxonomic significance of starch granules in botany, they are frequently used in archaeological research to identify ancient plant remains rich in starch [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. During the collection and processing of plants, starch granules readily adhere to the surfaces of tools such as stone artifacts, where they can be preserved for extended periods. Through the systematic extraction and identification of these residual starch grains on tool surfaces, archaeologists can infer the functions of the related tools, thereby revealing how early humans exploited plant resources.\u003c/p\u003e \u003cp\u003eAs one of the key methods in archaeological residue studies, starch grain analysis has been widely applied to prehistoric sites to explore ancient populations\u0026rsquo; utilization of plant food resources and the functions of stone tools. Regarding tool function, starch grains from cereals and other gathered plants were identified on stone tools from the Oldenburg LA 77 site in northern Germany(3270\u0026ndash;2920 BC), indicating that its inhabitants used stone tools to process a highly diverse of plant foods [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. Ma et al. analysed a large number of starch grains and phytoliths recovered from the surfaces of four stone knives unearthed at the Lajia site, demonstrating that one of their functions was crop harvesting [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. Regarding plant food resources, starch grains from Triticeae, Fabaceae, and Fagaceae were identified in four Neolithic dental calculus samples from Dehesilla Cave in Spain, providing new insights into dietary patterns at the site [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. In China, starch grains from Panicoideae, Triticeae, Job\u0026rsquo;s tears (\u003cem\u003eCoix\u003c/em\u003e spp.), and underground storage organs were discovered on 10 amphora sherds from the Liujiazhai site, providing evidence for investigating the subsistence economy of migrants in northwestern Sichuan[\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. Several pottery \u003cem\u003efu\u003c/em\u003e pots unearthed from the early strata of the Hemudu site contained black foodcrusts; residue analysis of one sample revealed starch grains from rice (\u003cem\u003eOryza\u003c/em\u003e sp.), Triticeae tribe, oak acorn (\u003cem\u003eQuercus\u003c/em\u003e sp.), Papilionoideae beans, and lily bulbs (\u003cem\u003eLilium\u003c/em\u003e sp.), reflecting the Hemudu people\u0026rsquo;s exploitation of diverse plant food resources [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eCurrently, while starch grain analysis has demonstrated significant potential in studying and discussing the plant food resources of prehistoric populations and the functions of artifacts, its application in Paleolithic sites remains relatively limited. In southern Italy and Sicily, starch grain analysis of dental calculus from unearthed human remains revealed starch grains from the Triticeae tribe, Paniceae tribe, Andropogoneae tribe, and Sambucus, providing evidence for the study of plant foods in the Late Paleolithic of this region [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. Starch grains from various plants, including Arundo donax (cane), \u003cem\u003eTypha\u003c/em\u003e sp., and species in the Triticeae tribe, were extracted from ground stones of the Paleolithic period in the Pontic steppe, revealing the processing behaviors of the past populations toward starchy plants[\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. In China, starch grain analysis was conducted on some specimens from the Zhanghuokou site. Combined with experimental results and specimen morphology, the analysis indicated that Luonan handaxes might have been used for digging underground roots and tubers [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]. Starch grains extracted from three grinding tools at the Shizitan site in the middle reaches of the Yellow River (23,000\u0026ndash;19,500 BP) revealed that ancient humans intensified the use of Paniceae and tuber plants more than ten thousand years earlier than the Neolithic period [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. The Wenquan Site is a large-scale, open-air Paleolithic site where a significant quantity of stone artifacts and animal fossils have been discovered. The stone artifacts include cores, flakes, hammerstones, choppers, scrapers, points, handaxes, picks, cleavers, spheres, and other types. The overall lithic assemblage exhibits characteristics of both a simple core-and-flake lithic industry and tool types commonly associated with the Acheulean tradition, such as handaxes and cleavers, dating to the Late Pleistocene. This study aims to conduct starch grain analysis by extracting residues from stone tools at the Wenquan site to explore how ancient humans utilized these tools 70,000 to 40,000 years ago, as well as the types of plant food resources they processed and utilized with them.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\n \u003ch2\u003e2.1 Sites and Stone Tools\u003c/h2\u003e\n \u003cp\u003eThe Wenquan Site is located in Wenquan Town, Ruzhou City, Pingdingshan City, Henan Province, China, at geographic coordinates 34\u0026deg;12\u0026prime;16.9\u0026Prime;N, 112\u0026deg;37\u0026prime;57.3\u0026Prime;E, and an altitude of 236 meters (Fig. \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e). Since 2020, the Henan Provincial Institute of Cultural Heritage and Archaeology has conducted three archaeological excavations at the site, yielding a substantial assemblage of stone artifacts. These include handaxes, cleavers, picks, large knives, heavy-duty scrapers, and choppers, which date to between 70,000 and 40,000 years ago [\u003cspan class=\"CitationRef\"\u003e12\u003c/span\u003e].\u003c/p\u003e\n \u003cp\u003eThe research materials consist of 11 stone artifacts recovered from the Wenquan Site during the 2022 and 2023 excavations. These include flake tools, Acheulean implements, scrapers, and a denticulate(Fig. \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e). The flake assemblage comprises four specimens: Flake (23wq:1480), Large Flake 1 (23wq:563), Large Flake 2 (23wq:1032), and Large Flake 3 (23wq:1344). The Acheulean tools are represented by three pieces: a Handaxe (22wq:160), a Pick (23wq:908), and a Cleaver (22wq:222). The scraper category includes Heavy-duty Scraper 1 (23wq:1138), Heavy-duty Scraper 2 (23wq:1341), and Scraper (23wq:965). Finally, the denticulate is represented by a single specimen: Denticulate (23wq:879). Detailed classification and descriptions of these stone artifacts are provided in Supplementary Material 1.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\n \u003ch2\u003e2.2 Sample collection\u003c/h2\u003e\n \u003cp\u003eThis study followed the residue extraction and laboratory processing methods established in published studies [\u003cspan class=\"CitationRef\"\u003e13\u003c/span\u003e\u0026ndash;\u003cspan class=\"CitationRef\"\u003e15\u003c/span\u003e]. In the processes of sample collection and pre-treatment, we consistently maintained a contamination-free environment for both the stone artifacts and the laboratory to prevent sample contamination. The main measures we took included:\u003c/p\u003e\n \u003cp\u003eSample preservation phase: From excavation to storage in the repository, the stone artifact samples were kept in contamination-isolated sample bags at all times.\u003c/p\u003e\n \u003cp\u003eLaboratory sampling phase: During the entire residue extraction process from stone tool surfaces, a disposable toothbrush head was used for each individual sample at every step, and powder-free gloves were worn to prevent cross-contamination.\u003c/p\u003e\n \u003cp\u003eExperimental processing phase: To prevent contamination during residue handling, laboratory personnel wore laboratory coats, masks, and powder-free gloves throughout the entire process.\u003c/p\u003e\n \u003cp\u003eContamination control phase: To account for potential airborne starch grain contamination in the sampling environment which could interfere with experimental results, Purified water was added to 10 cm Petri dishes, which were then exposed to ambient air for three days, serving as contamination controls.\u003c/p\u003e\n \u003cp\u003eSample Collection: A zoned approach was adopted, with sampling limited to one-half of the working edge surface area of each stone tool.\u003c/p\u003e\n \u003cp\u003eThe residue sampling procedure consisted of three steps:1. Collecting soil samples adhering to the tool surface by scraping with clean implements. 2.Cleaning the tool surfaces with purified water and a toothbrush to remove potential contaminants. 3.Repeatedly agitating and scrubbing the working edge surfaces with purified water using an ultrasonic toothbrush to collect ultrasonic samples. Some stone artifacts lacked adhering soil samples, so only ultrasonic samples were extracted from them. In total, 11 stone artifacts were selected for surface residue sampling in this study, yielding 19 residue samples (Table \u003cspan class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\n \u003cp\u003eLaboratory Processing of Samples: Pretreatment and slide preparation of the samples were conducted at the Henan Provincial International Joint Laboratory of Modern Human Origins. Three of these samples were processed at the Laboratory of Archaeometry at Xiamen University. The laboratory pretreatment and slide preparation procedures followed established protocols [\u003cspan class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e17\u003c/span\u003e] and were as follows:\u003c/p\u003e\u003cspan\u003e\n \u003cp\u003e1) Add deionized water to the tube, balance the centrifuge tubes, and centrifuge at 2000 rpm for 5 min. Discard the supernatant to concentrate the sample.\u003c/p\u003e\n \u003c/span\u003e \u003cspan\u003e\n \u003cp\u003e2) Add a 5% (NaPO\u003csub\u003e3\u003c/sub\u003e)\u003csub\u003e6\u003c/sub\u003e solution and place the tube on a shaker for 2 h to disperse sample particles. Then centrifuge at 5000 rpm for 5 min and discard the supernatant.\u003c/p\u003e\n \u003c/span\u003e \u003cspan\u003e\n \u003cp\u003e3) Add a heavy-liquid solution of sodium polytungstate (density 2.4 g/cm\u003csup\u003e3\u003c/sup\u003e). Agitate using a vortex mixer for 10 seconds, then centrifuge at 2000 rpm for 5 min. Using a micropipette, carefully extract the floating fraction (containing lighter plant remains such as starch grains) and transfer it to a new centrifuge tube.\u003c/p\u003e\n \u003c/span\u003e \u003cspan\u003e\n \u003cp\u003e4) Add deionized water, centrifuge at 5000 rpm for 5 min, and discard the supernatant. The resulting sediment contains the plant remains fraction. Repeat this step three times to thoroughly remove residual sodium polytungstate heavy liquid.\u003c/p\u003e\n \u003c/span\u003e \u003cspan\u003e\n \u003cp\u003e5) Extract a portion of the suspension containing plant remains and place a drop in the center of a clean glass microscope slide. Add a drop of 50% glycerol solution, cover with a coverslip, and seal the edges with nail polish. Allow the slide to dry before microscopic examination.\u003c/p\u003e\n \u003c/span\u003e\u0026nbsp;\u003ctable id=\"Tab1\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eInformation for Stone Tool Samples studied from the Wenquan Site\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSpecimen ID\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eLab ID\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eArtifact Type\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSample Description\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSampling Location\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1138\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHeavy-duty scraper\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1138\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHeavy-duty scraper\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1344\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1344\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1032\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1032\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1341\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHeavy-duty scraper\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1341\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHeavy-duty scraper\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:563\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:563\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eLarge flake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e22wq:160\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHandaxe-like\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e22wq:160\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eHandaxe-like\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:908\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePick\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:908\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePick\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e22wq:222\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eCleaver\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eSurface Soil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e22wq:222\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eCleaver\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:965\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eScraper\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:879\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003edenticulate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23wq:1480\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eFlake\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eUltrasonicated\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eBlade edge\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e\n \u003ch2\u003e2.3 Sample Microscopic Examination\u003c/h2\u003e\n \u003cp\u003eA total of 107 slides were prepared from the 21 samples in this study. Microscopic examination was conducted using a Sunny Optical biological microscope (model EX31) at the Henan Provincial International Joint Laboratory on Origins of Modern Humans in East Asia and a Zeiss Scope A1 biological microscope at the Laboratory of Archaeometry at the Xiamen University. Photomicrography and data measurement were performed using imaging systems and measurement software integrated with the respective microscopes. Identification of residues, particularly starch grains, followed published research on Chinese starch grain morphology[\u003cspan class=\"CitationRef\"\u003e18\u003c/span\u003e\u0026ndash;\u003cspan class=\"CitationRef\"\u003e20\u003c/span\u003e]. Starch grains were identified mainly according to their morphological characteristics, including size, shape, position of the hilum, presence of fissures and configuration, and extinction cross features under polarized light[\u003cspan class=\"CitationRef\"\u003e21\u003c/span\u003e].\u003c/p\u003e\n\u003c/div\u003e"},{"header":"3. Results and analysis","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e\n \u003ch2\u003e3.1 Contamination assessment\u003c/h2\u003e\n \u003cp\u003eA total of 205 starch grains were identified across the 19 samples obtained from 11 stone tools from the Wenquan site. Among these, 32 grains were found in the surface soil samples, and 173 grains were found in the ultrasonic samples. Furthermore, no starch grains were detected in the control samples, indicating that the sampling and slide preparation processes were free from contamination, and thus the starch grains discovered in this study are of ancient origin. The small quantity of starch grains found in the surface soil samples reflects their presence in the burial environment. However, their number is substantially lower than that found in the ultrasonic samples, demonstrating that the starch grains in the ultrasonic samples originate from residues preserved on the artifacts themselves, rather than from post-depositional contamination in the burial context. Consequently, the starch grains identified in the ultrasonic samples serve as the primary subjects for analysis in this study, while those from the surface soil samples serve as a comparativce reference. The detailed classification and comparison of starch grains from the surface soil samples and ultrasonic samples can be found in Supplementary material 2.\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\n \u003ch2\u003e3.2 Identification and statistical analysis\u003c/h2\u003e\n \u003cp\u003eAmong the starch grains discovered in this study, 159 grains were identifiable, accounting for 77.6% of the total starch grains found. Based on morphological characteristics, these were classified into eight types (Fig. \u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e). The remaining 46 grains could not be identified due to morphological damage or lack of diagnostic features (Table \u003cspan class=\"InternalRef\"\u003e2\u003c/span\u003e,Table\u0026nbsp;\u003cspan class=\"InternalRef\"\u003e3\u003c/span\u003e). To enable accurate quantitative analysis, this study employed three quantitative indicators: the absolute count of starch grains, percentage, and occurrence frequency. The absolute count refers to the total number of starch grains counted for each taxon. The percentage is the proportion of the absolute count of a specific starch grain type relative to the total absolute count of all starch grains. The occurrence frequency was calculated using the formula: (Number of artifacts bearing a specific starch grain type / Total number of artifacts) \u0026times; 100%.\u003c/p\u003e\n \u003cdiv class=\"gridtable\"\u003e\u0026nbsp;\u003ctable id=\"Tab2\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eStatistical Analysis of Starch Grains from Ultrasonicated Residues at the Wenquan Site\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003ccolgroup cols=\"12\"\u003e\u003c/colgroup\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCategory\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTriticeae\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePaniceae\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCoix\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTrichosanthes\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSetaria\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAgropyron\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eDioscorea\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePueraria\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eUnidentified\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eDamaged Grains\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eStarch Morphotype\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅠ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅡ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅢ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅣ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅤ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅥ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅦ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅧ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e42\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e173\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePercentage\u003c/p\u003e\n \u003cp\u003e(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.23\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e16.76\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5.78\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e11.56\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e6.36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.73\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1.16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e24.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e20.81\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eFrequency\u003c/p\u003e\n \u003cp\u003e(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e72\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e45\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e36\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e27\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e45\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003c/div\u003e\n \u003cdiv class=\"gridtable\"\u003e\n \u003cdiv align=\"left\" class=\"colspec\"\u003e\u003cbr\u003e\u003c/div\u003e\n \u003cdiv align=\"left\" class=\"colspec\"\u003e\u003cbr\u003e\u003c/div\u003e\u0026nbsp;\u003ctable id=\"Tab3\" border=\"1\"\u003e\n \u003ccaption language=\"En\"\u003e\n \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\n \u003cdiv class=\"CaptionContent\"\u003e\n \u003cp\u003eStatistical Analysis of Starch Grains from Surface Soil Residues at the Wenquan Site\u003c/p\u003e\n \u003c/div\u003e\n \u003c/caption\u003e\n \u003ccolgroup cols=\"12\"\u003e\u003c/colgroup\u003e\n \u003cthead\u003e\n \u003ctr\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCategory\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTriticeae\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePaniceae\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eCoix\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTrichosanthes\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eSetaria\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eAgropyron\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eDioscorea\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003ePueraria\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eUnidentified\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/th\u003e\n \u003cth align=\"left\"\u003e\n \u003cp\u003eDamaged Grains\u003c/p\u003e\n \u003c/th\u003e\n \u003c/tr\u003e\n \u003c/thead\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eStarch Morphotype\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅠ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅡ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅢ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅣ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅤ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅥ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅦ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eⅧ\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eS15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003ePercentage\u003c/p\u003e\n \u003cp\u003e(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e15.63\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e3.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e9.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e28.13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003eFrequency(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e37\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e-\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd align=\"left\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n \u003c/div\u003e\n \u003cp\u003eType I starch grains totaled 38, with 35 recovered from ultrasonic samples and 3 from surface soil samples. Type I starch grains are generally large, with a size range of 13.79\u0026ndash;29.61 \u0026micro;m and a mean size of 21.52 \u0026micro;m. The shapes are subcircular or oval, with the hilum located centrally, and faint lamellae are visible. The extinction cross arms appear as \u0026quot;+\u0026quot; or \u0026quot;X\u0026quot; shapes. Based on these morphological characteristics and comparisons with modern reference collections, Type Ⅰgrains are identified as originating from the tribe Triticeae.\u003c/p\u003e\n \u003cp\u003eType II starch grains totaled 33, with 29 from ultrasonic samples and 4 from surface soil samples. Type II starch grains are relatively small, with a size range of 9.78\u0026ndash;20.63 \u0026micro;m and a mean size of 14.50 \u0026micro;m. The shapes are subcircular or polygonal, with a visible central hilum; some grains exhibit \u0026quot;V\u0026quot; or \u0026quot;Y\u0026quot; shaped fissures. The extinction cross arms are perpendicular, forming a \u0026quot;+\u0026quot; shape. These morphological features resemble starch grains from seeds of the tribe Paniceae [\u003cspan class=\"CitationRef\"\u003e22\u003c/span\u003e]. Additionally, these starch grains could potentially originate from the stems or leaves of the subfamily Panicoideae plants [\u003cspan class=\"CitationRef\"\u003e23\u003c/span\u003e].\u003c/p\u003e\n \u003cp\u003eType III starch grains totaled 24, with 21 from ultrasonic samples and 3 from surface soil samples. Type III starch grains are predominantly single grains, with a size range of 11.30\u0026ndash;27.12 \u0026micro;m and a mean size of 18.33 \u0026micro;m. Shapes are subcircular or oval, sometimes with rounded rectangular forms. The hilum is central, and fissures are \u0026quot;V\u0026quot;, \u0026quot;Y\u0026quot;, or \u0026quot;\u0026mdash;\u0026quot; shaped. Under polarized light, they exhibit strong extinction with mutually perpendicular extinction arms that display a Z-shaped bend near the edges. Their morphological characteristics resemble starch grains from Job\u0026apos;s tears [\u003cspan class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan class=\"CitationRef\"\u003e25\u003c/span\u003e], hence they are identified as starch grains from the \u003cem\u003eCoix\u003c/em\u003e.\u003c/p\u003e\n \u003cp\u003eType IV starch grains totaled 15, with 10 from ultrasonic samples and 5 from surface soil samples. These grains have a size range of 9.64\u0026ndash;24.77 \u0026micro;m (mean: 14.81 \u0026micro;m) and exhibit diverse shapes, including subcircular, oval, bell-shaped, and polygonal forms. The hilum is eccentric, lamellae are mostly invisible, and the extinction cross arms are bent or exhibit a certain curvature [\u003cspan class=\"CitationRef\"\u003e11\u003c/span\u003e]. Based on morphological characteristics, Type IV grains are identified as originating from \u003cem\u003eTrichosanthes\u003c/em\u003e (snake gourd).\u003c/p\u003e\n \u003cp\u003eType V starch grains totaled 24, with 20 from ultrasonic samples and 4 from surface soil samples. Type V starch grains are small, with a size range of 6.02\u0026ndash;10.99 \u0026micro;m and a mean size of 8.25 \u0026micro;m. Their morphology is similar to Type II but smaller in size, being subcircular to oval in shape with a mostly visible, central hilum. The extinction cross arms are perpendicular, appearing as a \u0026quot;+\u0026quot; shape. Based on morphological characteristics, this type is identified as starch grains from \u003cem\u003eSetaria\u003c/em\u003e.\u003c/p\u003e\n \u003cp\u003eType VI starch grains totaled 12, with 11 from ultrasonic samples and 1 from surface soil samples. These grains are relatively large, with a size range of 14.39\u0026ndash;34.45 \u0026micro;m and a mean size of 26.02 \u0026micro;m. They are predominantly single grains, subcircular in shape, with a central hilum; some grains visible growth rings. The extinction cross is diffuse and appears as an \u0026quot;X\u0026quot; shape [\u003cspan class=\"CitationRef\"\u003e2\u003c/span\u003e]. Based on these morphological characteristics, Type VI grains are identified as originating from \u003cem\u003eAgropyron\u003c/em\u003e.\u003c/p\u003e\n \u003cp\u003eType VII starch grains totaled 6, with 3 from ultrasonic samples and 3 from surface soil samples. Type VII starch grains are large, with a size range of 18.80\u0026ndash;49.99 \u0026micro;m and a mean size of 28.34 \u0026micro;m. Shapes are mostly oval or rounded triangular, with a few irregular forms; the hilum is eccentric, often located near one end. Under polarized light, extinction is strong, and the extinction cross arms are non-perpendicular and bent. Based on these characteristics, Type VII is identified as starch grains from \u003cem\u003eDioscorea\u003c/em\u003e.\u003c/p\u003e\n \u003cp\u003eType VIII starch grains totaled 2, both from ultrasonic samples. Type VIII starch grains have a size range of 21.21\u0026ndash;22.69 \u0026micro;m and a mean size of 21.96 \u0026micro;m. The morphology is polygonal, with a visible central hilum; one grain exhibits a \u0026quot;\u0026mdash;\u0026quot; shaped fissure at the hilum, and the extinction cross arms are bent. The morphological characteristics of Type VIII starch grains resemble those from tubers of the genus \u003cem\u003ePueraria\u003c/em\u003e and support their identification as such [\u003cspan class=\"CitationRef\"\u003e26\u003c/span\u003e].\u003c/p\u003e\n \u003cp\u003eA total of 40 damaged starch grains were observed across the residue samples, with 36 recovered from ultrasonic samples and 4 from surface soil samples. The damaged starch grains in the ultrasonic samples account for 20.81% of the total ultrasonic sample grains, with a frequency of occurrence of 45%. These damaged starch grains exhibit features such as breakage, rough edges, and central depressions, which are consistent with mechanical damage caused by grinding [\u003cspan class=\"CitationRef\"\u003e27\u003c/span\u003e]. This pattern suggests that grinding processing was likely applied to the processed materials(Fig. \u003cspan class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\n\u003c/div\u003e"},{"header":"4. Discussions","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e4.1 Utilization of plant resources by past population at the Wenquan site\u003c/h2\u003e \u003cp\u003eA total of 173 starch grains, belonging to eight taxonomic groups, were identified across the 19 samples obtained from the 11 stone artifacts examined at the Wenquan Site. These groups include Triticeae, Paniceae, Coix, Trichosanthes, Setaria, Agropyron, Dioscorea, and Pueraria. Among these, starch grains from grasses were the most abundant, including those from the Panicoideae (Paniceae, Coix, Setaria) (70 grains) and the Triticeae (Triticeae, Agropyron (within Triticeae)) (46 grains), totaling 116 grains. This accounts for approximately 67% of the identifiable starch grains, indicating that grasses constituted a significant proportion of the diet of the ancient inhabitants at the Wenquan site. Furthermore, 15 starch grains were identified from rhizomatous plants (Trichosanthes, Dioscorea, and Pueraria), accounting for about 8.7% of the identifiable starch grains.\u003c/p\u003e \u003cp\u003eCurrent research indicates that grasses and rhizomatous plants have been widely discovered at Paleolithic sites across China, highlighting their importance as key food resources for ancient human populations during the Chinese Paleolithic period.\u003c/p\u003e \u003cp\u003eIn northern China, the earliest evidence for the collection and consumption of wild grasses by prehistoric humans dates back to the Shizitan Site Complex in Jixian County, Shanxi Province. Residue analysis of stone tools from Shizitan Locality 29 (ca. 28,000\u0026ndash;13,000 cal. BP), Locality 14 (ca. 23,000\u0026ndash;19,500 cal. BP), and Locality 9 (ca. 13,800\u0026ndash;8,500 cal. BP) revealed starch grains from wild grass seeds, primarily from tribes Triticeae and Paniceae (subfamilies Pooideae and Panicoideae) [\u003cspan additionalcitationids=\"CR29\" citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. Starch grains from Triticeae and Panicoideae grasses have also been found on stone artifacts excavated from the Gezishan site in Qingtongxia, Ningxia. This indicates that around 12,000\u0026ndash;8,000 years ago, the ancient inhabitants of Gezishan had already begun exploiting wild grass seeds, with grasses dominating their plant resource utilization [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eIn southern China, large numbers of Triticeae and Paniceae starch grains were found on shell tools unearthed from the Xianren Cave and Diaotonghuan sites in Wannian County, Jiangxi Province. These findings demonstrate that as early as approximately 20,000 years ago, the inhabitants of the these sites were gathering and consuming grass plants, including those from the Triticeae and Paniceae[\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. The discovery of Triticeae starch grains in dental calculus from hominins at the Fuyan Cave in Daoxian, Hunan, extends the earliest evidence for the consumption of wild grass seeds to approximately 80,000 years ago[\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eTrichosanthes, Dioscorea, and Pueraria all produce edible roots and tubers. Their underground storage organs are rich in starch, providing a high caloric return with relatively low foraging effort, making them prime targets for prehistoric collection and utilization. At the Gantangqing site in Jiangchuan, Yunnan, underground corms and rhizomes of aquatic plants were discovered. Combined with the presence of wooden tools exhibiting clear use-wear traces at the site (approximately 361,000\u0026ndash;250,000 years ago), researchers propose that the hominins at Gantangqing used fabricated tools of selected wood to exploit underground tubers, rhizomes, or corms[\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. On handaxe unearthed from the Zhanghuokou site in Luonan, Shaanxi (approximately 250,000\u0026ndash;100,000 years ago), definitive starch grains of rhizomatous plants have been discovered. This finding also provides evidence for investigating the utilization of rhizomatous plants by early hominin [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eStarch grain analysis of stone artifacts from Locality 2 of the Shuidonggou site in Lingwu, Ningxia, indicates that during the MIS 3, wild grass seeds, underground storage organs, and wild Fabaceae were all incorporated into the subsistence strategies of the hominins at the site[\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Similarly, starch grains of rhizomatous plants have been found on stone artifacts unearthed from the Shizitan site in Jixian County, Shanxi (ca. 28,000\u0026ndash;8,500 cal. BP) [\u003cspan additionalcitationids=\"CR29\" citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e]. Starch grain analysis of soil samples from the Paleolithic strata (approximately 17,000\u0026ndash;13,000 years ago) at the Qihe Cave site in Zhangping, Fujian, has identified starch grains from plants of \u003cem\u003ePueraria\u003c/em\u003e, suggesting that Pueraria served as a major source of starchy food for the inhabitants of the site during the late Pleistocene [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eOur study demonstrates that in northern China, Triticeae plants had a long history of exploitation and utilization prior to their domestication. As early as 70,000\u0026ndash;40,000 years ago, ancient hominins at the Wenquan site had already begun collecting and consuming wild grass seeds and rhizomatous plants. The gathering and consumption of grasses constituted a significant component of the subsistence economy of the inhabitants of the Wenquan site.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003e4.2 Functional study of stone tools at the Wenquan site\u003c/h2\u003e \u003cp\u003eThe lithic artifacts from the Wenquan site are predominantly made of quartzite and andesite, with tool production achieved primarily through hard-hammer percussion. Typological analysis reveals the presence of small scrapers and simple core-chopper tools at the Wenquan site, alongside large cutting tools such as handaxes, cleavers, picks, and large knives, representing characteristic of Acheulean-type lithics[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e].In this study, 11 stone tools were selected for the extraction and analysis of surface residues, including flakes (both regular and large flakes), Acheulean large tools (handaxes, picks, and cleavers), scrapers (both regular and heavy-duty scrapers), and denticulates. Based on the quantity and identification results of the starch grains, we conducted a preliminary discussion on how the early inhabitants of the Wenquan site used these stone tools.\u003c/p\u003e \u003cp\u003eBased on the absolute counts, morphological types, and presence of damaged starch grains found on the sampled stone tools, all 11 tools were likely used to collect or process at least one or multiple plant types (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor the flakes, the assemblage includes one regular flake and three giant flakes. On flake (23wq:1480) and giant flake 2 (23wq:1032), starch grains from grass plants were identified. Giant flake 1 (23wq:563) yielded starch grains from rhizome plants. Giant flake 3 (23wq:1344) was found contain starch grains from both grass and rhizome plants. Notably, this stone tool yielded the highest quantity and most diverse types of starch grains, with 72 grains identified from the ultrasonicated sample, belonging to six distinct types. These results suggest that the Wenquan inhabitants did not perform secondary modification to the flakes after initial production. Instead, they used them directly for harvesting grass plants and excavating rhizomatous plants.\u003c/p\u003e \u003cp\u003eRegarding the Acheulean large tools, these include handaxes, picks, and cleavers. Starch grains from both grass plants and rhizomatous plants were identified on handaxe 22wq:160 and cleaver 22wq:222. On pick 23wq:908, only starch grains from grass plants were found. The results of the starch grain analysis indicate that the ancient humans of the Wenquan Site likely used these Acheulean large tools (handaxes, picks, and cleavers) to process and handle both grass plants and rhizomatous plants. Guan Ying et al. [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e] previously reported starch grains from rhizomatous plants on handaxes from the Zhanghuokou site in Luonan, Shaanxi Province, and proposed that such tools may have been used for digging the underground storage organs of plants. Our findings align with and extend this interpretation: large tools from the Acheulean cultural tradition at Wenquan could have served for cutting and harvesting starch-rich grasses and rhizomes.\u003c/p\u003e \u003cp\u003eThese results further demonstrate the multifunctional nature of large Paleolithic tools such as handaxes and picks. They were not limited to woodworking but were also employed for exploiting and processing starch-containing grasses and rhizomes.\u003c/p\u003e \u003cp\u003eRegarding the scrapers, the assemblage includes both regular scrapers and heavy-duty scrapers. A relatively high quantity of starch grains was recovered from two scrapers: regular scraper 23wq:965 and heavy-duty scraper 23wq:1341. These included grains from Triticeae, Paniceae, \u003cem\u003eCoix\u003c/em\u003e, and others taxa.\u003c/p\u003e \u003cp\u003eOverall, the highest quantities of starch grains among all sampled tools were found on the scrapers. On Heavy-duty Scraper 1 (23wq:1138), starch grains from grass plants were identified. In contrast, Heavy-duty Scraper 2 (23wq:1341) and regular scraper (23wq:965) yielded starch from both grass plants and rhizomatous plants, with comparatively high counts on these two artifacts. The results suggest that scrapers at the Wenquan Site were likely used for collecting and processing wild grass seeds and rhizomatous plants.\u003c/p\u003e \u003cp\u003eRegarding the denticulate, only one denticulate(23wq:879) was sampled. The starch grains recovered from it resemble those from Triticeae, Trichosanthes, and \u003cem\u003eSetaria\u003c/em\u003e plants. This suggests that the denticulate may have been used for handling and processing both grass plants and rhizomatous plants.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eStatistical Analysis of Starch Grains from Stone Tool Residues at the Wenquan Site\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eTool Type\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eArtifact ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSample ID\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSample Description\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eIdentified Starch Taxa\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eFlakes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFlake\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:1480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Agropyron\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLarge flake 1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:563\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTrichosanthes\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLarge flake 2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:1032\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Setaria\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLarge flake 3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:1344\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Trichosanthes、Setaria、Agropyron、Pueraria\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eAcheulean large tools\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ehandaxe\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e22wq:160\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eCoix、Trichosanthes、Setaria、Agropyron\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePick\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:908\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePaniceae\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCleaver\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e22wq:222\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Dioscorea\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eScrapers\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHeavy-duty scraper 1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:1138\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eCoix\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHeavy-duty scraper 2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:1341\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Trichosanthes、Setaria、Dioscorea\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eScraper\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:965\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Paniceae、Coix、Setaria、Agropyron、Dioscorea\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDenticulate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDenticulate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e23wq:879\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eS18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eUltrasonicated\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eTriticeae、Trichosanthes、Setaria\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"5. Conclusions","content":"\u003cp\u003eThis study conducted residue extraction and analysis on 11 stone tool from the Wenquan site in Ruzhou, Henan Province. Starch grains assignable to eight plant types were identified, including those from the tribes Triticeae and Paniceae, as well as genera such as \u003cem\u003eCoix\u003c/em\u003e, \u003cem\u003eTrichosanthes\u003c/em\u003e, Seteria, \u003cem\u003eAgropyron\u003c/em\u003e, \u003cem\u003eDioscorea\u003c/em\u003e, and \u003cem\u003ePueraria\u003c/em\u003e. These findings indicate that wild grass seeds and root/tuber plants (rhizomatous plants) were important plant food resources for the local Paleolithic inhabitants.\u003c/p\u003e \u003cp\u003eThe inhabitants of the Wenquan Site used stone tools to collect and process wild grass seeds and rhizomatous plants available in their environment. This indicates that, early in the Paleolithic period, humans in the Central Plains region consciously incorporated wild grains into their diet. Furthermore, the diverse starch grain types recovered from the surfaces of various stone tools, including scrapers, handaxes and picks provide new evidence and insights into the functional roles of these stone tools at the Wenquan site. In particular, the presence of grass starch grains on Acheulean large tools, such as handaxes and picks, provides fresh perspectives on the use patterns and versatility of such tools in early Paleolithic subsistence strategies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAcknowledgements\u003c/p\u003e\n\u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;This study was supported by the Henan Cultural Relics Protection Research Project (Grant No. 25HNWWJ—KJ01), the Henan Provincial Department of Finance Basic Scientific Research Operating Funds Project (2025HKYJKY01), the Archaeological Talent Promotion Program of China (Grant No. 2025-191), the Chongqing Municipal Education Commission Science and Technology Youth Project (KJQN202500536), the Henan Social Science Project “Research on Late Pleistocene Lithic Technology in the Central Plains Based on the Ruzhou Wenquan Site”, and the Henan Provincial Institute of Cultural Heritage and Archaeology Special Conservation Project “Archaeological Survey and Exploration of the Ruzhou Wenquan Paleolithic Site”.\u003c/p\u003e\n\u003cp\u003eAuthor contributions\u003c/p\u003e\n\u003cp\u003eYuqian Yan, Wentai Lou, Jingyi Li, and Wei Ge conducted the experiments and data analysis. Yuqian Yan wrote the manuscript. Wentai Lou, Qingpo Zhao, Huanhuan Ma, and Wei Ge revised the paper. Shenglan Peng, Zizhen Zhao, Jia Huang, and Jun Chen participated in the research and were responsible for the organization of lithic materials and the compilation of supplementary data.\u0026nbsp;All authors reviewed and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003eCompeting interests\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\n\u003cp\u003eData availability\u003c/p\u003e\n\u003cp\u003eThe authors confirm that the data supporting the findings of the study are available within the article and supplementary materials.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eYang X,L\u0026uuml; H,Xia Z.The application of plant starch grain analysis in archaeology.Archaeology and Cultural Relics,2006,(03): 87-91.\u003c/li\u003e\n\u003cli\u003eGe W.The application of starch grain analysis in archaeology. 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Frontiers in Plant Science, 2022,13, 989308. https://doi.org/10.3389/fpls.2022.989308 \u003c/li\u003e\n\u003cli\u003eLiu J-H, Ruan Q-J, Ge J-Y, Huang Y-J, Zhang X-L, Liu J, Li S-F, Shen H, Wang Y, Stidham T A, Deng C-L, Li S-H, Han F, Jin Y-S, O\u0026rsquo;Gorman K, Li B, Dennell R, Gao X. 300,000-year-old wooden tools from Gantangqing, southwest China, Science, 2025,389(6755), 78-83. https://doi.org/10.1126/science.adr8540\u003c/li\u003e\n\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":"Wenquan Paleolithic Site, Starch grain analysis, Plant utilization, Stone tool function","lastPublishedDoi":"10.21203/rs.3.rs-9007914/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9007914/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe Wenquan Site is located in Wenquan Town, Ruzhou City, Pingdingshan City, Henan Province, China. It is a Paleolithic site dating from approximately 70,000 to 40,000 years ago, and the unearthed stone tools exhibit distinct Acheulean technical features, including handaxes, cleavers, and scrapers. This study conducted residue analysis on 11 typical artifacts from the Wenquan Site to investigate plant food resource utilization by its ancient inhabitants and the functions of stone tools. Results indicate that starch grains were found on all 11 stone tools (including flakes, large flakes, handaxes, picks, cleavers, scrapers, heavy-duty scrapers, and denticulate tools). These starch grains show a high degree of similarity with those from grasses and rhizomatous plants, such as species from the Triticeae, Paniceae, \u003cem\u003eCoix\u003c/em\u003e, \u003cem\u003eTrichosanthes\u003c/em\u003e, \u003cem\u003eSetaria\u003c/em\u003e, \u003cem\u003eAgropyron\u003c/em\u003e, \u003cem\u003eDioscorea\u003c/em\u003e, and \u003cem\u003ePueraria\u003c/em\u003e tribes or genera. The Triticeae starch grains discovered on stone tools from the Wenquan site represent the earliest recorded evidence of Triticeae starch grains in northern China. This indicates that ancient humans at the Wenquan site had already begun to collect and consume grass seeds, providing important physical evidence for the study of the exploitation and utilization of wild cereals prior to their domestication. Moreover, a higher concentration of starch grains from Poaceae and rhizomatous plants was discovered on handaxes, picks, and cleavers, clearly indicating that ancient humans at the Wenquan site between 70,000 and 40,000 years ago used these large-sized tools to process food resources. This finding provides new directions and insights for discussing the functions of large Acheulean tools.\u003c/p\u003e","manuscriptTitle":"Residue Analysis of Stone Artifacts from the Wenquan Paleolithic Site Reveals Early Exploitation of Wild Cereals and Food Processing Functions ofAcheulean Tools in Northern China (70,000–40,000 BP)","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-12 03:58:15","doi":"10.21203/rs.3.rs-9007914/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-13T13:02:31+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-13T12:46:35+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-05T08:43:39+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-30T12:22:26+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"147720022617491454952028055290761542454","date":"2026-03-12T20:10:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"111132954333779213578537729952436895664","date":"2026-03-12T12:28:41+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"285779525524456566510969487153323942871","date":"2026-03-10T11:24:36+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-09T06:46:06+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-03-09T01:10:44+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-03-09T01:09:54+00:00","index":"","fulltext":""},{"type":"submitted","content":"npj Heritage Science","date":"2026-03-02T08:48:20+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":"f51beb12-7aa1-4af7-a5bf-82fa37fcef6a","owner":[],"postedDate":"March 12th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"in-revision","subjectAreas":[],"tags":[],"updatedAt":"2026-04-13T13:10:02+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-12 03:58:15","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9007914","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9007914","identity":"rs-9007914","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}