Identification of Rubus Seeds in Chinese Archaeological Sites: Based on the Morphological Atlas of 126 Modern Rubus Seeds

Identification of Rubus Seeds in Chinese Archaeological Sites: Based on the Morphological Atlas of 126 Modern Rubus Seeds | 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 Identification of Rubus Seeds in Chinese Archaeological Sites: Based on the Morphological Atlas of 126 Modern Rubus Seeds Aoyu Ren, Xiaojiao Ma, Xinmin Xu, Minzhen Yin, Hetian Jin This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8367387/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 12 You are reading this latest preprint version Abstract Rubus seeds constitute the predominant plant remains in archaeobotanical assemblages from Chinese archaeological sites. Nevertheless, no studies have yet reported the precise species-level identification of archaeological Rubus remains, as a result of the lack of systematic research on Rubus seed morphology. The first seed morphology atlas was established by systematically collecting and analyzing the morphological characteristics of 126 Rubus plants' mature seeds, which represent Rubus taxa in China (including 22 varieties). Utilizing this atlas, we identified Rubus seeds from seven archaeological sites, which ranged from the Shang-Zhou dynasties to the present day. Of these seeds, one was carbonized and six were uncarbonized. These specimens were identified as Rubus lambertianus , R. parvifolius , R. flosculosus , R. coreanus var. tomentosus , R. trianthus , and R. adenophorus . Furthermore, the atlas was employed to precisely identify Rubus seeds from nine archaeological sites that had been previously reported, primarily assigning them to R. parvifolius , R. hirsutus , and R. rosifolius . The results indicate that the Rubus seed morphology atlas developed in this investigation can serve as a critical reference for the identification of Rubus seeds that have been excavated from archaeological sites in China. Archaeobotany Rubus Seed Morphological Atlas Identification Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction Fruit and seed remains (whole or fragmentary) constitute a substantial proportion of plant remains excavated from archaeological sites 1 – 4 . other plant remains that have been excavated include rhizomes 5 , tubers 6 , bulbs 7 , fibers 8 , 9 , and leaves or stems 10 . While fruit and seed remains recovered from archaeological sites are most commonly associated with food crops, they also include less frequent remains from wild plants. Notably, the majority of such fruit and seed remains can only be identified to the family or genus taxonomic level 11 . The main reason for the difficulty in identification is that the fruit and seed of closely related sibling species usually exhibit similar morphological characteristics, so identifying the accurate provenance of fruit and seed remains in archaeological sites is a challenge. According to relevant research reports, abundant Rubus seeds have been excavated from archaeological sites across China 12 – 22 . Among these Rubus seed remains, some are carbonized and others uncarbonized, yet both exhibit relatively well-preserved morphological features. In the aforementioned literature, only two sites yielded species-level identifications, while the remainder were identified only to the genus Rubus . Notably, China is home to a highly diverse Rubus flora, comprising over 200 species 23 , 24 . To date, there is no established morphological atlas of Rubus seeds in China, which affects the accurate identification of Rubus seeds excavated at Chinese archaeological sites. Identifying the seed remains of Rubus plants in archaeological sites can provide a reference for reconstructing the history of ancient ancestors' utilization of plants in the genus Rubus . This article aims to systematically organize the seed morphological characteristics of 126 species of Rubus plants in China and provide a scientific basis for the identification of Rubus seeds excavated from archaeological sites. The objectives include (1) establishing a seed morphology atlas for identifying Rubus seed in archaeological sites and (2) understanding the history of ancient Chinese ancestors using plants of the genus Rubus . 2. Research Ideas and Methods 2.1. Research Ideas Rubus L. is a large genus in the Rosaceae family. There are 208 species of Rubus plants in China, among which 139 species are endemic 23 , 24 , mainly deciduous shrubs, rarely evergreen shrubs, semi-shrubs, or perennial creeping herbs 25 – 29 . According to the sequence of phylogenetic evolution, Yu classified the Rubus plants in China into 8 sections and 24 subsections, including Sect. Idaeobatus, Sect. Lampobatus, Sect. Rubus, Sect. Malachobatus, Sect. Dalibardastrum, Sect. Chamaebatus, Sect. Cylactis, and Sect. Chamaemorus 30 , revising and establishing a taxonomic system for Rubus in China. The difficulties in identifying plants of the genus Rubus lie in their numerous species, similar morphological characteristics, small natural distribution range, rich morphological variation, and extensive convergent evolution, as well as polyploid hybridization and obvious frequent facultative apomixis 31 – 34 . From a morphological perspective, the genus Rubus exhibits enormous diversity, ranging from woody, semi-woody, shrubs to creeping dwarf herbaceous plants 28 . Seeds are vital organs for plant regeneration. Typically, the shape and structure of seeds serve as the basis for identifying many plant species. The sculptural patterns on the seed coat of many plants are highly diverse. Additionally, the structure and surface carving on the seed coat are maternal characteristics, so they vary among different species, and the seed coat ornamentation is the most prominent and variable feature for distinguishing genera and species 35 . There have been few descriptions of the seed morphology in the genus Rubus 36 – 38 , and comprehensive research is lacking. Therefore, this study characterized the genus Rubus based on seed morphology (e.g., shape, seed coat ornamentation) and evaluated the feasibility of using these traits to determine the taxonomic provenance of Rubus seeds from excavated archaeological sites. The core objective of this study is to accurately determine the taxonomic provenance of Rubus seed remains from archaeological sites, leveraging the known morphological characteristics of seeds within this genus. 2.2. Research methods To further investigate the seed morphology of Rubus , we randomly selected several representative seeds during sample collection. Priority was given to highly mature seeds to ensure that subsequent observations and measurements accurately reflect the typical characteristics of seeds in this taxonomic unit (the genus Rubus ). For observation and measurement, a stereomicroscope (model: Keyence Digital Microscopy System-VHX-7000N) was used. Seeds were positioned on the microscope stage, and comprehensive observations were conducted from multiple angles by adjusting the focus and angle of the microscope. Using the microscope’s integrated measurement software, precise measurements were taken along the long and short axes of seeds on high-resolution images. By switching the microscope to high magnification, the seed epidermis was focused on to meticulously examine its textural structure. Seed morphology terminology in this study follows the descriptions provided by Chernoff et al. 39 and Lersten et al. 40 . 3. Research on the seeds of the genus in species 3. Research on the seeds of the genus Rubus in species 3.1. Collection of samples Seed specimens are the material basis for conducting seed morphology identification and classification research, as well as an important reference for identifying seeds excavated from archaeological sites. This study collected mature seeds of Rubus from local populations in different regions of China. A total of 126 seeds (including 22 varieties) of Rubus were collected. According to the section and subsection classification system of Rubus in Flora of China, the collected seeds were classified at the species level, after which the micromorphological variation of seeds across 8 sections and 23 subsections was evaluated. The information on the investigated species is listed in Table 1 . Table 1 Seed source species and morphological determination of the genus Rubus L. No Species Section Subsection location 1 R. calycinus Sect. Chamaebatus Hunan province 2 R. calycinoides Sect. Chamaebatus Taiwan province 3 R. pectinellus Sect. Chamaebatus Hunan province 4 R. chamaemorus Sect. Chamaemorus Inner Mongolia 5 R. rubrisetulosus Sect. Cylactis Sichuan province 6 R. saxatilis Sect. Cylactis Heilongjiang province 7 R. nyalamensis Sect. Cylactis Xizang autonomous region 8 R. fockeanus Sect. Cylactis Sichuan province 9 R. tricolor Sect. Dalibardastrum Sichuan province 10 R. jinfoshanensis Sect. Dalibardastrum Chongqing city 11 R. multisetosus Sect. Dalibardastrum Yunnan province 12 R. tsangorum Sect. Dalibardastrum Jiangxi province 13 R. lucens Sect. Lampobatus Yunnan province 14 R. caesius Sect. Rubus Xinjiang autonomous region 15 R. pentagonus var. modestus Sect. Idaeobatus Subsect. Alpestres Sichuan province 16 R. quinquefoliolatus Sect. Idaeobatus Subsect. Alpestres Yunnan province 17 R. pentagonus Sect. Idaeobatus Subsect. Alpestres Sichuan province 18 R. corchorifolius Sect. Idaeobatus Subsect. Corchorifolii Anhui province 19 R. chingii Sect. Idaeobatus Subsect. Corchorifolii Anhui province 20 R. grayanus Sect. Idaeobatus Subsect. Corchorifolii Anhui province 21 R. glabricarpus Sect. Idaeobatus Subsect. Corchorifolii Fujian province 22 R. trianthus Sect. Idaeobatus Subsect. Corchorifolii Henan province 23 R. idaeus var. glabratus Sect. Idaeobatus Subsect. Idaeanthi Heilongjiang province 24 R. sachalinensis Sect. Idaeobatus Subsect. Idaeanthi Heilongjiang province 25 R. austrotibetanus Sect. Idaeobatus Subsect. Idaeanthi Xizang autonomous region 26 R. eucalyptus var. trullisatus Sect. Idaeobatus Subsect. Idaeanthi Sichuan province 27 R. niveus Sect. Idaeobatus Subsect. Idaeanthi Xizang autonomous region 28 R. trijugus Sect. Idaeobatus Subsect. Idaeanthi Xizang autonomous region 29 R. eucalyptus var. yunnanensis Sect. Idaeobatus Subsect. Idaeanthi Yunnan province 30 R. idaeus Sect. Idaeobatus Subsect. Idaeanthi Hebei province 31 R. idaeus var. borealisinensis Sect. Idaeobatus Subsect. Idaeanthi Hebei province 32 R. piluliferus Sect. Idaeobatus Subsect. Idaeanthi Gansu province 33 R. irritans Sect. Idaeobatus Subsect. Idaeanthi Gansu province 34 R. leucanthus Sect. Idaeobatus Subsect. Leucanthi Anhui province 35 R. delavayi Sect. Idaeobatus Subsect. Leucanthi Yunnan province 36 R. zhaogoshanensis Sect. Idaeobatus Subsect. Leucanthi Yunnan province 37 R. columellaris Sect. Idaeobatus Subsect. Leucanthi Jiangxi province 38 R. impressinervius Sect. Idaeobatus Subsect. Leucanthi Hunan province 39 R. peltatus Sect. Idaeobatus Subsect. Peltati Zhejiang province 40 R. komarovi Sect. Idaeobatus Subsect. Pileati Jilin province 41 R. pseudopileatus Sect. Idaeobatus Subsect. Pileati Sichuan province 42 R. pileatus Sect. Idaeobatus Subsect. Pileati Shaanxi province 43 R. macilentus Sect. Idaeobatus Subsect. Pungentes Yunnan province 44 R. pinfaensis Sect. Idaeobatus Subsect. Pungentes Gansu province 45 R. pungens Sect. Idaeobatus Subsect. Pungentes Shaanxi province 46 R. stans Sect. Idaeobatus Subsect. Pungentes Sichuan province 47 R. amabilis var. aculeatissimus Sect. Idaeobatus Subsect. Pungentes Sichuan province 48 R. coreanus var. tomentosus Sect. Idaeobatus Subsect. Pungentes Anhui province 49 R. coreanus Sect. Idaeobatus Subsect. Pungentes Anhui province 50 R. amabilis Sect. Idaeobatus Subsect. Pungentes Shaanxi province 51 R. inopertus Sect. Idaeobatus Subsect. Pungentes Hunan province 52 R. tsangii Sect. Idaeobatus Subsect. Rosaefolii Hunan province 53 R. eustephanus var. glanduliger Sect. Idaeobatus Subsect. Rosaefolii Sichuan province 54 R. glandulosocalycinus Sect. Idaeobatus Subsect. Rosaefolii Taiwan province 55 R. piptopetalus Sect. Idaeobatus Subsect. Rosaefolii Taiwan province 56 R. rosaefolius Sect. Idaeobatus Subsect. Rosaefolii Anhui province 57 R. hirsutus Sect. Idaeobatus Subsect. Rosaefolii Anhui province 58 R. sumatranus Sect. Idaeobatus Subsect. Rosaefolii Anhui province 59 R. glandulosocarpus Sect. Idaeobatus Subsect. Rosaefolii Guizhou province 60 R. eustephanus Sect. Idaeobatus Subsect. Rosaefolii Anhui province 61 R. erythrocarpus Sect. Idaeobatus Subsect. Stimulantes Yunnan province 62 R. kulinganus Sect. Idaeobatus Subsect. Stimulantes Jiangxi province 63 R. mesogaeus var. oxycomus Sect. Idaeobatus Subsect. Stimulantes Gansu province 64 R. ellipticus var. obcordatus Sect. Idaeobatus Subsect. Stimulantes Sichuan province 65 R. alexeterius var. acaenocalyx Sect. Idaeobatus Subsect. Stimulantes Yunnan province 66 R. biflorus Sect. Idaeobatus Subsect. Stimulantes Gansu province 67 R. parvifolius Sect. Idaeobatus Subsect. Stimulantes Anhui province/ Hunan province 68 R. mesogaeus Sect. Idaeobatus Subsect. Stimulantes Shanxi province 69 R. subornatus var. melanadenus Sect. Idaeobatus Subsect. Stimulantes Sichuan province 70 R. ellipticus Sect. Idaeobatus Subsect. Stimulantes Sichuan province 71 R. idaeopsis Sect. Idaeobatus Subsect. Thyrsidaei Guizhou province 72 R. innominatus var. macrosepalus Sect. Idaeobatus Subsect. Thyrsidaei Zhejiang province 73 R. innominatus var. aralioides Sect. Idaeobatus Subsect. Thyrsidaei Fujian province 74 R. chiliadenus Sect. Idaeobatus Subsect. Thyrsidaei Hunan province 75 R. flosculosus Sect. Idaeobatus Subsect. Thyrsidaei Shanxi province 76 R. grandipaniculatus Sect. Idaeobatus Subsect. Thyrsidaei Sichuan province 77 R. cockburnianus Sect. Idaeobatus Subsect. Thyrsidaei Henan province 78 R. adenophorus Sect. Idaeobatus Subsect. Thyrsidaei Fujian province 79 R. innominatus Sect. Idaeobatus Subsect. Thyrsidaei Anhui province 80 R. lasiostylus var. hubeiensis Sect. Idaeobatus Subsect. Wushanenses Hubei province 81 R. lasiostylus var. dizygos Sect. Idaeobatus Subsect. Wushanenses Sichuan province 82 R. lasiostylus Sect. Idaeobatus Subsect. Wushanenses Gansu province 83 R. poliophyllus Sect. Malachobatus Subsect. Acuminati Yunnan province 84 R. lambertianus var. paykouangensis Sect. Malachobatus Subsect. Acuminati Chongqing city 85 R. laxus Sect. Malachobatus Subsect. Acuminati Yunnan province 86 R. lambertianus Sect. Malachobatus Subsect. Acuminati Anhui province 87 R. cochinchinensis Sect. Malachobatus Subsect. Cochinchinenses Hainan province 88 R. angustibracteatus Sect. Malachobatus Subsect. Dolichophylli Sichuan province 89 R. dolichophyllus var. pubescens Sect. Malachobatus Subsect. Dolichophylli Guizhou province 90 R. parkeri Sect. Malachobatus Subsect. Dolichophylli Hunan province 91 R. howii Sect. Malachobatus Subsect. Dolichophylli Sichuan province 92 R. ichangensis Sect. Malachobatus Subsect. Dolichophylli Jiangxi province 93 R. feddei Sect. Malachobatus Subsect. Elongati Guizhou province 94 R. salwinensis Sect. Malachobatus Subsect. Elongati Yunnan province 95 R. luchunensis var. coriaceus Sect. Malachobatus Subsect. Elongati Yunnan province 96 R. fuscorubens Sect. Malachobatus Subsect. Elongati Chongqing city 97 R. lobophyllus Sect. Malachobatus Subsect. Elongati Guangxi province 98 R. tephrodes Sect. Malachobatus Subsect. Elongati Anhui province 99 R. paniculatus Sect. Malachobatus Subsect. Elongati Yunnan province 100 R. reflexus var. lanceolobus Sect. Malachobatus Subsect. Fuscifolii Anhui province 101 R. malipoensis Sect. Malachobatus Subsect. Fuscifolii Yunnan province 102 R. reflexus Sect. Malachobatus Subsect. Fuscifolii Anhui province 103 R. jambosoides Sect. Malachobatus Subsect. Jambosoides Guangdong province 104 R. lineatus var. glabrescens Sect. Malachobatus Subsect. Lineati Yunnan province 105 R. lineatus var. angustifolius Sect. Malachobatus Subsect. Lineati Yunnan province 106 R. lineatus Sect. Malachobatus Subsect. Lineati Yunnan province 107 R. wardii Sect. Malachobatus Subsect. Metoenses Yunnan province 108 R. echinoides Sect. Malachobatus Subsect. Moluccani Guangxi province 109 R. nagasawanus Sect. Malachobatus Subsect. Moluccani Taiwan province 110 R. lasiotrichos Sect. Malachobatus Subsect. Moluccani Yunnan province 111 R. hunanensis Sect. Malachobatus Subsect. Moluccani Hunan province 112 R. buergeri Sect. Malachobatus Subsect. Moluccani Anhui province 113 R. setchuenensis Sect. Malachobatus Subsect. Moluccani Hubei province 114 R. faberi Sect. Malachobatus Subsect. Moluccani Sichuan province 115 R. alceaefolius Sect. Malachobatus Subsect. Moluccani Jiangxi province 116 R. multibracteatus Sect. Malachobatus Subsect. Moluccani Anhui province 117 R. reticulatus Sect. Malachobatus Subsect. Moluccani Sichuan province 118 R. pirifolius Sect. Malachobatus Subsect. Pirifolii Fujian province 119 R. doyonensis Sect. Malachobatus Subsect. Sozostyli Yunnan province 120 R. swinhoei Sect. Malachobatus Subsect. Sozostyli Anhui province 121 R. preptanthus Sect. Malachobatus Subsect. Sozostyli Yunnan province 122 R. henryi Sect. Malachobatus Subsect. Sozostyli Hunan province 123 R. fujianensis Sect. Malachobatus Subsect. Sozostyli Fujian province 124 R. irenaeus Sect. Malachobatus Subsect. Stipulosi Anhui province 125 R. pacificus Sect. Malachobatus Subsect. Stipulosi Anhui province 126 R. fanjingshanensis Guizhou province 3.2 Identification of the seeds of the genus Rubus in China In this study, the seed micromorphology of Rubus taxa native to China was first systematically documented. At the micromorphological level, certain morphological features of Rubus seeds (such as size, shape, color, etc.) may overlap across taxa, but the surface ornamentation of the seed coat exhibits significant variation among different species (Fig. 1 – 4 ). 3.2.1. Seed size The seed length ranges from 1.23 mm to 6.11 mm, with R. chamaemorus having the longest seeds, while R. howii has the shortest. Based on length, the seeds can be divided into four categories: Class A: Above 3.5 mm (e.g., R. pirifolius , R. quinquefoliolatus , R. chamaemorus ); Class B: 2.5 mm − 3.5 mm (e.g., R. parkeri , R. henryi , R. flosculosus ); Class C: 1.5 mm − 2.5 mm (e.g., R. pacificus , R. adenophorus , R. piluliferus ); Class D: Below 1.5 mm (e.g., R. reticulatus , R. sumatranus , R. ellipticus ). 3.2.2. Seed shape Most seeds exhibit an elliptical shape (e.g., R. mesogaeus , R. trijugus ), semi-elliptical shape (e.g., R. tephrodes , R. idaeus ), semi-circular shape (e.g., R. lambertianus , R. trianthus ), ovate shape (e.g., R. feddei , R. parkeri ), long-oval shape (e.g., R. cinclidodictyus ), or shield shape (e.g., R. impressinervius , R. glabricarpus ). A few seeds show a square shape (e.g., R. ichangensis ), crescent shape (e.g., R. henryi , R. laxus ), long-strip shape (e.g., R. idaeus var. borealisinensis , R. multisetosus ), or irregular shape (e.g., R. retinulatus ). 3.2.3. Surface ornamentation of seeds The seed coat ornamentation of Rubus exhibits specific changes, which seem to have taxonomic significance. The seed coat ornamentation of seeds is reticular or striped, but different species have their own characteristics in terms of the depth of the holes, the composition of the reticular pattern, and the thickness, arrangement, and direction of the stripes. It mainly includes two categories: The first group is seeds with prominent reticulate sculpture (e.g., R. reflexus , R. parvifolius , R. flosculosus ). The second group is seeds with less prominent reticulate sculpture (e.g., R. salwinensis , R. cinclidodictyus , R. fujianensis ). Among them, the first group can be further divided into two types based on the depth of the holes: deep-hole type (e.g., R. lobophyllus , R. cochinchinensis ) and shallow-hole type (e.g., R. faberi , R. lambertianus ). 4. Identification of Seeds from Archaeological Sites 4. Identification of Rubus Seeds from Archaeological Sites The climate in China has undergone certain changes throughout history, which may have led to subtle differences in the distribution range of Rubus plants between ancient and modern times, thereby affecting the identification of seeds excavated from archaeological sites. Therefore, this study evaluated the seed characteristics of Rubus taxa distributed in different regions. The results indicated that the seeds selected for this study were representative, covering the morphological variation range of the genus across geographic populations. Considering that seed size is closely related to the plant's growth environment and may be affected by environmental factors during burial, causing expansion or shrinkage, in this study's identification work, seed morphology and seed coat ornamentation were mainly used as identification criteria, while seed size was only used as a reference. This study collected Rubus seed samples from seven archaeological sites (Table 2 ). Table 2 Plant Remains Information of Rubus Serial number Site name Location 14 C Dating Period Origin(Grains) Fruit period 1 Wutongtu Site * Jiangjing Chongqing 1416 − 1235 cal BC (3190 − 3184 cal BP) the Shang and Zhou dynasties R. coreanus var. tomentosus (1 grains) Jun. to Aug. 2 Nanshan Site Shaoxing, Zhejiang - the Eastern Zhou Dynasty R. lambertianus (25 grains) Sep. to Nov. 3 Tianshengcheng Site Wanzhou, Chongqing - the Song Dynasty R. flosculosus (5 grains)/ R. parvifolius ༈17 grains༉ Aug. to Sep. / Jul. to Aug. 4 Fanjiayan Site Hechuan, Chongqing 1222–1286 cal AD(728 − 664 cal BP) the Southern Song Dynasty R. trianthus (15 grains)/ R. parvifolius ༈15 grains༉ May to Jun. / Jul. to Aug. 5 Taipingmen Site Yuzhong, Chongqing 1165–1265 cal AD (785 − 685 cal BP) the Southern Song Dynasty R. adenophorus (6 grains) Jun. to Jul. 6 Drum Tower Yamen Site Yuzhong, Chongqing 1221–1282 cal AD (729 − 668 cal BP) the Southern Song Dynasty R. trianthus (20 grains) May to Jun. 7 Yuanmingyuan site Haidian, Beijing - The Ming and Qing Dynasties / the Modern Times R. lambertianus (1 grains /1 grains) Sep. to Nov. 4.1. Wutongtu Site The Wutongtu Site is located on a crescent-shaped platform on the left bank of the Yangtze River in Youxi Town, Jiangjin District, Chongqing City. In this study, one Rubus seed was selected from an ash pit dating to the Shang and Zhou dynasties. Although the seed has been carbonized, the seed coat ornamentation remains clearly visible under microscopy. Observations show that it is elliptical in shape (Fig. 5 j), with reticular intersecting ornamentation. Based on these characteristics and comparison with modern specimens, it is identified as R. coreanus var. tomentosus (Table 2 ). 4.2. Nanshan Site The Nanshan Site is located in Yuecheng District, Shaoxing City, Zhejiang Province. In this study, 25 Rubus seeds were obtained from the site dating to the Eastern Zhou Dynasty. The Rubus seeds are uncarbonized, with distinct seed coat ornamentation. They are half-moon shaped (Fig. 5 g), with bluntly rounded ends, and the seed coat ornamentation branches from the hilum in a tree-branch pattern. Based on these characteristics and correspondence with morphological descriptions of R. lambertianus in taxonomic literature, the seeds excavated from the Nanshan Site are inferred to originate from R. lambertianus (Table 2 ). 4.3. Tianshengcheng Site The Tianshengcheng Site is located in Zhoujiaba Street, Wanzhou District, Chongqing City, adjacent to the confluence of the Zhuxi River and the Yangtze River to the south. It is an important Song and Yuan dynasties mountain city site. In this study, 22 Rubus seeds were selected from Southern Song Dynasty water wells. The seeds are uncarbonized. Based on morphological characteristics, they can be divided into two types: semicircular (Fig. 5 c) and oval (Fig. 5 d). Semi-circular type: Seeds are semi-circular, with bluntly rounded ends and a small aspect ratio. The seed coat ornamentation consists of intersecting polygonal meshes with relatively dense reticulation. Identification confirms they belong to R. flosculosus . Oval type: Seeds are oval, with one end sharp and the other bluntly rounded. The seed coat exhibits polygonal intersecting patterns with relatively loose overall reticulation. Identification confirms they belong to R. parvifolius (Table 2 ). 4.4. Fanjiayan Site The Fanjiayan Site is located on Diaoyu Mountain in the northeastern part of Dongcheng Peninsula, Hechuan District, Chongqing City. As a significant component of the Fishing Town Site, it is a well-preserved relic of governmental establishments from the Song and Yuan Dynasties. In this study, 30 Rubus seeds were selected from a Southern Song Dynasty pool (archaeologically identified as a water storage facility). Notably, the excavated Rubus seeds are uncarbonized and exhibit diverse shapes. Based on morphology, they can be categorized into two types: oval (Fig. 5 a) and semi-oval (Fig. 5 b). Oval type: Seeds are oval, with one end pointed and the other bluntly rounded. The seed coat exhibits intersecting polygonal patterns with relatively sparse reticulation, consistent with the seed characteristics of R. parvifolius . Semi-oval type: Seeds are semi-oval, with one end slightly protruding and the other bluntly rounded, showing a large aspect ratio. The seed coat ornamentation features branching patterns originating from the hilum, with overall sparse reticulation, matching the seed characteristics of R. trianthus (Table 2 ). 4.5. Taipingmen Site The Taipingmen Site is located on Wanglongmen Street, Yuzhong District, Chongqing City. Situated on the first river terrace of the left bank of the Yangtze River in the Yuzhong Peninsula, it is one of the important city gates of Chongqing. In this study, six Rubus seeds were selected from Southern Song Dynasty drainage ditches of the city walls. The seeds are uncarbonized, semi-oval with slightly blunt ends. The seed coat ornamentation is indistinct, faintly appearing as an irregular cross pattern (Fig. 5 f). Based on these characteristics and comparison with R. adenophorus specimens in collections, the excavated Rubus seeds are inferred to derive from R. adenophorus (Table 2 ). 4.6. Drum Tower Yashu Site The Drum Tower Yashu Site is located in Wanglongmen Street, Yuzhong District, Chongqing City. Situated on a terrace at the foot of Jinbi Mountain and the left bank of the Yangtze River, it served as the administrative center of Sichuan and Chongqing Prefecture during the Southern Song Dynasty. In this study, 20 Rubus seeds were selected from Southern Song Dynasty ponds. The seeds are uncarbonized, semi-oval with one end slightly protruding and the other blunt and round, and have a large aspect ratio. The seed coat ornamentation features a network-like, cross-shaped pattern with relatively loose overall reticulation. Based on these characteristics and comparison with specimens of R. trianthus, they are identified as originating from R. trianthus (Table 2 ). 4.7. Yuanmingyuan Site The Yuanmingyuan Site is located in Haidian District, Beijing City. One Rubus seed was excavated from each of the Ming and Qing Dynastic relics (Fig. 5 h) and modern deposits (Fig. 5 i). The seeds are uncarbonized, all crescent-shaped with bluntly rounded ends. The seed coat ornamentation branches from the hilum in a tree-branch pattern, with ornamentation lines approximately parallel. Based on morphological consistency across historical and modern specimens, the Rubus seeds excavated from both periods at this site are identified as R. lambertianus (Table 2 ). 5. Discussion 5.1. Application of This Identification Methodology to Rubus Seed Remains from Other Archaeological Sites in China This study preliminarily demonstrates that the established identification method is effective for identifying seeds of the genus Rubus excavated from archaeological contexts, based on the collection and analysis of a large dataset of modern Rubus seed samples. However, the archaeological Rubus seed assemblages discussed above are predominantly from Chongqing, Zhejiang, and Beijing. To further validate the universality of this methodology, we applied it to the re-identification of Rubus seed remains excavated from additional archaeological sites across China, spanning diverse geographical regions and chronological periods (Table 3 ). Table 3 Re-identification of Rubus seed remains in reported archaeological sites. No Site Location Grains re-identification Result references 1 Bashidang Site Changde, Hunan - R. parvifolius 1 2 Yuchisi Site Bozhou, Anhui 2 R. parvifolius 11,13 3 Liangzhu Ancient City Site Hangzhou, Zhejiang 24 R. parvifolius 21 4 Chengtoushan Site & Changde, Hunan - R. parvifolius 1 5 Guangfulin Site Shanghai 81 R. hirsutus 14 6 Daxinzhuang Site * Jinan, Shandong 18 R. parvifolius 18 7 Changdun Han Tomb Cemetery Nanchang, Jiangxi 11 R. rosifolius 16 8 Nanyue Palace Site: J264 Water Well Guangzhou, Guangdong 15000 R. rosifolius 20,13 9 Nanyue Palace Garden: Nanhan Kingdom Palace Pool Site Guangzhou, Guangdong 1 R. rosifolius 12,13 *Carbonized; & Identification error Through identification, Rubus seeds excavated from the Bashidang Site, Yuchisi Site, Liangzhu Ancient City Site, Chengtoushan Site, and Daxinzhuang Site all exhibit a pointed-obtuse shape. The seed coat ornamentation consists of polygonal intersections with an overall loose pattern. These seeds are uniformly identified as R. parvifolius . Notably, the previously reported identification of Rubus seeds from the Chengtoushan Site as R. lambertianus is revised herein. This study clarifies that R. lambertianus seeds are distinctively characterized by bluntly rounded ends and forked, hilum-branching ornamentation, whereas the Chengtoushan seeds display reticulate interwoven ornamentation, consistent with R. parvifolius rather than R. lambertianus . Therefore, this study posits that the provenance of the Rubus seeds excavated from the Chengtoushan Site should be R. parvifolius rather than R. lambertianus . The Rubus seeds excavated from the Guangfulin Site exhibit a semicircular shape with seed coat ornamentation featuring reticulate intersections in a hexagonal pattern, consistent with the seed morphology of R. hirsutus . Based on these diagnostic traits—specifically the semicircular outline and dense hexagonal reticulation distinct from other sympatric species (e.g., R. parvifolius with looser polygonal patterns)—these seeds are taxonomically identified as R. hirsutus . The Rubus seeds excavated from the Changdun Han Tomb Cemetery, Nanyue Palace Site (Well J264), and Nanyue Palace Garden (Palace Pool Site of the Southern Han Kingdom) exhibit a relatively high length-to-width ratio with bluntly rounded ends. Their seed coat ornamentation features densely interwoven reticulate patterns, diagnostic of R. rosifolius . These morphological traits—specifically the elongated oval shape and anastomosing reticulation distinct from other congeners (e.g., R. lambertianus with bifurcating hilum patterns)—consistently align with the seed characteristics of R. rosifolius . The above identification results fully demonstrate that the identification method established in this study can be effectively applied to the identification of Rubus seed remains in Chinese archaeological sites (Table 3 ). 5.2 Established an illustrated guide to the seeds of common Rubus plants in China Stereoscopic microscopy and scanning electron microscopy (SEM) have revealed many important seed characteristics, which contribute to resolving taxonomic problems in botanical origin identification 41 , 42 . This article is the first to use a stereomicroscope to study the microscopic morphological characteristics of seeds in Chinese plants of the genus Rubus and systematically describes the microscopic morphological characteristics of many taxonomic groups. In recent years, the use of stereomicroscopes as a means of studying microstructural changes in plants has gained advantages in plant systems 43 – 45 , with observed data achieving high precision and being irreplaceable by other techniques. At different classification levels, there are significant differences in the seed micromorphology of the genus Rubus in terms of different traits such as seed size, shape, and seed coat ornamentation. The micromorphological characteristics of seeds are key features for plant species identification and classification, reflecting the genetic, physiological, and ecological characteristics of plants. This paper uses stereomicroscopy to study the micromorphological characteristics of the seeds of Rubus plants in China and systematically describes the micromorphological characteristics of many taxonomic groups for the first time. In recent years, the use of stereomicroscopy as a means to study plant micromorphological changes has gained an advantage in plant systems 43 – 45 . Stereomicroscopic observations yield highly precise data and cannot be replaced by other technologies. At different taxonomic levels, there are significant differences in the seed micromorphology of the genus Rubus in different traits (such as seed size, shape, and seed coat ornamentation). Seed micromorphological characteristics are key features for plant species identification and classification, reflecting the genetic, physiological, and ecological characteristics of plants. The micromorphological characteristics of seeds can provide useful information for the species classification of this genus. For example, although R. ichangensis and R. angustibraceteatus are very similar in plant morphological characteristics, their seed morphological features are completely different. However, many species assigned to different sections or subsections with distantly related macromorphological relationships exhibit similar seed morphological characteristics, such as R. multibraceteatus and R. mesogeaus , R. fuscorubens and R. parkeri . In other words, apart from the seed coat ornamentation, no section (except Sect. Ghamaemorus) or subsection (except Subsect. Pirifolii) exhibits unique seed morphology, and species of different subgenera or families have similar seed morphology (such as size, shape, color, etc.). Whether seed coat ornamentation can be used for intrageneric classification depends on the degree of variation between species 46 . Although this study observed the seed morphology of common plants in the Chinese Rubus genus, which covers almost all sections and subsections of the genus, it remains difficult to identify the evolutionary patterns of seed size, shape, and seed coat ornamentation, as well as their correlations with macromorphological characteristics. The accurate identification of plant seed remains requires the support of a large number of modern seed specimens. Based on this, this study constructed the first seed morphology atlas of the genus Rubus in China according to key characteristics such as seed shape and seed coat ornamentation and applied it to the identification of seed remains in archaeological sites. Through the method of comparing ancient and modern specimens, this study provides an important reference for the identification of Rubus seeds unearthed from sites. 5.3 The Uses of Ancient Rubus Plants In this study, some Rubus seed remains were unearthed from water pool remains at archaeological sites 12 , 13 . Meanwhile, in some sites, Rubus seeds were unearthed with abundant crop and fruit remains. Based on these comprehensive archaeological findings, it is inferred that the fruits or seeds of Rubus at the sites were intentionally collected and preserved 16 , 18 . Additionally, identification shows that the fruits of the original plants corresponding to Rubus seeds from archaeological sites are all red (Table 2 ), with a sweet and sour taste. Based on this, it can be inferred that the fruit of the original seed plant of the genus Rubus is highly likely to have been an important component of the ancient ancestors' plant-based diet. The Rubus genus is a widely distributed plant and grows in mountainous areas, hills, and wasteland hillsides in most parts of southern and central China. The fruiting period of Rubus plants varies from May to November, making them common wild fruits. Its fruits are plump and juicy and taste delicious. It is highly probable that ancient people would pick and eat ripe Rubus fruits during their outdoor activities. Additionally, from the perspective of traditional medicine, some fruits of the Rubus genus have medicinal values and are rich in nutrients 47 , 48 . Numerous studies have shown that Rubus seed remains are prevalent in Chinese archaeological sites 12 – 22 . In some sites, a large number of seeds were unearthed; this is possibly because each Rubus fruit contains numerous seeds. This quantitative advantage naturally increases the probability of Rubus seeds being discarded in archaeological deposits, resulting in their general dominance in absolute numbers among unearthed seeds 13 , 20 . In addition, the seeds of plants in the genus Rubus are relatively small in size. When consuming the fruits, ancient people generally did not deliberately spit out the seeds, which were excreted through bodily waste. In natural environments, these seeds might be wrapped in other excrement; this process reduces erosion from the external environment and thus enables relatively complete preservation in archaeological sites. It is worth noting that plants of the genus Rubus also have the possibility of being cultivated as ornamental plants 20 . The abundant and well-preserved Rubus seed remains in some sites may be linked to special burial contexts or specific functions. Nevertheless, clarifying their specific roles demands more archaeological evidence and targeted research. 6. Conclusions In this study, 126 mature seeds from wild Rubus populations across various regions of China were collected for systematic seed morphology analysis, leading to the construction of the first morphological atlas of seeds for common Chinese Rubus species. Through the analysis of Rubus seed remains from 7 unreported sites and 9 reported sites, the taxonomic identity of seeds was accurately identified. This study employed modern plant seed morphology research methods to link modern samples to archaeological samples, thereby providing new ideas and methods for plant archaeology research. As seed identification of Rubus from more sites progresses and research deepens, this study is expected to further reveal the edible history and species of Rubus fruits, provide important historical background information for exploring the distribution shifts and utilization patterns of Rubus plants, and offer strong archaeological evidence for studying the biodiversity evolution of Rubus . Declarations Conflicts of Interest: The authors declare no conflicts of interest. Fundings: The National Key Research and Development Program of China (2022YFC3500900、2022YFC3500902), Key project at central government level: The ability establishment of sustainable use for valuable Chinese medicine resources(2060302), China Academy of Chinese Medical Sciences: Science and Technology Innovation Project(CI2024E001), The Major Increase and Reduction 516 Project at the Central Level (2060302) and the Innovation Team and Talents Cultivation Program 517 of National Administration of Traditional Chinese Medicine (ZYYCXTD-C-202409). Author Contribution A. R.: Writing-original draft, Writing-review and editing, Investigation; X. M.: Writing-original draft, Writing-review and editing, Investigation;; X. X.: Resources, Data curation; M. Y.: Writing-original draft, Writing-review and editing, Methodology, Investigation, Funding acquisition; H. 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Moreno-Medina B, Casierra-Posada F, Cutler J. Phytochemical Composition and Potential Use of Rubus Species. Gesunde Pflanz. 2018;70:65–74. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 25 Jan, 2026 Reviews received at journal 21 Jan, 2026 Reviews received at journal 15 Jan, 2026 Reviews received at journal 12 Jan, 2026 Reviewers agreed at journal 25 Dec, 2025 Reviewers agreed at journal 24 Dec, 2025 Reviewers agreed at journal 23 Dec, 2025 Reviewers agreed at journal 23 Dec, 2025 Reviewers invited by journal 23 Dec, 2025 Editor assigned by journal 21 Dec, 2025 Submission checks completed at journal 21 Dec, 2025 First submitted to journal 15 Dec, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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(II)\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8367387/v1/5e050ef3e518fabe439d79ee.jpeg"},{"id":99311912,"identity":"4288b314-744c-42d7-b9d5-5156dfb9e9f0","added_by":"auto","created_at":"2025-12-31 16:17:22","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":576031,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of Seed Morphology in the \u003cem\u003eRubus\u003c/em\u003e L. (III)\u003c/p\u003e","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8367387/v1/0cb6666576bf81071b3fe995.jpeg"},{"id":98999772,"identity":"663dae2b-985d-4a99-8d7a-cd08d2e5ae7b","added_by":"auto","created_at":"2025-12-25 14:04:17","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":160335,"visible":true,"origin":"","legend":"\u003cp\u003eComparison of Seed Morphology in the \u003cem\u003eRubus\u003c/em\u003e L. (IV)\u003c/p\u003e","description":"","filename":"Onlinefloatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8367387/v1/d99b1657a9059f5dc9a1dd23.png"},{"id":98999770,"identity":"9c74f1fc-f1cb-4ba1-9706-76b4f87c1afb","added_by":"auto","created_at":"2025-12-25 14:04:16","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":199259,"visible":true,"origin":"","legend":"\u003cp\u003eSeeds of the genus \u003cem\u003eRubus\u003c/em\u003e excavated from archaeological sites (a, b: Fanjiayan site; c: Drum Tower Yashu Site; d. E: Tianshengcheng Site; f: Taipingmen Site; g: Nanshan Site; h, I: Yuanmingyuan Ruins; j: Wutongtu Site)\u003c/p\u003e\n\u003cp\u003eNote: a, d, g; b, c, e, f; h, I, j: each has a consistent scale.\u003c/p\u003e","description":"","filename":"Onlinefloatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8367387/v1/531ab12e7554ed9e724191b8.png"},{"id":99322903,"identity":"7ac006bd-06dc-4b17-a0e7-c7e53e2acfd0","added_by":"auto","created_at":"2025-12-31 16:44:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3632561,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8367387/v1/325b7ab5-c4fe-4675-ae20-e85ab17e4b95.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Identification of Rubus Seeds in Chinese Archaeological Sites: Based on the Morphological Atlas of 126 Modern Rubus Seeds","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eFruit and seed remains (whole or fragmentary) constitute a substantial proportion of plant remains excavated from archaeological sites \u003csup\u003e\u003cspan additionalcitationids=\"CR2 CR3\" citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u003c/sup\u003e. other plant remains that have been excavated include rhizomes\u003csup\u003e\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e\u003c/sup\u003e, tubers\u003csup\u003e\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u003c/sup\u003e, bulbs\u003csup\u003e\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u003c/sup\u003e, fibers\u003csup\u003e\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e,\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u003c/sup\u003e, and leaves or stems\u003csup\u003e\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e\u003c/sup\u003e. While fruit and seed remains recovered from archaeological sites are most commonly associated with food crops, they also include less frequent remains from wild plants. Notably, the majority of such fruit and seed remains can only be identified to the family or genus taxonomic level\u003csup\u003e\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e\u003c/sup\u003e. The main reason for the difficulty in identification is that the fruit and seed of closely related sibling species usually exhibit similar morphological characteristics, so identifying the accurate provenance of fruit and seed remains in archaeological sites is a challenge.\u003c/p\u003e \u003cp\u003eAccording to relevant research reports, abundant \u003cem\u003eRubus\u003c/em\u003e seeds have been excavated from archaeological sites across China\u003csup\u003e\u003cspan additionalcitationids=\"CR13 CR14 CR15 CR16 CR17 CR18 CR19 CR20 CR21\" citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u003c/sup\u003e. Among these \u003cem\u003eRubus\u003c/em\u003e seed remains, some are carbonized and others uncarbonized, yet both exhibit relatively well-preserved morphological features. In the aforementioned literature, only two sites yielded species-level identifications, while the remainder were identified only to the genus \u003cem\u003eRubus\u003c/em\u003e. Notably, China is home to a highly diverse Rubus flora, comprising over 200 species\u003csup\u003e\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e,\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e. To date, there is no established morphological atlas of \u003cem\u003eRubus\u003c/em\u003e seeds in China, which affects the accurate identification of \u003cem\u003eRubus\u003c/em\u003e seeds excavated at Chinese archaeological sites. Identifying the seed remains of \u003cem\u003eRubus\u003c/em\u003e plants in archaeological sites can provide a reference for reconstructing the history of ancient ancestors' utilization of plants in the genus \u003cem\u003eRubus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThis article aims to systematically organize the seed morphological characteristics of 126 species of \u003cem\u003eRubus\u003c/em\u003e plants in China and provide a scientific basis for the identification of \u003cem\u003eRubus\u003c/em\u003e seeds excavated from archaeological sites. The objectives include (1) establishing a seed morphology atlas for identifying \u003cem\u003eRubus\u003c/em\u003e seed in archaeological sites and (2) understanding the history of ancient Chinese ancestors using plants of the genus \u003cem\u003eRubus\u003c/em\u003e.\u003c/p\u003e"},{"header":"2. Research Ideas and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Research Ideas\u003c/h2\u003e \u003cp\u003e\u003cem\u003eRubus\u003c/em\u003e L. is a large genus in the Rosaceae family. There are 208 species of \u003cem\u003eRubus\u003c/em\u003e plants in China, among which 139 species are endemic\u003csup\u003e\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e,\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e\u003c/sup\u003e, mainly deciduous shrubs, rarely evergreen shrubs, semi-shrubs, or perennial creeping herbs\u003csup\u003e\u003cspan additionalcitationids=\"CR26 CR27 CR28\" citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e\u003c/sup\u003e. According to the sequence of phylogenetic evolution, Yu classified the \u003cem\u003eRubus\u003c/em\u003e plants in China into 8 sections and 24 subsections, including Sect. Idaeobatus, Sect. Lampobatus, Sect. Rubus, Sect. Malachobatus, Sect. Dalibardastrum, Sect. Chamaebatus, Sect. Cylactis, and Sect. Chamaemorus\u003csup\u003e\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e\u003c/sup\u003e, revising and establishing a taxonomic system for \u003cem\u003eRubus\u003c/em\u003e in China.\u003c/p\u003e \u003cp\u003eThe difficulties in identifying plants of the genus \u003cem\u003eRubus\u003c/em\u003e lie in their numerous species, similar morphological characteristics, small natural distribution range, rich morphological variation, and extensive convergent evolution, as well as polyploid hybridization and obvious frequent facultative apomixis\u003csup\u003e\u003cspan additionalcitationids=\"CR32 CR33\" citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u003c/sup\u003e. From a morphological perspective, the genus \u003cem\u003eRubus\u003c/em\u003e exhibits enormous diversity, ranging from woody, semi-woody, shrubs to creeping dwarf herbaceous plants\u003csup\u003e\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e\u003c/sup\u003e. Seeds are vital organs for plant regeneration. Typically, the shape and structure of seeds serve as the basis for identifying many plant species. The sculptural patterns on the seed coat of many plants are highly diverse. Additionally, the structure and surface carving on the seed coat are maternal characteristics, so they vary among different species, and the seed coat ornamentation is the most prominent and variable feature for distinguishing genera and species\u003csup\u003e\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u003c/sup\u003e. There have been few descriptions of the seed morphology in the genus \u003cem\u003eRubus\u003c/em\u003e \u003csup\u003e\u003cspan additionalcitationids=\"CR37\" citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e\u003c/sup\u003e, and comprehensive research is lacking. Therefore, this study characterized the genus \u003cem\u003eRubus\u003c/em\u003e based on seed morphology (e.g., shape, seed coat ornamentation) and evaluated the feasibility of using these traits to determine the taxonomic provenance of \u003cem\u003eRubus\u003c/em\u003e seeds from excavated archaeological sites.\u003c/p\u003e \u003cp\u003eThe core objective of this study is to accurately determine the taxonomic provenance of \u003cem\u003eRubus\u003c/em\u003e seed remains from archaeological sites, leveraging the known morphological characteristics of seeds within this genus.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2. Research methods\u003c/h2\u003e \u003cp\u003eTo further investigate the seed morphology of \u003cem\u003eRubus\u003c/em\u003e, we randomly selected several representative seeds during sample collection. Priority was given to highly mature seeds to ensure that subsequent observations and measurements accurately reflect the typical characteristics of seeds in this taxonomic unit (the genus \u003cem\u003eRubus\u003c/em\u003e).\u003c/p\u003e \u003cp\u003eFor observation and measurement, a stereomicroscope (model: Keyence Digital Microscopy System-VHX-7000N) was used. Seeds were positioned on the microscope stage, and comprehensive observations were conducted from multiple angles by adjusting the focus and angle of the microscope. Using the microscope\u0026rsquo;s integrated measurement software, precise measurements were taken along the long and short axes of seeds on high-resolution images. By switching the microscope to high magnification, the seed epidermis was focused on to meticulously examine its textural structure. Seed morphology terminology in this study follows the descriptions provided by Chernoff et al.\u003csup\u003e39\u003c/sup\u003e and Lersten et al.\u003csup\u003e40\u003c/sup\u003e.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Research on the seeds of the genus in species","content":"\u003cdiv class=\"Heading\"\u003e3. Research on the seeds of the genus \u003cem\u003eRubus\u003c/em\u003e in species\u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003e3.1. Collection of samples\u003c/h2\u003e \u003cp\u003eSeed specimens are the material basis for conducting seed morphology identification and classification research, as well as an important reference for identifying seeds excavated from archaeological sites. This study collected mature seeds of \u003cem\u003eRubus\u003c/em\u003e from local populations in different regions of China. A total of 126 seeds (including 22 varieties) of \u003cem\u003eRubus\u003c/em\u003e were collected. According to the section and subsection classification system of \u003cem\u003eRubus\u003c/em\u003e in Flora of China, the collected seeds were classified at the species level, after which the micromorphological variation of seeds across 8 sections and 23 subsections was evaluated. The information on the investigated species is listed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSeed source species and morphological determination of the genus \u003cem\u003eRubus\u003c/em\u003e L.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNo\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003elocation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. calycinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Chamaebatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. calycinoides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Chamaebatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTaiwan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pectinellus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Chamaebatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. chamaemorus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Chamaemorus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eInner Mongolia\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. rubrisetulosus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Cylactis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. saxatilis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Cylactis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHeilongjiang province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. nyalamensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Cylactis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eXizang autonomous region\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. fockeanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Cylactis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. tricolor\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Dalibardastrum\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. jinfoshanensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Dalibardastrum\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eChongqing city\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. multisetosus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Dalibardastrum\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. tsangorum\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Dalibardastrum\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJiangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lucens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Lampobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. caesius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Rubus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eXinjiang autonomous region\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pentagonus\u003c/em\u003e var. \u003cem\u003emodestus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Alpestres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. quinquefoliolatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Alpestres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pentagonus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Alpestres\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. corchorifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Corchorifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. chingii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Corchorifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. grayanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Corchorifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. glabricarpus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Corchorifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFujian province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. trianthus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Corchorifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHenan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. idaeus\u003c/em\u003e var. \u003cem\u003eglabratus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHeilongjiang province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. sachalinensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHeilongjiang province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. austrotibetanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eXizang autonomous region\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. eucalyptus\u003c/em\u003e var. \u003cem\u003etrullisatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. niveus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eXizang autonomous region\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. trijugus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eXizang autonomous region\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. eucalyptus\u003c/em\u003e var. \u003cem\u003eyunnanensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. idaeus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHebei province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. idaeus\u003c/em\u003e var. \u003cem\u003eborealisinensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHebei province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. piluliferus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. irritans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Idaeanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. leucanthus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Leucanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. delavayi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Leucanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. zhaogoshanensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Leucanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. columellaris\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Leucanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJiangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. impressinervius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Leucanthi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. peltatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Peltati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eZhejiang province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. komarovi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pileati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJilin province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pseudopileatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pileati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pileatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pileati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eShaanxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. macilentus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pinfaensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pungens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eShaanxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. stans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. amabilis\u003c/em\u003e var. \u003cem\u003eaculeatissimus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. coreanus\u003c/em\u003e var. \u003cem\u003etomentosus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. coreanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. amabilis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eShaanxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. inopertus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pungentes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. tsangii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. eustephanus\u003c/em\u003e var. \u003cem\u003eglanduliger\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. glandulosocalycinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTaiwan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. piptopetalus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTaiwan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. rosaefolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. hirsutus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. sumatranus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. glandulosocarpus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuizhou province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. eustephanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Rosaefolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. erythrocarpus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. kulinganus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJiangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. mesogaeus\u003c/em\u003e var. \u003cem\u003eoxycomus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. ellipticus\u003c/em\u003e var. \u003cem\u003eobcordatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. alexeterius\u003c/em\u003e var. \u003cem\u003eacaenocalyx\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. biflorus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province/ Hunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. mesogaeus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eShanxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. subornatus\u003c/em\u003e var. \u003cem\u003emelanadenus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. ellipticus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stimulantes\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. idaeopsis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuizhou province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. innominatus\u003c/em\u003e var. \u003cem\u003emacrosepalus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eZhejiang province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. innominatus\u003c/em\u003e var. \u003cem\u003earalioides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFujian province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. chiliadenus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. flosculosus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eShanxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. grandipaniculatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. cockburnianus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHenan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. adenophorus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFujian province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. innominatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Thyrsidaei\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lasiostylus\u003c/em\u003e var. \u003cem\u003ehubeiensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Wushanenses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHubei province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lasiostylus\u003c/em\u003e var. \u003cem\u003edizygos\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Wushanenses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lasiostylus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Idaeobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Wushanenses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGansu province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. poliophyllus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Acuminati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lambertianus\u003c/em\u003e var. \u003cem\u003epaykouangensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Acuminati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eChongqing city\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. laxus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Acuminati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lambertianus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Acuminati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. cochinchinensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Cochinchinenses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHainan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. angustibracteatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Dolichophylli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. dolichophyllus\u003c/em\u003e var. \u003cem\u003epubescens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Dolichophylli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuizhou province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. parkeri\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Dolichophylli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. howii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Dolichophylli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. ichangensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Dolichophylli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJiangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. feddei\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuizhou province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. salwinensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. luchunensis\u003c/em\u003e var. \u003cem\u003ecoriaceus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. fuscorubens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eChongqing city\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lobophyllus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. tephrodes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. paniculatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Elongati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. reflexus\u003c/em\u003e var. \u003cem\u003elanceolobus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Fuscifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e101\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. malipoensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Fuscifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e102\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. reflexus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Fuscifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e103\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. jambosoides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Jambosoides\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuangdong province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e104\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lineatus\u003c/em\u003e var. \u003cem\u003eglabrescens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Lineati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e105\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lineatus\u003c/em\u003e var. \u003cem\u003eangustifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Lineati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e106\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lineatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Lineati\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e107\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. wardii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Metoenses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e108\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. echinoides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. nagasawanus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTaiwan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e110\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. lasiotrichos\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. hunanensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e112\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. buergeri\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e113\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. setchuenensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHubei province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e114\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. faberi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. alceaefolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eJiangxi province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. multibracteatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e117\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. reticulatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Moluccani\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSichuan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e118\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pirifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Pirifolii\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFujian province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. doyonensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Sozostyli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e120\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. swinhoei\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Sozostyli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e121\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. preptanthus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Sozostyli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYunnan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e122\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. henryi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Sozostyli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eHunan province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e123\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. fujianensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Sozostyli\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eFujian province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e124\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. irenaeus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stipulosi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e125\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. pacificus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSect. Malachobatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSubsect. Stipulosi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAnhui province\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e126\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eR. fanjingshanensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGuizhou province\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 \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Identification of the seeds of the genus Rubus in China\u003c/h2\u003e \u003cp\u003eIn this study, the seed micromorphology of \u003cem\u003eRubus\u003c/em\u003e taxa native to China was first systematically documented. At the micromorphological level, certain morphological features of \u003cem\u003eRubus\u003c/em\u003e seeds (such as size, shape, color, etc.) may overlap across taxa, but the surface ornamentation of the seed coat exhibits significant variation among different species (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section3\"\u003e \u003ch2\u003e3.2.1. Seed size\u003c/h2\u003e \u003cp\u003eThe seed length ranges from 1.23 mm to 6.11 mm, with \u003cem\u003eR. chamaemorus\u003c/em\u003e having the longest seeds, while \u003cem\u003eR. howii\u003c/em\u003e has the shortest. Based on length, the seeds can be divided into four categories: Class A: Above 3.5 mm (e.g., \u003cem\u003eR. pirifolius\u003c/em\u003e, \u003cem\u003eR. quinquefoliolatus\u003c/em\u003e, \u003cem\u003eR. chamaemorus\u003c/em\u003e); Class B: 2.5 mm \u0026minus;\u0026thinsp;3.5 mm (e.g., \u003cem\u003eR. parkeri\u003c/em\u003e, \u003cem\u003eR. henryi\u003c/em\u003e, \u003cem\u003eR. flosculosus\u003c/em\u003e); Class C: 1.5 mm \u0026minus;\u0026thinsp;2.5 mm (e.g., \u003cem\u003eR. pacificus\u003c/em\u003e, \u003cem\u003eR. adenophorus\u003c/em\u003e, \u003cem\u003eR. piluliferus\u003c/em\u003e); Class D: Below 1.5 mm (e.g., \u003cem\u003eR. reticulatus\u003c/em\u003e, \u003cem\u003eR. sumatranus\u003c/em\u003e, \u003cem\u003eR. ellipticus\u003c/em\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section3\"\u003e \u003ch2\u003e3.2.2. Seed shape\u003c/h2\u003e \u003cp\u003eMost seeds exhibit an elliptical shape (e.g., \u003cem\u003eR. mesogaeus\u003c/em\u003e, \u003cem\u003eR. trijugus\u003c/em\u003e), semi-elliptical shape (e.g., \u003cem\u003eR. tephrodes\u003c/em\u003e, \u003cem\u003eR. idaeus\u003c/em\u003e), semi-circular shape (e.g., \u003cem\u003eR. lambertianus\u003c/em\u003e, \u003cem\u003eR. trianthus\u003c/em\u003e), ovate shape (e.g., \u003cem\u003eR. feddei\u003c/em\u003e, \u003cem\u003eR. parkeri\u003c/em\u003e), long-oval shape (e.g., \u003cem\u003eR. cinclidodictyus\u003c/em\u003e), or shield shape (e.g., \u003cem\u003eR. impressinervius\u003c/em\u003e, \u003cem\u003eR. glabricarpus\u003c/em\u003e). A few seeds show a square shape (e.g., \u003cem\u003eR. ichangensis\u003c/em\u003e), crescent shape (e.g., \u003cem\u003eR. henryi\u003c/em\u003e, \u003cem\u003eR. laxus\u003c/em\u003e), long-strip shape (e.g., \u003cem\u003eR. idaeus\u003c/em\u003e var. \u003cem\u003eborealisinensis\u003c/em\u003e, \u003cem\u003eR. multisetosus\u003c/em\u003e), or irregular shape (e.g., \u003cem\u003eR. retinulatus\u003c/em\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section3\"\u003e \u003ch2\u003e3.2.3. Surface ornamentation of seeds\u003c/h2\u003e \u003cp\u003eThe seed coat ornamentation of \u003cem\u003eRubus\u003c/em\u003e exhibits specific changes, which seem to have taxonomic significance. The seed coat ornamentation of seeds is reticular or striped, but different species have their own characteristics in terms of the depth of the holes, the composition of the reticular pattern, and the thickness, arrangement, and direction of the stripes. It mainly includes two categories: The first group is seeds with prominent reticulate sculpture (e.g., \u003cem\u003eR. reflexus\u003c/em\u003e, \u003cem\u003eR. parvifolius\u003c/em\u003e, \u003cem\u003eR. flosculosus\u003c/em\u003e). The second group is seeds with less prominent reticulate sculpture (e.g., \u003cem\u003eR. salwinensis\u003c/em\u003e, \u003cem\u003eR. cinclidodictyus\u003c/em\u003e, \u003cem\u003eR. fujianensis\u003c/em\u003e). Among them, the first group can be further divided into two types based on the depth of the holes: deep-hole type (e.g., \u003cem\u003eR. lobophyllus\u003c/em\u003e, \u003cem\u003eR. cochinchinensis\u003c/em\u003e) and shallow-hole type (e.g., \u003cem\u003eR. faberi\u003c/em\u003e, \u003cem\u003eR. lambertianus\u003c/em\u003e).\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e"},{"header":"4. Identification of Seeds from Archaeological Sites","content":"\u003cdiv class=\"Heading\"\u003e4. Identification of \u003cem\u003eRubus\u003c/em\u003e Seeds from Archaeological Sites\u003c/div\u003e \u003cp\u003eThe climate in China has undergone certain changes throughout history, which may have led to subtle differences in the distribution range of \u003cem\u003eRubus\u003c/em\u003e plants between ancient and modern times, thereby affecting the identification of seeds excavated from archaeological sites. Therefore, this study evaluated the seed characteristics of \u003cem\u003eRubus\u003c/em\u003e taxa distributed in different regions. The results indicated that the seeds selected for this study were representative, covering the morphological variation range of the genus across geographic populations.\u003c/p\u003e \u003cp\u003eConsidering that seed size is closely related to the plant's growth environment and may be affected by environmental factors during burial, causing expansion or shrinkage, in this study's identification work, seed morphology and seed coat ornamentation were mainly used as identification criteria, while seed size was only used as a reference. This study collected \u003cem\u003eRubus\u003c/em\u003e seed samples from seven archaeological sites (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePlant Remains Information of \u003cem\u003eRubus\u003c/em\u003e\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSerial number\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSite name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003csup\u003e14\u003c/sup\u003eC Dating\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePeriod\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eOrigin(Grains)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFruit period\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eWutongtu Site \u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eJiangjing Chongqing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1416\u0026thinsp;\u0026minus;\u0026thinsp;1235 cal BC (3190\u0026thinsp;\u0026minus;\u0026thinsp;3184 cal BP)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Shang and Zhou dynasties\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. coreanus\u003c/em\u003e var. \u003cem\u003etomentosus\u003c/em\u003e(1 grains)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eJun. to Aug.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNanshan Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eShaoxing, Zhejiang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Eastern Zhou Dynasty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. lambertianus\u003c/em\u003e(25 grains)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSep. to Nov.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTianshengcheng Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eWanzhou, Chongqing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Song Dynasty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. flosculosus\u003c/em\u003e(5 grains)/ \u003cem\u003eR. parvifolius\u003c/em\u003e༈17 grains༉\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAug. to Sep. / Jul. to Aug.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFanjiayan Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHechuan, Chongqing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1222\u0026ndash;1286 cal AD(728\u0026thinsp;\u0026minus;\u0026thinsp;664 cal BP)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Southern Song Dynasty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. trianthus\u003c/em\u003e(15 grains)/ \u003cem\u003eR. parvifolius\u003c/em\u003e༈15 grains༉\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMay to Jun. / Jul. to Aug.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTaipingmen Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYuzhong, Chongqing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1165\u0026ndash;1265 cal AD (785\u0026thinsp;\u0026minus;\u0026thinsp;685 cal BP)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Southern Song Dynasty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. adenophorus\u003c/em\u003e(6 grains)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eJun. to Jul.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDrum Tower Yamen Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYuzhong, Chongqing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1221\u0026ndash;1282 cal AD (729\u0026thinsp;\u0026minus;\u0026thinsp;668 cal BP)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ethe Southern Song Dynasty\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. trianthus\u003c/em\u003e(20 grains)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMay to Jun.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eYuanmingyuan site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHaidian, Beijing\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eThe Ming and Qing Dynasties / the Modern Times\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eR. lambertianus\u003c/em\u003e(1 grains /1 grains)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSep. to Nov.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e4.1. Wutongtu Site\u003c/h2\u003e \u003cp\u003eThe Wutongtu Site is located on a crescent-shaped platform on the left bank of the Yangtze River in Youxi Town, Jiangjin District, Chongqing City. In this study, one \u003cem\u003eRubus\u003c/em\u003e seed was selected from an ash pit dating to the Shang and Zhou dynasties.\u003c/p\u003e \u003cp\u003eAlthough the seed has been carbonized, the seed coat ornamentation remains clearly visible under microscopy. Observations show that it is elliptical in shape (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ej), with reticular intersecting ornamentation. Based on these characteristics and comparison with modern specimens, it is identified as \u003cem\u003eR. coreanus\u003c/em\u003e var. \u003cem\u003etomentosus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003e4.2. Nanshan Site\u003c/h2\u003e \u003cp\u003eThe Nanshan Site is located in Yuecheng District, Shaoxing City, Zhejiang Province. In this study, 25 \u003cem\u003eRubus\u003c/em\u003e seeds were obtained from the site dating to the Eastern Zhou Dynasty.\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eRubus\u003c/em\u003e seeds are uncarbonized, with distinct seed coat ornamentation. They are half-moon shaped (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eg), with bluntly rounded ends, and the seed coat ornamentation branches from the hilum in a tree-branch pattern. Based on these characteristics and correspondence with morphological descriptions of \u003cem\u003eR. lambertianus\u003c/em\u003e in taxonomic literature, the seeds excavated from the Nanshan Site are inferred to originate from \u003cem\u003eR. lambertianus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003e4.3. Tianshengcheng Site\u003c/h2\u003e \u003cp\u003eThe Tianshengcheng Site is located in Zhoujiaba Street, Wanzhou District, Chongqing City, adjacent to the confluence of the Zhuxi River and the Yangtze River to the south. It is an important Song and Yuan dynasties mountain city site. In this study, 22 \u003cem\u003eRubus\u003c/em\u003e seeds were selected from Southern Song Dynasty water wells.\u003c/p\u003e \u003cp\u003eThe seeds are uncarbonized. Based on morphological characteristics, they can be divided into two types: semicircular (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec) and oval (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ed).\u003c/p\u003e \u003cp\u003eSemi-circular type: Seeds are semi-circular, with bluntly rounded ends and a small aspect ratio. The seed coat ornamentation consists of intersecting polygonal meshes with relatively dense reticulation. Identification confirms they belong to \u003cem\u003eR. flosculosus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eOval type: Seeds are oval, with one end sharp and the other bluntly rounded. The seed coat exhibits polygonal intersecting patterns with relatively loose overall reticulation. Identification confirms they belong to \u003cem\u003eR. parvifolius\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003e4.4. Fanjiayan Site\u003c/h2\u003e \u003cp\u003eThe Fanjiayan Site is located on Diaoyu Mountain in the northeastern part of Dongcheng Peninsula, Hechuan District, Chongqing City. As a significant component of the Fishing Town Site, it is a well-preserved relic of governmental establishments from the Song and Yuan Dynasties. In this study, 30 \u003cem\u003eRubus\u003c/em\u003e seeds were selected from a Southern Song Dynasty pool (archaeologically identified as a water storage facility).\u003c/p\u003e \u003cp\u003eNotably, the excavated \u003cem\u003eRubus\u003c/em\u003e seeds are uncarbonized and exhibit diverse shapes. Based on morphology, they can be categorized into two types: oval (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea) and semi-oval (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003eOval type: Seeds are oval, with one end pointed and the other bluntly rounded. The seed coat exhibits intersecting polygonal patterns with relatively sparse reticulation, consistent with the seed characteristics of \u003cem\u003eR. parvifolius\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eSemi-oval type: Seeds are semi-oval, with one end slightly protruding and the other bluntly rounded, showing a large aspect ratio. The seed coat ornamentation features branching patterns originating from the hilum, with overall sparse reticulation, matching the seed characteristics of \u003cem\u003eR. trianthus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003e4.5. Taipingmen Site\u003c/h2\u003e \u003cp\u003eThe Taipingmen Site is located on Wanglongmen Street, Yuzhong District, Chongqing City. Situated on the first river terrace of the left bank of the Yangtze River in the Yuzhong Peninsula, it is one of the important city gates of Chongqing. In this study, six \u003cem\u003eRubus\u003c/em\u003e seeds were selected from Southern Song Dynasty drainage ditches of the city walls.\u003c/p\u003e \u003cp\u003eThe seeds are uncarbonized, semi-oval with slightly blunt ends. The seed coat ornamentation is indistinct, faintly appearing as an irregular cross pattern (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ef). Based on these characteristics and comparison with R. adenophorus specimens in collections, the excavated \u003cem\u003eRubus\u003c/em\u003e seeds are inferred to derive from R. adenophorus (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003e4.6. Drum Tower Yashu Site\u003c/h2\u003e \u003cp\u003eThe Drum Tower Yashu Site is located in Wanglongmen Street, Yuzhong District, Chongqing City. Situated on a terrace at the foot of Jinbi Mountain and the left bank of the Yangtze River, it served as the administrative center of Sichuan and Chongqing Prefecture during the Southern Song Dynasty. In this study, 20 \u003cem\u003eRubus\u003c/em\u003e seeds were selected from Southern Song Dynasty ponds.\u003c/p\u003e \u003cp\u003eThe seeds are uncarbonized, semi-oval with one end slightly protruding and the other blunt and round, and have a large aspect ratio. The seed coat ornamentation features a network-like, cross-shaped pattern with relatively loose overall reticulation. Based on these characteristics and comparison with specimens of R. trianthus, they are identified as originating from R. trianthus (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003e4.7. Yuanmingyuan Site\u003c/h2\u003e \u003cp\u003eThe Yuanmingyuan Site is located in Haidian District, Beijing City. One \u003cem\u003eRubus\u003c/em\u003e seed was excavated from each of the Ming and Qing Dynastic relics (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eh) and modern deposits (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ei).\u003c/p\u003e \u003cp\u003eThe seeds are uncarbonized, all crescent-shaped with bluntly rounded ends. The seed coat ornamentation branches from the hilum in a tree-branch pattern, with ornamentation lines approximately parallel. Based on morphological consistency across historical and modern specimens, the \u003cem\u003eRubus\u003c/em\u003e seeds excavated from both periods at this site are identified as \u003cem\u003eR. lambertianus\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"5. Discussion","content":"\u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003e5.1. Application of This Identification Methodology to Rubus Seed Remains from Other Archaeological Sites in China\u003c/h2\u003e \u003cp\u003eThis study preliminarily demonstrates that the established identification method is effective for identifying seeds of the genus \u003cem\u003eRubus\u003c/em\u003e excavated from archaeological contexts, based on the collection and analysis of a large dataset of modern Rubus seed samples. However, the archaeological \u003cem\u003eRubus\u003c/em\u003e seed assemblages discussed above are predominantly from Chongqing, Zhejiang, and Beijing. To further validate the universality of this methodology, we applied it to the re-identification of Rubus seed remains excavated from additional archaeological sites across China, spanning diverse geographical regions and chronological periods (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eRe-identification of \u003cem\u003eRubus\u003c/em\u003e seed remains in reported archaeological sites.\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=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNo\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eGrains\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ere-identification Result\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ereferences\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBashidang Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChangde, Hunan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eYuchisi Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBozhou, Anhui\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e11,13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLiangzhu Ancient City Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eHangzhou, Zhejiang\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChengtoushan Site\u003csup\u003e\u0026amp;\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eChangde, Hunan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eGuangfulin Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eShanghai\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. hirsutus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDaxinzhuang Site\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eJinan, Shandong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. parvifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChangdun Han Tomb Cemetery\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNanchang, Jiangxi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. rosifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNanyue Palace Site: J264 Water Well\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGuangzhou, Guangdong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. rosifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e20,13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNanyue Palace Garden: Nanhan Kingdom Palace Pool Site\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eGuangzhou, Guangdong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eR. rosifolius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e12,13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003e*Carbonized; \u003csup\u003e\u0026amp;\u003c/sup\u003eIdentification error\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThrough identification, \u003cem\u003eRubus\u003c/em\u003e seeds excavated from the Bashidang Site, Yuchisi Site, Liangzhu Ancient City Site, Chengtoushan Site, and Daxinzhuang Site all exhibit a pointed-obtuse shape. The seed coat ornamentation consists of polygonal intersections with an overall loose pattern. These seeds are uniformly identified as \u003cem\u003eR. parvifolius\u003c/em\u003e. Notably, the previously reported identification of \u003cem\u003eRubus\u003c/em\u003e seeds from the Chengtoushan Site as \u003cem\u003eR. lambertianus\u003c/em\u003e is revised herein. This study clarifies that \u003cem\u003eR. lambertianus\u003c/em\u003e seeds are distinctively characterized by bluntly rounded ends and forked, hilum-branching ornamentation, whereas the Chengtoushan seeds display reticulate interwoven ornamentation, consistent with \u003cem\u003eR. parvifolius\u003c/em\u003e rather than \u003cem\u003eR. lambertianus\u003c/em\u003e. Therefore, this study posits that the provenance of the \u003cem\u003eRubus\u003c/em\u003e seeds excavated from the Chengtoushan Site should be \u003cem\u003eR. parvifolius\u003c/em\u003e rather than \u003cem\u003eR. lambertianus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eRubus\u003c/em\u003e seeds excavated from the Guangfulin Site exhibit a semicircular shape with seed coat ornamentation featuring reticulate intersections in a hexagonal pattern, consistent with the seed morphology of \u003cem\u003eR. hirsutus\u003c/em\u003e. Based on these diagnostic traits\u0026mdash;specifically the semicircular outline and dense hexagonal reticulation distinct from other sympatric species (e.g., \u003cem\u003eR. parvifolius\u003c/em\u003e with looser polygonal patterns)\u0026mdash;these seeds are taxonomically identified as \u003cem\u003eR. hirsutus\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eRubus\u003c/em\u003e seeds excavated from the Changdun Han Tomb Cemetery, Nanyue Palace Site (Well J264), and Nanyue Palace Garden (Palace Pool Site of the Southern Han Kingdom) exhibit a relatively high length-to-width ratio with bluntly rounded ends. Their seed coat ornamentation features densely interwoven reticulate patterns, diagnostic of \u003cem\u003eR. rosifolius\u003c/em\u003e. These morphological traits\u0026mdash;specifically the elongated oval shape and anastomosing reticulation distinct from other congeners (e.g., \u003cem\u003eR. lambertianus\u003c/em\u003e with bifurcating hilum patterns)\u0026mdash;consistently align with the seed characteristics of \u003cem\u003eR. rosifolius\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThe above identification results fully demonstrate that the identification method established in this study can be effectively applied to the identification of \u003cem\u003eRubus\u003c/em\u003e seed remains in Chinese archaeological sites (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003e5.2 Established an illustrated guide to the seeds of common Rubus plants in China\u003c/h2\u003e \u003cp\u003eStereoscopic microscopy and scanning electron microscopy (SEM) have revealed many important seed characteristics, which contribute to resolving taxonomic problems in botanical origin identification\u003csup\u003e\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e,\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e\u003c/sup\u003e. This article is the first to use a stereomicroscope to study the microscopic morphological characteristics of seeds in Chinese plants of the genus \u003cem\u003eRubus\u003c/em\u003e and systematically describes the microscopic morphological characteristics of many taxonomic groups. In recent years, the use of stereomicroscopes as a means of studying microstructural changes in plants has gained advantages in plant systems\u003csup\u003e\u003cspan additionalcitationids=\"CR44\" citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e, with observed data achieving high precision and being irreplaceable by other techniques. At different classification levels, there are significant differences in the seed micromorphology of the genus \u003cem\u003eRubus\u003c/em\u003e in terms of different traits such as seed size, shape, and seed coat ornamentation. The micromorphological characteristics of seeds are key features for plant species identification and classification, reflecting the genetic, physiological, and ecological characteristics of plants.\u003c/p\u003e \u003cp\u003eThis paper uses stereomicroscopy to study the micromorphological characteristics of the seeds of \u003cem\u003eRubus\u003c/em\u003e plants in China and systematically describes the micromorphological characteristics of many taxonomic groups for the first time. In recent years, the use of stereomicroscopy as a means to study plant micromorphological changes has gained an advantage in plant systems\u003csup\u003e\u003cspan additionalcitationids=\"CR44\" citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e\u003c/sup\u003e. Stereomicroscopic observations yield highly precise data and cannot be replaced by other technologies. At different taxonomic levels, there are significant differences in the seed micromorphology of the genus \u003cem\u003eRubus\u003c/em\u003e in different traits (such as seed size, shape, and seed coat ornamentation). Seed micromorphological characteristics are key features for plant species identification and classification, reflecting the genetic, physiological, and ecological characteristics of plants.\u003c/p\u003e \u003cp\u003eThe micromorphological characteristics of seeds can provide useful information for the species classification of this genus. For example, although \u003cem\u003eR. ichangensis\u003c/em\u003e and \u003cem\u003eR. angustibraceteatus\u003c/em\u003e are very similar in plant morphological characteristics, their seed morphological features are completely different. However, many species assigned to different sections or subsections with distantly related macromorphological relationships exhibit similar seed morphological characteristics, such as \u003cem\u003eR. multibraceteatus\u003c/em\u003e and \u003cem\u003eR. mesogeaus\u003c/em\u003e, \u003cem\u003eR. fuscorubens\u003c/em\u003e and \u003cem\u003eR. parkeri\u003c/em\u003e. In other words, apart from the seed coat ornamentation, no section (except Sect. Ghamaemorus) or subsection (except Subsect. Pirifolii) exhibits unique seed morphology, and species of different subgenera or families have similar seed morphology (such as size, shape, color, etc.). Whether seed coat ornamentation can be used for intrageneric classification depends on the degree of variation between species\u003csup\u003e\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e\u003c/sup\u003e. Although this study observed the seed morphology of common plants in the Chinese \u003cem\u003eRubus\u003c/em\u003e genus, which covers almost all sections and subsections of the genus, it remains difficult to identify the evolutionary patterns of seed size, shape, and seed coat ornamentation, as well as their correlations with macromorphological characteristics.\u003c/p\u003e \u003cp\u003eThe accurate identification of plant seed remains requires the support of a large number of modern seed specimens. Based on this, this study constructed the first seed morphology atlas of the genus \u003cem\u003eRubus\u003c/em\u003e in China according to key characteristics such as seed shape and seed coat ornamentation and applied it to the identification of seed remains in archaeological sites. Through the method of comparing ancient and modern specimens, this study provides an important reference for the identification of \u003cem\u003eRubus\u003c/em\u003e seeds unearthed from sites.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003e5.3 The Uses of Ancient Rubus Plants\u003c/h2\u003e \u003cp\u003eIn this study, some \u003cem\u003eRubus\u003c/em\u003e seed remains were unearthed from water pool remains at archaeological sites\u003csup\u003e\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e,\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u003c/sup\u003e. Meanwhile, in some sites, \u003cem\u003eRubus\u003c/em\u003e seeds were unearthed with abundant crop and fruit remains. Based on these comprehensive archaeological findings, it is inferred that the fruits or seeds of \u003cem\u003eRubus\u003c/em\u003e at the sites were intentionally collected and preserved\u003csup\u003e\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e,\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e\u003c/sup\u003e. Additionally, identification shows that the fruits of the original plants corresponding to \u003cem\u003eRubus\u003c/em\u003e seeds from archaeological sites are all red (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), with a sweet and sour taste. Based on this, it can be inferred that the fruit of the original seed plant of the genus \u003cem\u003eRubus\u003c/em\u003e is highly likely to have been an important component of the ancient ancestors' plant-based diet.\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eRubus\u003c/em\u003e genus is a widely distributed plant and grows in mountainous areas, hills, and wasteland hillsides in most parts of southern and central China. The fruiting period of \u003cem\u003eRubus\u003c/em\u003e plants varies from May to November, making them common wild fruits. Its fruits are plump and juicy and taste delicious. It is highly probable that ancient people would pick and eat ripe \u003cem\u003eRubus\u003c/em\u003e fruits during their outdoor activities. Additionally, from the perspective of traditional medicine, some fruits of the \u003cem\u003eRubus\u003c/em\u003e genus have medicinal values and are rich in nutrients\u003csup\u003e\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e,\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eNumerous studies have shown that \u003cem\u003eRubus\u003c/em\u003e seed remains are prevalent in Chinese archaeological sites\u003csup\u003e\u003cspan additionalcitationids=\"CR13 CR14 CR15 CR16 CR17 CR18 CR19 CR20 CR21\" citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u003c/sup\u003e. In some sites, a large number of seeds were unearthed; this is possibly because each \u003cem\u003eRubus\u003c/em\u003e fruit contains numerous seeds. This quantitative advantage naturally increases the probability of \u003cem\u003eRubus\u003c/em\u003e seeds being discarded in archaeological deposits, resulting in their general dominance in absolute numbers among unearthed seeds\u003csup\u003e\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e,\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eIn addition, the seeds of plants in the genus \u003cem\u003eRubus\u003c/em\u003e are relatively small in size. When consuming the fruits, ancient people generally did not deliberately spit out the seeds, which were excreted through bodily waste. In natural environments, these seeds might be wrapped in other excrement; this process reduces erosion from the external environment and thus enables relatively complete preservation in archaeological sites. It is worth noting that plants of the genus \u003cem\u003eRubus\u003c/em\u003e also have the possibility of being cultivated as ornamental plants\u003csup\u003e\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e\u003c/sup\u003e. The abundant and well-preserved \u003cem\u003eRubus\u003c/em\u003e seed remains in some sites may be linked to special burial contexts or specific functions. Nevertheless, clarifying their specific roles demands more archaeological evidence and targeted research.\u003c/p\u003e \u003c/div\u003e"},{"header":"6. Conclusions","content":"\u003cp\u003eIn this study, 126 mature seeds from wild \u003cem\u003eRubus\u003c/em\u003e populations across various regions of China were collected for systematic seed morphology analysis, leading to the construction of the first morphological atlas of seeds for common Chinese \u003cem\u003eRubus\u003c/em\u003e species. Through the analysis of \u003cem\u003eRubus\u003c/em\u003e seed remains from 7 unreported sites and 9 reported sites, the taxonomic identity of seeds was accurately identified.\u003c/p\u003e \u003cp\u003eThis study employed modern plant seed morphology research methods to link modern samples to archaeological samples, thereby providing new ideas and methods for plant archaeology research. As seed identification of \u003cem\u003eRubus\u003c/em\u003e from more sites progresses and research deepens, this study is expected to further reveal the edible history and species of \u003cem\u003eRubus\u003c/em\u003e fruits, provide important historical background information for exploring the distribution shifts and utilization patterns of \u003cem\u003eRubus\u003c/em\u003e plants, and offer strong archaeological evidence for studying the biodiversity evolution of \u003cem\u003eRubus\u003c/em\u003e.\u003c/p\u003e"},{"header":"Declarations","content":" \u003ch2\u003eConflicts of Interest:\u003c/h2\u003e \u003cp\u003eThe authors declare no conflicts of interest.\u003c/p\u003e \u003ch2\u003eFundings:\u003c/h2\u003e \u003cp\u003eThe National Key Research and Development Program of China (2022YFC3500900、2022YFC3500902), Key project at central government level: The ability establishment of sustainable use for valuable Chinese medicine resources(2060302), China Academy of Chinese Medical Sciences: Science and Technology Innovation Project(CI2024E001), The Major Increase and Reduction 516 Project at the Central Level (2060302) and the Innovation Team and Talents Cultivation Program 517 of National Administration of Traditional Chinese Medicine (ZYYCXTD-C-202409).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eA. R.: Writing-original draft, Writing-review and editing, Investigation; X. M.: Writing-original draft, Writing-review and editing, Investigation;; X. X.: Resources, Data curation; M. Y.: Writing-original draft, Writing-review and editing, Methodology, Investigation, Funding acquisition; H. J.: Writing-review and editing, Conceptualization, Project administration, Funding acquisition. All authors have read and agreed to the published version of the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe would like to express our gratitude to Professor Peng Huasheng for his constructive comments.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData are contained within the article. Further information can be obtained by contacting the corresponding author.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eLiu C, Jin G, Kong Z. Archaeobotany: Research on Seeds and Fruits. Beijing: Sci. 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Pr. 194\u0026ndash;243 (2003).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWaugh R, et al. Chloroplast DNA diversity in the genus \u003cem\u003eRubus\u003c/em\u003e (Rosaceae) revealed by Southern hybridization. Plant Syst Evol. 1990;172:65\u0026ndash;75.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAlice L, Campbell C. Phylogeny of \u003cem\u003eRubus\u003c/em\u003e (Rosaceae) based on nuclear ribosomal DNA internal transcribed spacer region sequences. Am J Bot. 1999;86:81\u0026ndash;97.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKurtto A, Weber H, Lampinen R. Atlas florae Europaeae: distribution of vascular plants in Europe: Rosaceae (\u003cem\u003eRubus\u003c/em\u003e). Com. Map Flora Europe \u0026amp; Societas Biol. Fennica Vanamo. (2010).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXiong X, et al. Pollen morphology in \u003cem\u003eRubus\u003c/em\u003e (Rosaceae) and its taxonomic implications. Plant Syst Evol. 2019;305:705\u0026ndash;16.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGovaerts R, et al. The World Checklist of Vascular Plants, a continuously updated resource for exploring global plant diversity. Sci Data. 2021;8:215.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYu D, Lu L, Gu C. Flora of China (37). Beijing: Sci Pr 10\u0026ndash;218 (1985).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKollmann J, Steinger T, Roy B. Evidence of sexuality in European \u003cem\u003eRubus\u003c/em\u003e (Rosaceae) species based on AFLP and allozyme analysis. Am J Bot. 2000;87:1592\u0026ndash;8.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeng Q, Manghwar H, Hu W. Study on supergenus \u003cem\u003eRubus\u003c/em\u003e L.: Edible, medicinal, and phylogenetic characterization. Plants. 2022;11:1211.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eŠarhanov\u0026aacute; P, et al. 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Seed morphology and anatomy of \u003cem\u003eRubus\u003c/em\u003e geoides Sm. Chi J Agric Res. 2016;76:385\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSurya M, Destri D, Ismaini L. Seed variation of \u003cem\u003eRubus\u003c/em\u003e spp. in Cibodas Botanical Garden. In Journal of Physics: Conference Series (Vol. 1943, No. 1, p. 012075). IOP Publishing. (2021).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChernoff M, Plitmann U, Kislev M. Seed characters and testa texture in species of Vicieae: Their taxonomic significance. Isr J Bot. 1992;41:167\u0026ndash;86.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLersten N, Gunn C. Testa characters in tribe Vicieae, with notes about tribes Abreae, Cicereae, and Trifolieae (Fabaceae) (1982).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRashid N, et al. Intraspecific variation in seed morphology of tribe vicieae (Papilionoidae) using scanning electron microscopy techniques. Microsc Res Tech. 2018;81:298\u0026ndash;307.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eUllah F, et al. Seed micromorphology and its taxonomic evidence in subfamily Alsinoideae (Caryophyllaceae). Microsc Res Tech. 2019;82:250\u0026ndash;9.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGanie A, et al. Fruit morphology of the genus Potamogeton L. in Kashmir Himalaya and its utility in taxonomic delimitation. J Asia Pac Biodivers. 2017;10:274\u0026ndash;8.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eShah H, et al. First report of ultra-sculpturing and taxonomic implications of seed morphology of selected genus \u003cem\u003eEuphorbia\u003c/em\u003e L. Genetic Resour Crop Evol. 2025;72:3357\u0026ndash;81.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eUllah A, et al. Ultra-sculpturing of nutlet morphology and its taxonomic significance for identifications of Boraginaceae species. Genet Resour Crop Evol. 2025;72:2743\u0026ndash;66.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003e\u0026Ccedil;ilden E, Yıldırımlı Ş. The impact of seed micromorphology in the subgeneric classification of the genus \u003cem\u003eReseda\u003c/em\u003e L. (Resedaceae) in Turkey. Microsc Res Tech. 2021;84:1992\u0026ndash;2003.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBuczyński K, Kapłan M, Jarosz Z. Review of the report on the nutritional and health-promoting values of species of the \u003cem\u003eRubus\u003c/em\u003e L. genus. Agr. 2024;14:1324.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMoreno-Medina B, Casierra-Posada F, Cutler J. Phytochemical Composition and Potential Use of \u003cem\u003eRubus\u003c/em\u003e Species. Gesunde Pflanz. 2018;70:65\u0026ndash;74.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"npj-heritage-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"hsci","sideBox":"Learn more about [Heritage Science](http://heritagesciencejournal.springeropen.com)","snPcode":"40494","submissionUrl":"https://submission.nature.com/new-submission/40494/3","title":"npj Heritage Science","twitterHandle":"@SpringerOpen","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Archaeobotany, Rubus, Seed Morphological Atlas, Identification","lastPublishedDoi":"10.21203/rs.3.rs-8367387/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8367387/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eRubus\u003c/em\u003e seeds constitute the predominant plant remains in archaeobotanical assemblages from Chinese archaeological sites. Nevertheless, no studies have yet reported the precise species-level identification of archaeological \u003cem\u003eRubus\u003c/em\u003e remains, as a result of the lack of systematic research on \u003cem\u003eRubus\u003c/em\u003e seed morphology. The first seed morphology atlas was established by systematically collecting and analyzing the morphological characteristics of 126 \u003cem\u003eRubus\u003c/em\u003e plants' mature seeds, which represent \u003cem\u003eRubus\u003c/em\u003e taxa in China (including 22 varieties). Utilizing this atlas, we identified \u003cem\u003eRubus\u003c/em\u003e seeds from seven archaeological sites, which ranged from the Shang-Zhou dynasties to the present day. Of these seeds, one was carbonized and six were uncarbonized. These specimens were identified as \u003cem\u003eRubus lambertianus\u003c/em\u003e, \u003cem\u003eR. parvifolius\u003c/em\u003e, \u003cem\u003eR. flosculosus\u003c/em\u003e, \u003cem\u003eR. coreanus\u003c/em\u003e var. \u003cem\u003etomentosus\u003c/em\u003e, \u003cem\u003eR. trianthus\u003c/em\u003e, and \u003cem\u003eR. adenophorus\u003c/em\u003e. Furthermore, the atlas was employed to precisely identify \u003cem\u003eRubus\u003c/em\u003e seeds from nine archaeological sites that had been previously reported, primarily assigning them to \u003cem\u003eR. parvifolius\u003c/em\u003e, \u003cem\u003eR. hirsutus\u003c/em\u003e, and \u003cem\u003eR. rosifolius\u003c/em\u003e. The results indicate that the \u003cem\u003eRubus\u003c/em\u003e seed morphology atlas developed in this investigation can serve as a critical reference for the identification of \u003cem\u003eRubus\u003c/em\u003e seeds that have been excavated from archaeological sites in China.\u003c/p\u003e","manuscriptTitle":"Identification of Rubus Seeds in Chinese Archaeological Sites: Based on the Morphological Atlas of 126 Modern Rubus Seeds","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-25 14:04:12","doi":"10.21203/rs.3.rs-8367387/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-01-26T03:27:31+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-21T09:49:52+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-15T18:54:03+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-13T04:21:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"161310834624494131068464656925778836942","date":"2025-12-25T20:45:11+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"256512999398589246578268028161684479735","date":"2025-12-24T12:40:58+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"72777250579412016083654272717675088411","date":"2025-12-24T02:37:13+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"168278741114890648195027218581079557794","date":"2025-12-23T23:50:02+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-12-23T23:26:09+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-12-22T04:45:50+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-12-22T04:44:58+00:00","index":"","fulltext":""},{"type":"submitted","content":"npj heritage science","date":"2025-12-15T14:27:00+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":"b9650658-1fdd-442f-8305-68574eb3848a","owner":[],"postedDate":"December 25th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-15T20:08:25+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-25 14:04:12","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8367387","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8367387","identity":"rs-8367387","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}