山茶属植物系统发育及分类研究进展

doi:10.16420/j.issn.0513-353x.2023-0192

摘要/Abstract

摘要：

基于形态学特征，目前山茶属植物分类主要有Sealy、张宏达和闵天禄所创立的3种系统。由于所关注的表型特征不同，这3种分类观点存在较大争议。本文从国内外山茶属分类研究历史出发，对山茶属分类和系统发育研究进展进行综述，主要对基于不同的标记类型对山茶属植物系统发育重建的相关研究进行整理总结，评价不同标记在山茶属物种系统发育分析中的利弊。提出目前分类中的问题和今后的重点研究方向，以期为重建山茶属物种间系统关系和解决分类争议提供一定的参考。

关键词: 山茶属, 分类学, 系统发育, 研究进展

Abstract:

The genus Camellia comprises a vast number of species that are widely distributed across various regions. The frequent inter-specific hybridization results in，controversial and unstable taxonomy and systematics among different species，which also poses a greater challenge for infrageneric classification within Camellia. Based on morphological features，Sealy，Chang Hongda，and Min Tianlu have established three classification systems for species in the genus. However they hold different opinions on the taxonomic significance of each character. This paper presents a comprehensive overview of the classification and phylogenetic research progress on the Camellia genus from a global perspectives. Pertinent research on phylogeny reconstruction of Camellia genus based on various marker types were summarized，appraise the merits and demerits of these markers in elucidating phylogenetic relationships，and propose future research directions to address current classification issues. The paper provides valuable insights into reconstructing the systematic relationship within the Camellia genus and resolving classification disputes.

Key words: Camellia, taxonomy, phylogeny, research progress

邝雪琨, 郑倩, 孔庆博, 周佳思, 周莉君, 冯士令, 丁春邦, 陈涛. 山茶属植物系统发育及分类研究进展[J]. 园艺学报, 2024, 51(1): 53-66.

KUANG Xuekun, ZHENG Qian, KONG Qingbo, ZHOU Jiasi, ZHOU Lijun, FENG Shiling, DING Chunbang, CHEN Tao. Research Progress on Phylogeny and Classification of Camellia[J]. Acta Horticulturae Sinica, 2024, 51(1): 53-66.

图/表 4

参考文献 64

[1]	Ao C Q, Chen G X, Zhang H D. 2002. Leaf epidermis morphology of Camellia and its taxonomic significance. Plant Diversity, 24 (1):1.
[2]	Ao Cheng-qi. 2003. Floral organogenesis of Camellia Japonica and C.ptilophylla. Journal of Wenzhou Teachers College, 24 (2):12-15. (in Chinese)
	敖成齐. 2003. 山茶和毛叶茶的花器官形态发生. 温州师范学院学报, 24 (2):12-15.
[3]	Chang Hung-ta, Ren Shan-xiang. 1998. Flora of China. Volume 49,Part 3. Beijing: Science Press. (in Chinese)
	张宏达, 任善湘. 1998. 中国植物志. 第49卷,第3分册. 北京: 科学出版社.
[4]	Chen Chun-mei, Ma Chun-lei, Ma Jian-qiang, Liu Sheng-chuan, Chen Liang. 2014. Sequencing of chloroplast genome of Camellia sinensis and genetic relationship for Camellia plants based on chloroplast DNA sequences. Journal of Tea Science, 34 (4):371-380. (in Chinese)
	陈春梅, 马春雷, 马建强, 刘声传, 陈亮. 2014. 茶树cpDNA测序及基于cpDNA序列的山茶属植物亲缘关系研究. 茶叶科学, 34 (4):371-380.
[5]	Chen Chun-ru, Xie Pei-wu, Zou Jia-tong, Lian Hui-ming, Wang Ying-li, Qiu Wen-xia, He Bo-xiang, Cai Yan-ling, Liang Dong-cheng, Chen Yi-qun, Chen Jie-lian. 2019. Comparative leaf morphology study of 12 species of Sect. Chrysantha (Theaceae,Camellia). Forestry and Environmental Science, 35:7-15. (in Chinese)
	陈春如, 谢佩吾, 邹家通, 连辉明, 汪迎利, 邱文霞, 何波祥, 蔡燕灵, 梁东成, 陈一群, 陈杰连. 2019. 12种金花茶组植物叶片的比较形态学研究. 林业与环境科学, 35:7-15.
[6]	Chen Shi-lin, Pang Xiao-hui, Luo Kun, Yao Hui, Han Jian-ping, Song Jing-Yuan. 2013. DNA barcoding of biological resources. Chinese Bulletin of Life Sciences, 25 (5):451-459. (in Chinese)
	陈士林, 庞晓慧, 罗焜, 姚辉, 韩建萍, 宋经元. 2013. 生物资源的DNA条形码技术. 生命科学, 25 (5):451-459.
[7]	Committee F O C E. 2018. Flora of China. Massachusetts,USA:Missouri Botanical Garden and Harvard University Herbaria.
[8]	Cui X, Li C, Qin S, Huang Z, Gan B, Jiang Z, Huang X, Yang X, Li Q, Xiang X. 2022. High-throughput sequencing-based microsatellite genotyping for polyploids to resolve allele dosage uncertainty and improve analyses of genetic diversity,structure and differentiation:a case study of the hexaploid Camellia oleifera. Molecular Ecology Resources, 22 (1):199-211. doi: 10.1111/men.v22.1 URL
[9]	Deng Bai-luo, Tan Xiao-feng, Qi Long-lin, He Jin, Hu Fang-ming. 2006. RAPD analysis and taxonomy of Sect. Camellia Species in Camellia. Scientia Silvae Sinicae, 42 (5):36-41. (in Chinese)
	邓白罗, 谭晓风, 漆龙霖, 贺晶, 胡芳名. 2006. 山茶属红山茶组植物的RAPD分析及分类研究. 林业科学, 42 (5):36-41.
[10]	Deng Yu-jun, Li Yi, He Ping, Deng Hong-ping, Li Wen-qiao, Yang Xu-dong. 2018. Study on leaf epidermal micro-morphology of 20 species of Theaceae. Acta Botanica Boreali-Occidentalia Sinica,(8):1476-1486. (in Chinese)
	邓禹君, 李懿, 贺萍, 邓洪平, 李文巧, 杨旭东. 2018. 20种山茶科植物的叶表皮微形态研究. 西北植物学报,(8):1476-1486.
[11]	Fang Wei, Yang Jun-bo, Yang Shi-xiong, Li De-zhu. 2010. Phylogeny of Camellia sects. Longipedicellata,Chrysantha and Longissima(Theaceae)based on sequence data of four chloroplast DNA loci. Acta Botanica Yunnanica, 32 (1):1-13. (in Chinese) doi: 10.3724/SP.J.1143.2010.00001 URL
	方伟, 杨俊波, 杨世雄, 李德铢. 2010. 基于叶绿体四个DNA片段联合分析探讨山茶属长柄山茶组、金花茶组和超长柄茶组的系统位置与亲缘关系. 云南植物研究, 32 (1):1-13.
[12]	Gao Yuan, Xie Yuan, Yang Zi-yun, Chen Long-qing, Wu Tian. 2022. Genetic relationship analysis of different Camellia sasanqua cultivars based on the difference of flowering season and ISSR-PCR. Non-wood Forest Research, 40 (4):209-217. (in Chinese)
	高园, 解元, 杨自云, 陈龙清, 吴田. 2022. 基于花期差异和ISSR-PCR分析的不同茶梅品种间亲缘关系. 经济林研究, 40 (4):209-217.
[13]	Jiang Zheng-dong. 2017. Preliminary study of molecular phylogenetics and biogeography of the genus Camellia L. based on chloroplast DNA[M. D. Dissertation ]. Hangzhou: Zhejiang Sci-tech University. (in Chinese)
	江正栋. 2017. 基于叶绿体DNA的山茶属植物的分子系统学和生物地理学初探[硕士论文]. 杭州: 浙江理工大学.
[14]	Li Guang-qing, Sun Li, Liu Yan. 2005. Pollen features of 6 species in Sect. Theopsis of genus Camellia. Journal of Tropical and Subtropical Botany, 13 (1):40-44. (in Chinese)
	李广清, 孙立, 刘燕. 2005. 山茶属连蕊茶组6种植物花粉形态特征研究. 热带亚热带植物学报, 13 (1):40-44.
[15]	Li L, Hu Y, He M, Zhang B, Wu W, Cai P, Huo D, Hong Y. 2021. Comparative chloroplast genomes:insights into the evolution of the chloroplast genome of Camellia sinensis and the phylogeny of Camellia. BMC Genomics, 22 (1):1-22. doi: 10.1186/s12864-020-07350-y
[16]	Li Qian, Guo Qiqiang, Gao Chao, Li Hui’e. 2020. Characterization of complete chloroplast genome of Camellia weiningensis in Weining,Guizhou Province. Acta Horticulturae Sinica, 47 (4):779-787. (in Chinese)
	李倩, 郭其强, 高超, 李慧娥. 2020. 贵州威宁红花油茶的叶绿体基因组特征分析. 园艺学报, 47 (4):779-787. doi: 10.16420/j.issn.0513-353x.2019-0410 URL
[17]	Liang Guo-xiao, Liu Kai, Ma Jin-lin, Chen Guo-chen, Ye Hang, Jiang Ze-peng. 2017. Molecular taxonomy and identification of Camellia osmantha. Nonwood Forest Research, 35 (1):26-29,58. (in Chinese)
	梁国校, 刘凯, 马锦林, 陈国臣, 叶航, 江泽鹏. 2017. 香花油茶分子分类与鉴定. 经济林研究, 35 (1):26-29,58.
[18]	Liao Chu-qin, Miao Shen-yu. 2023. Advances in the traditional and modern classification of Camellia. Chinese Journal of Tropical Agriculture, 43 (7):37-45. (in Chinese)
	廖初琴, 缪绅裕. 2023. 山茶属植物传统及现代分类的研究进展. 热带农业科学, 43 (7):37-45.
[19]	Lin Ping, Wang Kai-liang, Yao Xiao-hua, Ren Hua-dong. 2023. Development of DNA molecular ID in Camellia oleifera germplasm based on Transcriptome-Wide SNPs. Scientia Agricultura Sinica, 56 (2):217-235. (in Chinese) doi: 10.3864/j.issn.0578-1752.2023.02.002
	林萍, 王开良, 姚小华, 任华东. 2023. 基于转录组SNP构建油茶主要品种资源的分子身份证. 中国农业科学, 56 (2):217-235. doi: 10.3864/j.issn.0578-1752.2023.02.002
[20]	Lin Xiu-yan, Peng Qiu-fa, Lü Hong-fei, Du Yue-qiang, Tang Bi-ying. 2008. Leaf anatomy of Camellia sect. Oleifera and sect. Paracamellia (Theaceae) with reference to their taxonomic significance. Journal of Systematics and Evolution, 46 (2):183-193. (in Chinese) doi: 10.3724/SP.J.1002.2008.07057
	林秀艳, 彭秋发, 吕洪飞, 杜跃强, 汤妣颖. 2008. 山茶属油茶组和短柱茶组叶解剖特征及其分类学意义. 植物分类学报, 46 (2):183-193.
[21]	Lin P, Yin H, Wang K, Gao H, Liu L, Yao X. 2022a. Comparative genomic analysis uncovers the chloroplast genome variation and phylogenetic relationships of Camellia species. Biomolecules, 12 (10):1474. doi: 10.3390/biom12101474 URL
[22]	Lin P, Wang K, Wang Y, Hu Z, Yan C, Huang H, Ma X, Cao Y, Long W, Liu W. 2022b. The genome of oil-Camellia and population genomics analysis provide insights into seed oil domestication. Genome Biology, 23:1-21. doi: 10.1186/s13059-021-02568-9
[23]	Lu H, Jiang B, Shen Z, Shen J, Peng Q, Cheng C. 2008. Comparative leaf anatomy,FTIR discrimination and biogeographical analysis of Camellia section Tuberculata(Theaceae)with a discussion of its taxonomic treatments. Plant Systematics and Evolution, 274:223-235. doi: 10.1007/s00606-008-0047-6 URL
[24]	Lu H, Jiang W, Ghiassi M, Lee S, Nitin M. 2012. Classification of Camellia(Theaceae) species using leaf architecture variations and pattern recognition techniques. PLoS ONE, 7 (1):e29704. doi: 10.1371/journal.pone.0029704 URL
[25]	Ming Tien-lu. 1998. The classification,differentiation and distribution of the genus Camellia Sect. Camellia. Acta Botanica Yunnanica, (2):2-23. (in Chinese)
	闵天禄. 1998. 山茶属山茶组植物的分类,分化和分布. 植物分类与资源学报,(2):2-23.
[26]	Ming Tien-lu. 1999. A systematic synopsis of the genus Camellia. Acta Botanica Yunnanica, 21 (2):149-159. (in Chinese)
	闵天禄. 1999. 山茶属的系统大纲. 云南植物研究, 21 (2):149-159.
[27]	Mu Xiao-ting, Liao Zhen-cheng, Lin Cai-jin, Zheng Xi, Lin Jian-yong, He Jian-ping, Jiang Li-bing. 2023. Genetic diversity and relationship analysis of 55 wild tea germplasm from Qingyuan based on SSR markers. China Tea, 45 (2):36-43. (in Chinese)
	穆小婷, 廖侦成, 凌彩金, 郑溪, 林建勇, 何建平, 江丽冰. 2023. 基于SSR标记的55份清远野生茶树种质资源遗传多样性和亲缘关系分析. 中国茶叶, 45 (2):36-43.
[28]	Pan Li-qin. 2019. Research on molecular mechanism and substance related to red color in leaves of Camellia‘Chunjiang Hongxia’[Ph. D. Dissertation]. Beijing: Chinese Academy of Forestry. (in Chinese)
	潘丽芹. 2019. 山茶杂交品种‘春江红霞’红叶相关呈色物质及其分子基础研究[博士论文]. 北京: 中国林业科学研究院.
[29]	Pierre L M. 1887. Flore forestière de la Cochinchine. Paris: Octave Doin.
[30]	Qiu Lu, Guan De-hong, Fan Shu-guo, Xu Wen-bo. 2019. Fourier infrared spectroscopic analysis of plants DNA from Genus Camellia. Journal of Chuxiong Normal University, 34 (3):34-45. (in Chinese)
	邱璐, 关德红, 范树国, 徐文博. 2019. 山茶亚属植物DNA傅立叶红外光谱分析. 楚雄师范学院学报, 34 (3):34-45.
[31]	Shen J B, Lv H F, Peng Q F, Zheng J F, Tian Y M. 2008. FTIR spectra of Camellia sect. Oleifera,sect. Paracamellia,and sect. Camellia(Theaceae) with reference to their taxonomic significance. Journal of Systematics and Evolution, 46 (2):194-204.
[32]	Shi Si-yun, Wu Wen-fei, Cui Jia, Zhang Yi-yi, Li Zhi-hui, Wang Yi. 2022. Pollen morphology and taxonomic significance of ten species of sect. Chrysantha. Guihaia, 42 (1):68-77. (in Chinese)
	石思云, 吴雯霏, 崔佳, 张伊伊, 李志辉, 王翊. 2022. 10种金花茶组植物的花粉形态及分类学意义. 广西植物, 42 (1):68-77.
[33]	Song Yu, Shi Li-ying, Lu Xuan, Feng Bao-min, Yu Da-yong. 2018. Study on chemical constituents and pharmacological activities of Camellia. China Pharmacy, 29 (15):2143-2148. (in Chinese)
	宋昱, 史丽颖, 卢轩, 冯宝民, 于大永. 2018. 山茶属植物的化学成分及药理活性研究. 中国药房, 29 (15):2143-2148.
[34]	Sealy J R. 1958. A revision of the genus Camellia. London:The Royal Horticultural Society.
[35]	Stuart C P C. 1916. Voorbereidende onderzoekingen ten dienste van de selektie der theeplant:JH de Bussy.
[36]	Tan Xiao-feng, Qi Long-lin, He Jin, Gui Jun, Luo Chun-qing, Hu Fang-ming. 2005. Molecular classification of section Oleifera chang and Section Chrysantha Chang of Camellia L.. Journal of Central South Forestry University, 25 (4):31-34. (in Chinese)
	谭晓风, 漆龙霖, 贺晶, 桂君, 罗春清, 胡芳名. 2005. 山茶属植物油茶组与金花茶组的分子分类. 中南林学院学报, 25 (4):31-34.
[37]	Tang Shao-qing, Shi Su-hua, Chen Yue-qin, Qu Liang-hu, Zhang Hong-da. 1998. Phylogenetic relationship of Camellia nitidissima Chi and its allied species based on random amplified polymorphic DNA. Acta Scientiarum Naturalium Universitatis Sunyatseni, (4):29-33. (in Chinese)
	唐绍清, 施苏华, 陈月琴, 屈良鹄, 张宏达. 1998. 金花茶与近缘种的RAPD分析及分类学意义. 中山大学学报(自然科学版),(4):29-33.
[38]	Tang Shao-qing, Shi Su-hua, Zhong Yang, Wang Yan. 2004. Phylogenetic relationships of golden camellias(sect.Chrysantha,Camellia)in China:evidence from ITS sequences of nuclear ribosomal DNA. Guihaia,(6):488-492,487. (in Chinese)
	唐绍清, 施苏华, 钟杨, 王燕. 2004. 基于ITS序列探讨山茶属金花茶组的系统发育关系. 广西植物,(6):488-492,487.
[39]	Tang Shao-qing, Zhong yang, Shi Su-hua, Chang Hung-ta. 2001. Relationship between Camellia nitidissima Chi and C. petelotii (Merr.) Sealy based on nrDNA ITS. Journal of Wuhan Botanical Research,(6):449-452 (in Chinese)
	唐绍清, 钟扬, 施苏华, 张宏达. 2001. Camellia nitidissima与C. petelotii之间的关系研究——来自nrDNA ITS的证据. 武汉植物学研究(6):449-452
[40]	Tian Min, Li Ji-yuan, Ni Sui, Fan Zheng-qi, Li Xin-lei. 2008. Phylogenetic study on Section Camellia based on ITS sequences data. Acta Horticulturae Sinica, 35 (11):1685-1688. (in Chinese)
	田敏, 李纪元, 倪穗, 范正琪, 李辛雷. 2008. 基于 ITS 序列的红山茶组植物系统发育关系的研究. 园艺学报, 35 (11):1685-1688.
[41]	Vijayan K, Tsou C H. 2008. Technical report on the molecular phylogeny of Camellia with nrITS:the need for high quality DNA and PCR amplification with Pfu-DNA polymerase. Bot Stud, 49 (2):177-188.
[42]	Vijayan K, Zhang W J, Tsou C H. 2009. Molecular taxonomy of Camellia (Theaceae) inferred from nrITS sequences. American Journal of Botany, 96 (7):1348-1360. doi: 10.3732/ajb.0800205 URL
[43]	Wang P, Su J, Wu H, Chen Y, Xie Y, Wang H, He G, Chen N, Wei C, Yang L. 2023. Analysis of germplasm genetic diversity and construction of a core collection in Camellia oleifera C. Abel by integrating novel simple sequence repeat markers. Genetic Resources and Crop Evolution,DOI:10.1007/s10722-022-01519-y.
[44]	Wang Jie, He Wen-chuang, Xiang Kun-li, Wu Zhi-qiang, Gu Cui-hua. 2023. Advances in plant phylogeny in the genome era. Journal of Zhejiang A & F University, 40 (1):227-236. (in Chinese)
	王杰, 贺文闯, 向坤莉, 武志强, 顾翠花. 2023. 基因组时代的植物系统发育研究进展. 浙江农林大学学报, 40 (1):227-236.
[45]	Wen Qiang, Zhu Heng, Ye Jin-shan, Xu Li-an, Xu Lin-chu, Jiang Xiang-mei. 2015. Authentication of Camellia chekiangoleosa and its close species using RPB 2 gene sequences. Molecular Plant Breeding, 13 (11):2559-2565. (in Chinese)
	温强, 朱恒, 叶金山, 徐立安, 徐林初, 江香梅. 2015. 利用RPB2基因序列甄别浙江红山茶及其近缘种. 分子植物育种, 13 (11):2559-2565.
[46]	Wei Yun-hua. 2020. Research progress on phenotypic genetic diversity of Camellia germplasm resources in China. South China Agriculture, 14 (28):66-68. (in Chinese)
	魏云华. 2020. 我国山茶花种质资源表型遗传多样性研究进展. 南方农业, 14 (28):66-68.
[47]	Wu Q, Tong W, Zhao H, Ge R, Li R, Huang J, Li F, Wang Y, Mallano A I, Deng W. 2022. Comparative transcriptomic analysis unveils the deep phylogeny and secondary metabolite evolution of 116 Camellia plants. Plant Journal, 111 (2):406-421. doi: 10.1111/tpj.v111.2 URL
[48]	Xiao Zheng, Li Ji-yuan, Li Zhi-hui, Huang Lian-dong, Fan Zheng-qi, Jiang Chang-jie. 2014. Genetic relationships among species from Camellia Sect. Chrysantha Chang revealed by ISSR analysis. Forest Research, 27 (1):71-76. (in Chinese)
	肖政, 李纪元, 李志辉, 黄连冬, 范正琪, 蒋昌杰. 2014. 金花茶组物种遗传关系的ISSR分析. 林业科学研究, 27(1):71-76.
[49]	Xue Xiao-ming, Zhou Yong-wu, Fang Yan. 2012. Observation of pollen morphology of four species of genus Camellia under scanning electron microscope. Guizhou Agricultural Sciences, 40 (5):25-28. (in Chinese)
	薛晓明, 周用武, 方彦. 2012. 4种山茶属植物花粉形态的扫描电子显微观察. 贵州农业科学, 40 (5):25-28.
[50]	Yang J B, Yang S X, Li H T, Yang J, Li D Z. 2013. Comparative chloroplast genomes of Camellia species. PLoS ONE, 8 (8):e73053. doi: 10.1371/journal.pone.0073053 URL
[51]	Yang Jun-bo, Li Hong-tao, Yang Shi-xiong, Li De-zhu, Yang Ying-yan. 2006. The application of four DNA sequences to studying molecular phylogeny of Camellia(Theaceae). Acta Botanica Yunnanica, (2):108-114. (in Chinese)
	杨俊波, 李洪涛, 杨世雄, 李德铢, 杨莹燕. 2006. 四个DNA片段在山茶属分子系统学研究中的应用. 云南植物研究,(2):108-114.
[52]	Yao Xiao-hua, Ren Hua-dong. 2020. Oil-tea Camellia genetic resource in China. Beijing: Science Press. (in Chinese)
	姚小华, 任华东. 2020. 中国油茶遗传资源. 北京: 科学出版社.
[53]	Ye Chuang-xing. 1988. The subdivisions of genus Camellia with a discussion on their phylogenetic relationship. Acta Botanica Yunnanica,(1):61-67. (in Chinese)
	叶创兴. 1988. 山茶属的分群及它们亲缘关系的探讨. 云南植物研究,(1):61-67.
[54]	Ye Chuang-xing, Chang Hung-ta. 1997. Diagnosis on the systematic development of Theaceae primitive characters and evolutionary trends of the genus Camellia. Acta Scientiarum Naturalium Universitatis Sunyatseni,(3):77-82. (in Chinese)
	叶创兴, 张宏达. 1997. 山茶科系统发育诠释:Ⅸ.山茶属的原始特征及其演化趋向. 中山大学学报:自然科学版,(3):77-82.
[55]	Yu X Q, Gao L M, Soltis D E, Soltis P S, Yang J B, Fang L, Yang S X, Li D Z. 2017. Insights into the historical assembly of East Asian subtropical evergreen broadleaved forests revealed by the temporal history of the tea family. New Phytologist, 215 (3):1235-1248. doi: 10.1111/nph.2017.215.issue-3 URL
[56]	Zhang Min-huan, Zhang Han-qing, Cai Xiu-lan, Wang Hai, Ye Ye, Wu Yi. 2018. SSR analysis of genetic relationship between Camellia germplasm resources. Non-wood Forest Research, 36 (4):130-134. (in Chinese)
	张旻桓, 张汉卿, 蔡秀兰, 王海, 叶烨, 吴毅. 2018. 山茶属种质资源亲缘关系的SSR分析. 经济林研究, 36 (4):130-134.
[57]	Zhang T, Qiu F, Chen L, Liu R, Wang X. 2020. Identification and in vitro anti-inflammatory activity of different forms of phenolic compounds in Camellia oleifera oil. Food Chemistry, 344 (49):128660. doi: 10.1016/j.foodchem.2020.128660 URL
[58]	Zhang W, Kan S L, Zhao H, Li Z Y, Wang X Q. 2014. Molecular phylogeny of tribe Theeae(Theaceae s.s.)and its implications for generic delimitation. PLoS ONE, 9 (5):e98133. doi: 10.1371/journal.pone.0098133 URL
[59]	Zhang Wen-Ju, Rong Jun, Wei Chao-ling, Gao Lian-ming, Chen Jia-kuan. 2018. Domestication origin and spread of cultivated tea plants. Biodiversity Science, 26 (4):16. (in Chinese)
	张文驹, 戎俊, 韦朝领, 高连明, 陈家宽. 2018. 栽培茶树的驯化起源与传播. 生物多样性, 26 (4):16.
[60]	Zhao D W, Hodkinson T R, Parnell J A. 2023. Phylogenetics of global Camellia(Theaceae)based on three nuclear regions and its implications for systematics and evolutionary history. Journal of Systematics and Evolution, 61 (2):356-368. doi: 10.1111/jse.v61.2 URL
[61]	Zhou An-pei, Chen Xi-yan, Liang Jian-kun. 2021. The chloroplast genome characteristics and phylogenetic classification of Camellia oleifera. Journal of Jiujiang University(Natural Science Edition), 36 (1):5-8,14. (in Chinese)
	周安佩, 陈曦颜, 梁坚坤. 2021. 油茶叶绿体基因组特征及其系统分类. 九江学院学报(自然科学版), 36 (1):5-8,14.
[62]	Zhou L, Luo S, Li J, Zhou Y, Wang X, Kong Q, Chen T, Feng S, Yuan M, Ding C. 2021. Optimization of the extraction of polysaccharides from the shells of Camellia oleifera and evaluation on the antioxidant potential in vitro and in vivo. Journal of Functional Foods, 86:104678. doi: 10.1016/j.jff.2021.104678 URL
[63]	Zhu Y, Liang D, Song Z, Tan Y, Guo X, Wang D. 2022. Genetic diversity analysis and core germplasm collection construction of Camellia oleifera based on fruit phenotype and SSR data. Genes, 13 (12):2351. doi: 10.3390/genes13122351 URL
[64]	Zhuang Rui-lin. 2008. Chinese oil tea. 2nd Ed. Beijing: China Forestry Publishing House. (in Chinese)
	庄瑞林. 2008. 中国油茶. 第2版. 北京: 中国林业出版社.

所用标记 Marker	物种 Species	主要结果 Main results	欠缺 Deficiency	分化时间 Diversification ages	参考文献 Reference
16 RAPD	5种油茶组和20种金花茶组植物 5 species from Sect. Oleifera，and 20 in Sect. Chrysantha	油茶组分为3大类，金花茶组分为4大类 Sec. Oleifera and Sect. Chrysantha were classified into three and four clusters，respectively.	茶梅分类不明 C. sasanqua needs further verification.	—	谭晓风等,2005
23 RAPD	红山茶组29个种 29 species from Sect. Camellia	29个种划分为光果红山茶亚组、毛蕊系、滇山茶系、短管红山茶。 29 species grouped Subsect. Lucidissima，Series Chrysanthae and Reticulata，C. glabsipelata.	—	—	邓白罗等,2006
核基因 Nuclear：ITS，waxy；叶绿体 cp DNA： trnL-F，rpl16	山茶属10组21种 Total 21species from 10 sections in Camellia	支持山茶属为一单系，评价了不同类型序列 The results supported the monophyly of genus Camellia，and different types of sequences were evaluated.	属下系统有待研究 The phylogentic system needs further study due to sampling and other reasons	—	杨俊波等,2006
ITS	红山茶组30个种。 30 species from Sect. Camellia	分布于中国中南、华东南地区的多数二倍体物种形成一个单系群，分布于云贵高原及金沙江流域的倍性复杂多样的物种构成了另一个单系群 The Clade Ⅰ included the species distributed in central south and southeast of China formed monophyletic clade，the species distributed in southwest China formed another clade	需采集更多的红山茶组样品 It is necessary to collect more samples from Sect. Camellia	—	田敏等,2008
ITS	山茶属15个组112个种 Total 112 species from 15 sections in Camellia	茶组和糙果茶组为单系类群，支持连蕊茶组和毛蕊茶组合并，单列油茶组，红山茶组、半宿萼茶组和瘤果茶组为多系类群以及金花茶组为并系类群，不支持古茶组、越南茶组和实果茶组的分类 The results supported monophyly of Sect. Thea and Furfuracea，merged section of Theopsis and Eriandra and the formation of Sect. Oleifera. It suggested polyphyletic nature of the Sect. Camellia，Paracamellia，Pseudocamellia，and Tuberculata and paraphyletic nature of Sect. Chrysantha；but did not support sectional status of Sect. Archecamellia，Piquetia，and Sterocarpus	需研究短蕊茶组以揭示山茶属早期的物种进化，需研究红山茶组、短柱茶组和瘤果茶组以描绘山茶属系谱关系 Detailed studies focusing on Sect. Brachyandra are needed to shed light on the early evolution of the genus. Further study of Sect. Camellia，Paracamellia，and Tuberculata are crucial	—	Vijayan et al.,2009
叶绿体cpDNA：rpl16，psbA-trnH，trnL-F，rpl32-trnL	山茶属4个亚属11组28个种 A total of 28 species from 11 sections 4 subgenus in Camellia	金花茶组可能是一个并系或复系类群，与长柄山茶组亲缘关系紧密；超长柄茶组非单系类群 Sect. Chrysantha is paraphyletic or polyphyletic cluste，closely related to Sect. Longipedicellata，Sect. Longissima is not monophyletic clade.	取样面窄，超长柄茶的系统位置仍不明确 The systematic position of C. longissima are still unclear	—	方伟等,2010
叶绿体cpDNA：atpI-H，matK，psbA5′R-ALS-11F，rbc；核基因Nuclear：LEAFY	山茶科圆籽荷属、山茶属、大头茶属和核果茶属的30个物种 A total 30 species from genus Apterosperma，Camellia，Polyspora，and Pyrenaria	山茶属为并系类群，核果茶为多系起源，山茶属和其他3个属之间存在广泛的杂交 Camellia is paraphyletic clade and Pyrenaria is polyphyletic clade. Widespread hybridization between Camellia and Pyrenaria，Polyspora and Parapyrenaria were detected	属间广泛的杂交和形态学的相似性增加了分类的难度 Hybridization，rather than morphological homoplasy has confused the current classification of Theeae	核基因冠层年龄为12.3百万年，叶绿体为15.9百万年 LEAFY data （12.3，10.8-17.1）and plastid DNA（15.9，7.3-18.7）	Zhang et al.,2014
叶绿体cpDNA：matK，rbcL，ycf1，trnL-F	4 个山茶亚属164个物种 A total of 164 species from 4 subgenus in Camellia.	张宏达依据形态学处理的分组均不为单系 All sections in Chang’s taxonomy were polyphyly	系统发育树分支之间关系仍未得到解决 The phylogeny between clusters has not been well solved	—	江正栋,2017
叶绿体全基因组和nrDNA Plastome and nuclear ribosomal DNA	山茶科9个属53个物种 Forty-three representatives from nine genera of Theaceae.	山茶属为独立分支，起源可追溯到中新世晚期 Camellia is an independent clade whose origin can be traced back to the late Miocene	—	预估山茶属冠层分化年龄为6.8百万年 The estimated crown ages of Camellia was 6.8 Ma	Yu et al.,2017
核基因RPB3内含子11-15、23 和waxy序列 Nuclear RPB2 introns 11-15 and 23，and waxy.	山茶属21个组，99 个物种，增加了2个分布于越南的新物种和18个植物志未收录的物种 Total 99 samples in 21 sections of Camellia. 2 new species from Vietnam and 18 species missed in Flora of China	山茶属分为2个分支和10个亚分支，其中“离生雄蕊”和“匹克茶”构成原始分支，“实果茶”“金花茶”“糙果茶” “离蕊茶”“茶”“山茶”“毛连蕊茶”、“短柱油茶”构成进化分支 Two main clades and ten sub-clades bearing taxonomically section，the clades “Corallina”and“Piquetia”are the original taxa，while the“Stereocarpus”，“Furfuracea”，“Heterogenea”，“Thea”，“Camellia”，“Camelliopsis”and “Parcamellia”constructed the evolutionary clade	—	山茶属分化的冠层年龄约为39.5 百万年 Crown age of Camellia is 39.5 Ma	Zhao et al.,2022
405个低拷贝直系同源基因 405 low-copy orthologous genes from core Camellia.	116个山茶属物种 116 plants from almost all sections of Camellia	山茶属系统关系划分为7个组，支持合并茶组和秃茶组、油茶组和短柱茶组、半宿萼茶组和瘤果茶组、连蕊茶组和毛蕊茶组、实果茶组和古茶组和红山茶组；茶组和糙果茶组和金花茶组各自独立 Camellia could be divided into seven clusters，supporting the combinations of Sect. Glaberrima and Thea，Sect. Paracamellia and Oleifera，Sect. Pseudocamellia and Tuberculata，Sect. Eriandria and Theopsis. Sect. Thea and Furfuracea and Chrysantha were monophyletic groups	—	预估山茶属分化冠层年龄为14.3百万年 Crown ages of Camellia was 14.3 Ma	Wu et al.,2022

所用标记 Marker	物种 Species	主要结果 Main results	欠缺 Deficiency	分化时间 Diversification ages	参考文献 Reference
16 RAPD	5种油茶组和20种金花茶组植物 5 species from Sect. Oleifera，and 20 in Sect. Chrysantha	油茶组分为3大类，金花茶组分为4大类 Sec. Oleifera and Sect. Chrysantha were classified into three and four clusters，respectively.	茶梅分类不明 C. sasanqua needs further verification.	—	谭晓风等,2005
23 RAPD	红山茶组29个种 29 species from Sect. Camellia	29个种划分为光果红山茶亚组、毛蕊系、滇山茶系、短管红山茶。 29 species grouped Subsect. Lucidissima，Series Chrysanthae and Reticulata，C. glabsipelata.	—	—	邓白罗等,2006
核基因 Nuclear：ITS，waxy；叶绿体 cp DNA： trnL-F，rpl16	山茶属10组21种 Total 21species from 10 sections in Camellia	支持山茶属为一单系，评价了不同类型序列 The results supported the monophyly of genus Camellia，and different types of sequences were evaluated.	属下系统有待研究 The phylogentic system needs further study due to sampling and other reasons	—	杨俊波等,2006
ITS	红山茶组30个种。 30 species from Sect. Camellia	分布于中国中南、华东南地区的多数二倍体物种形成一个单系群，分布于云贵高原及金沙江流域的倍性复杂多样的物种构成了另一个单系群 The Clade Ⅰ included the species distributed in central south and southeast of China formed monophyletic clade，the species distributed in southwest China formed another clade	需采集更多的红山茶组样品 It is necessary to collect more samples from Sect. Camellia	—	田敏等,2008
ITS	山茶属15个组112个种 Total 112 species from 15 sections in Camellia	茶组和糙果茶组为单系类群，支持连蕊茶组和毛蕊茶组合并，单列油茶组，红山茶组、半宿萼茶组和瘤果茶组为多系类群以及金花茶组为并系类群，不支持古茶组、越南茶组和实果茶组的分类 The results supported monophyly of Sect. Thea and Furfuracea，merged section of Theopsis and Eriandra and the formation of Sect. Oleifera. It suggested polyphyletic nature of the Sect. Camellia，Paracamellia，Pseudocamellia，and Tuberculata and paraphyletic nature of Sect. Chrysantha；but did not support sectional status of Sect. Archecamellia，Piquetia，and Sterocarpus	需研究短蕊茶组以揭示山茶属早期的物种进化，需研究红山茶组、短柱茶组和瘤果茶组以描绘山茶属系谱关系 Detailed studies focusing on Sect. Brachyandra are needed to shed light on the early evolution of the genus. Further study of Sect. Camellia，Paracamellia，and Tuberculata are crucial	—	Vijayan et al.,2009
叶绿体cpDNA：rpl16，psbA-trnH，trnL-F，rpl32-trnL	山茶属4个亚属11组28个种 A total of 28 species from 11 sections 4 subgenus in Camellia	金花茶组可能是一个并系或复系类群，与长柄山茶组亲缘关系紧密；超长柄茶组非单系类群 Sect. Chrysantha is paraphyletic or polyphyletic cluste，closely related to Sect. Longipedicellata，Sect. Longissima is not monophyletic clade.	取样面窄，超长柄茶的系统位置仍不明确 The systematic position of C. longissima are still unclear	—	方伟等,2010
叶绿体cpDNA：atpI-H，matK，psbA5′R-ALS-11F，rbc；核基因Nuclear：LEAFY	山茶科圆籽荷属、山茶属、大头茶属和核果茶属的30个物种 A total 30 species from genus Apterosperma，Camellia，Polyspora，and Pyrenaria	山茶属为并系类群，核果茶为多系起源，山茶属和其他3个属之间存在广泛的杂交 Camellia is paraphyletic clade and Pyrenaria is polyphyletic clade. Widespread hybridization between Camellia and Pyrenaria，Polyspora and Parapyrenaria were detected	属间广泛的杂交和形态学的相似性增加了分类的难度 Hybridization，rather than morphological homoplasy has confused the current classification of Theeae	核基因冠层年龄为12.3百万年，叶绿体为15.9百万年 LEAFY data （12.3，10.8-17.1）and plastid DNA（15.9，7.3-18.7）	Zhang et al.,2014
叶绿体cpDNA：matK，rbcL，ycf1，trnL-F	4 个山茶亚属164个物种 A total of 164 species from 4 subgenus in Camellia.	张宏达依据形态学处理的分组均不为单系 All sections in Chang’s taxonomy were polyphyly	系统发育树分支之间关系仍未得到解决 The phylogeny between clusters has not been well solved	—	江正栋,2017
叶绿体全基因组和nrDNA Plastome and nuclear ribosomal DNA	山茶科9个属53个物种 Forty-three representatives from nine genera of Theaceae.	山茶属为独立分支，起源可追溯到中新世晚期 Camellia is an independent clade whose origin can be traced back to the late Miocene	—	预估山茶属冠层分化年龄为6.8百万年 The estimated crown ages of Camellia was 6.8 Ma	Yu et al.,2017
核基因RPB3内含子11-15、23 和waxy序列 Nuclear RPB2 introns 11-15 and 23，and waxy.	山茶属21个组，99 个物种，增加了2个分布于越南的新物种和18个植物志未收录的物种 Total 99 samples in 21 sections of Camellia. 2 new species from Vietnam and 18 species missed in Flora of China	山茶属分为2个分支和10个亚分支，其中“离生雄蕊”和“匹克茶”构成原始分支，“实果茶”“金花茶”“糙果茶” “离蕊茶”“茶”“山茶”“毛连蕊茶”、“短柱油茶”构成进化分支 Two main clades and ten sub-clades bearing taxonomically section，the clades “Corallina”and“Piquetia”are the original taxa，while the“Stereocarpus”，“Furfuracea”，“Heterogenea”，“Thea”，“Camellia”，“Camelliopsis”and “Parcamellia”constructed the evolutionary clade	—	山茶属分化的冠层年龄约为39.5 百万年 Crown age of Camellia is 39.5 Ma	Zhao et al.,2022
405个低拷贝直系同源基因 405 low-copy orthologous genes from core Camellia.	116个山茶属物种 116 plants from almost all sections of Camellia	山茶属系统关系划分为7个组，支持合并茶组和秃茶组、油茶组和短柱茶组、半宿萼茶组和瘤果茶组、连蕊茶组和毛蕊茶组、实果茶组和古茶组和红山茶组；茶组和糙果茶组和金花茶组各自独立 Camellia could be divided into seven clusters，supporting the combinations of Sect. Glaberrima and Thea，Sect. Paracamellia and Oleifera，Sect. Pseudocamellia and Tuberculata，Sect. Eriandria and Theopsis. Sect. Thea and Furfuracea and Chrysantha were monophyletic groups	—	预估山茶属分化冠层年龄为14.3百万年 Crown ages of Camellia was 14.3 Ma	Wu et al.,2022

序号Code	物种Species	登录号Genbank Number	长度/bp Length	GI
1	小果油茶C. meiocarpa	MN078088.1	156 512	2187768699
1	普通油茶C. oleifera	MZ151356.1	156 512	2258873380
2	短柱茶C. brevistyla	MW256435.1	156 550	1999716664
2	小果油茶C. meiocarpa	MZ665482.1	156 550	2282748010
3	东兴金花茶C. tunghinensis	ON208846.1	156 621	2432720715
3	平果金花茶C. pingguoensis	NC_061051.1	156 621	2199204139
4	簇蕊金花茶C. fascicularis	OM868265.1	156 903	2326926816
4	贵州金花茶C. huana	KY626040.1	156 903	1150563370

序号Code	物种Species	登录号Genbank Number	长度/bp Length	GI
1	小果油茶C. meiocarpa	MN078088.1	156 512	2187768699
1	普通油茶C. oleifera	MZ151356.1	156 512	2258873380
2	短柱茶C. brevistyla	MW256435.1	156 550	1999716664
2	小果油茶C. meiocarpa	MZ665482.1	156 550	2282748010
3	东兴金花茶C. tunghinensis	ON208846.1	156 621	2432720715
3	平果金花茶C. pingguoensis	NC_061051.1	156 621	2199204139
4	簇蕊金花茶C. fascicularis	OM868265.1	156 903	2326926816
4	贵州金花茶C. huana	KY626040.1	156 903	1150563370