Research Progress on Phylogeny and Classification of Camellia

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

Acta Horticulturae Sinica ›› 2024, Vol. 51 ›› Issue (1): 53-66.doi: 10.16420/j.issn.0513-353x.2023-0192

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Research Progress on Phylogeny and Classification of Camellia

KUANG Xuekun, ZHENG Qian, KONG Qingbo, ZHOU Jiasi, ZHOU Lijun, FENG Shiling, DING Chunbang, CHEN Tao^*()

College of Life Sciences，Sichuan Agricultural University，Ya’an，Sichuan 625014，China

Received:2023-06-09 Revised:2023-08-15 Online:2024-01-25 Published:2024-01-16
Contact: *（E-mail：chentao293@163.com）

Abstract

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

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.

Figures/Tables 4

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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

Research Progress on Phylogeny and Classification of Camellia

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