世界紫堇属（罂粟科）的分子系统学与生物地理学研究

	世界紫堇属（罂粟科）的分子系统学与生物地理学研究
	陈俊通
导师	孙航
关键词	紫堇亚科，紫堇属，系统发育，生物地理，组级新分类系统 Fumarioideae, Corydalis, Phylogeny, Biogeography, New classification
摘要	紫堇属Corydalis是罂粟科Papaveraceae紫堇亚科Fumarioideae中最大的一个属，约有500种，主要分布于北温带。该属在我国的西南地区有着极高的物种多样性。前期关于紫堇属的系统发育研究往往基于少数几个叶绿体基因片段和少数类群取样，未能很好解决本属内的组间系统发育关系，且所提出的亚属级和组级分类系统一直存在争议。因此，本研究对紫堇属的所有组进行全面系统地取样，并基于叶绿体基因组水平构建完整的属下系统发育关系，探讨了该属的生物地理起源和现代分布格局形成机制。 1. 紫堇属的分子系统学研究本研究对紫堇属约235种的280个样品的叶绿体基因组进行分析，其中273个为新测序。这些样品涵盖了当前紫堇属分类系统的42组和5个独立的“系”，即Flora of China（FOC）中的紫堇属分类系统40组和5“系”加上欧洲紫堇组sect. Radix-cava（中国无分布）和中尼黄堇组（nomen nudum，中国新纪录）。同时，这些样品也涵盖了早期Wu et al.系统中的41组，包括曾独立为组的疆堇组sect. Roborowskia和变根紫堇组sect. Linstowianae。基于叶绿体基因组中65个CDS串联形成的超级矩阵，分别构建贝叶斯树、最大似然树、最大简约树，重建了紫堇属较为完整的系统发育关系。同时结合形态学、地理分布，系统而深入地探讨了该属的属下系统发育关系。结果表明，基于叶绿体基因组数据构建的紫堇属系统发育树具有很高的分辨率，且在组级水平和物种水平上强有力地解释了紫堇属的属下系统发育关系，同时也发现一些以前未识别的关系。具体结果如下：1）ML树、BI树和MP树得出了基本一致的拓扑结构；2）系统发育结果再次证实了紫堇属是一个单系类群，与烟堇属Fumaria-藤堇属Ceratocapnos-假烟堇属Pseudofumaria分支互为姊妹关系，并具有极高的支持率；3）属下三个亚属（直茎黄堇亚属subg. Chremnocapnos、黄堇亚属subg. Sophorocapnos、紫堇亚属subg. Corydalis）的单系性也得到很好的支持；4）三个亚属内可识别出6个高支持率的超级分支，即直茎黄堇亚属和黄堇亚属各对应1个超级分支，紫堇亚属内包括4个超级分支，这些超级分支也得到了地理分布证据的支持；5）当前42组和5“系”的紫堇属分类系统中23组和1“系”为单系类群；6）本研究在紫堇属下定义了40个高支持率的组分支，其中15个大致与当前的紫堇属分类系统一致，并重新定义或命名了25个；除Orthopodae超级分支内Elatae-Flexuosae-Priapos-Hamatae-Fasciculatae分支互相之间的关系尚未得到充分解决完外，其它分支之间的关系都得到了充分解决。 2. 紫堇属生物地理学研究运用PAML中的MCMCTree对起源分化时间估算以及运用RASP进行祖先分布区重建。结果表明，紫堇属于晚始新世（36.28Ma）可能在喜马拉雅-横断山地区起源，随后迅速分化（35.49 Ma）。6个超级分支均起源于晚始新世至早渐新世的喜马拉雅-横断山地区，除黄堇亚属于晚渐新世的东亚地区开始分化，其余5个超级分支均于早渐新世的喜马拉雅-横断山地区开始分化。后续各个组的分化主要集中在中新世，这可能与青藏高原的隆升及亚洲季风气候的形成与加剧密切相关，进而在喜马拉雅-横断山地区形成本属的分化中心。其中，东亚-北美间断的类群（黄堇组和古紫堇组）的迁移扩散主要发生在中新世暖期，此间白令陆桥可能发挥了桥梁作用。 3. 紫堇属组级新分类系统基于紫堇属组级系统发育关系，并结合形态学证据、地理分布、分化时间和起源地，本研究提出了一个全新的完整的紫堇属组级新分类系统，包括3亚属，41组（直茎黄堇亚属2组，黄堇亚属3组，紫堇亚属36组）。在紫堇亚属内新建立4个超组，即古紫堇超组supersect. Archaeocapnos、延胡索超组supersect. Corydalis、直梗紫堇超组supersect. Orthopodae和弯梗黄堇超组supersect. Campylopodae。在该新系统中，新建立6个组，即半裸茎紫堇组sect. Potaninianae、川东紫堇组sect. Acuminatae、双斑黄堇组sect. Bimaculatae、米林紫堇组sect. Lupinoides、卡惹拉黄堇组sect. Inopinatae、穆坪紫堇组sect. Flexuosae（新等级）。此外，本研究将积鳞紫堇C. lidenii Z.Y.Su处理为叶状苞紫堇C. microflora (C.Y.Wu & H.Chuang) Z.Y.Su & Lidén的异名。; The genus Corydalis, the largest genus of Papaveraceae subfamily Fumarioideae, contains ca. 500 species mainly distributed in the North Temperate zone with the highest species diversity in SW China. The relationships within this genus based on few markers and very limited taxon sampling have long been controversial, resulting in contrasting taxonomic proposals in delimitation of subgenera and sections. This study aims to clarify infrageneric relationship based on the chloroplast genome data and comprehensive taxon sampling, and to analyze the mechanisms of origination of biogeographical pattern and formation of modern geographical pattern of Corydalis. 1. Molecular phylogeny of Corydalis In this study, a total of 280 accessions of 235 species of Corydalis were analyzed, 273 of which were newly sequenced. These samples cover all the 42 sections and five independent “series” of the current classification of Corydalis, which includes the 40 sections and five independent “series” of FOC plus sect. Radix-cava (no distribution in China) and sect. Filicinae (nomen nudum, new record of China). These samples also cover all 41 sections (including sect. Roborowskia, sect. Linstowianae that were once independent sections) recognized by Wu et al. Phylogenetic trees were reconstructed using maximum likelihood, Bayesian inference and maximum parsimony based on 65 protein-coding sequences. Using the reconstructed complete phylogeny of Corydalis, combined with morphology and geographical distribution, the infrageneric relationship of the genus were systematically and deeply discussed. Our results show that the phylogenetic trees of Corydalis were reconstructed based on chloroplast genome data with high resolution, and can clarify the infrageneric phylogenetic relationship of Corydalis at the section level and species level, and uncover some previously unrecognized relationships. The major results are: 1) The ML, BI and MP analyses yielded almost identical tree topologies; 2) Our phylogenetic results confirmed the monophyly of Corydalis and the Fumaria-Ceratocapnos-Pseudofumaria clade is resolved as sister to Corydalis; 3) the monophyly of the three subgenera is again established; 4) six superclades with strong support can be identified in the three genera: subg. Chremnocapnos and subg. Sophorocapnos respectively correspond to one superclade, and subg. Corydalis includes four superclades (named as four new supersections), which also supported by evidence of geographic distribution; 5) in the current classification of Corydalis (including 42 sections and five “series”), 23 sections and one “series” are monophyletic groups; 6) our results support that Corydalis is divided into 40 clades with high support, 15 of which are roughly consistent with the current classification of Corydalis, and 25 sections have been redefined or named; only the internal relationship of the Elatae-Flexuosae-Priapos-Hamatae-Fasciculatae clade within supersect. Orthopodae has not been fully
语种	中文
	2022-05
学位授予单位	中国科学院大学
文献类型	学位论文
条目标识符	http://ir.kib.ac.cn/handle/151853/75140
专题	昆明植物所硕博研究生毕业学位论文
推荐引用方式 GB/T 7714	陈俊通. 世界紫堇属（罂粟科）的分子系统学与生物地理学研究[D]. 中国科学院大学,2022.

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