落新妇属和鬼灯檠属的分子系统学及生物地理学研究
朱卫东
学位类型博士
导师孙航
2010-05
学位授予单位中国科学院研究生院
学位专业植物学
摘要落新妇属(Astilbe Buch.-Ham. ex D.Don)、鬼灯檠属(Rodgersia A.Gray)以及大叶子属(Astilboides Engler)同为传统虎耳草科落新妇族植物。大叶子属为中国-日本特有分布类型,鬼灯檠属为东亚特有分布类型,落新妇属为东亚-北美洲际间断分布类型。落新妇族被认为是典型的“北极-第三纪植物类群”。细胞学、类黄酮化学和近年来的分子系统学证据表明落新妇族不是一个自然组合类群。本文对选择东亚特有属——鬼灯檠属、东亚-北美间断分布属——落新妇属,利用多DNA片段,分别重建了两属属下系统发育关系,结合形态学、细胞学和分子系统学研究结果,分别讨论了他们与近缘类群的系统学关系,对属下划分和种的界限进行了认定。结合现代地理分布格局,探讨了两属的生物地理学意义。1. 落新妇属的分子系统学和生物地理学研究,利用叶绿体trnL-F、matK和psbA-trnH及核糖体ITS基因四个DNA片段对东亚-北美东部间断分布落新妇属以及北美分布的近缘类群Saxifragopsis属、Boykinia Group、Heuchera属等进行了系统发育研究。对世界范围内分布的落新妇属近缘类群和属下各种进行了广泛采集,先后获得了中国大陆、台湾岛、朝鲜、日本、尼泊尔、印度尼西亚巴布亚岛以及北美东南部分布种的65份样品。研究结果表明:落新妇属是一个单系类群。落新妇属与鬼灯檠属、大叶子属关系较远。其姊妹群为北美西部分布的Saxifragopsis属。形态学、细胞学和分子系统学证据均支持两属组成一个自然类群组合。新大陆(北美)东南部分布的特有种二回落新妇(Astilbe biternata (Ventenat) Britton ex Kearney)在系统树中没有与旧大陆(亚洲)分布种构成姊妹关系,其系统位置在基于核糖体基因和叶绿体基因构建的拓扑结构中不同。分子证据结合形态学、染色体组型和繁育生物学证据,支持A. biternata为杂交起源,其亲本为两个不同形态型(完全花瓣/无花瓣)的二倍体落新妇属植物。分子证据不支持多花落新妇(Astilbe myriantha Diels)作为溪畔落新妇(A. rivularis Buch.-Ham. ex D.Don)的变种,前者与具有完整花瓣的东亚分布种近缘,基于其植株杂性等特征,本文建议将其重新提升到种级水平。中国东北部和朝鲜半岛分布的朝鲜落新妇(A. koreana)与大落新妇(A. grandis)关系较远,而与落新妇(A. chinensis)近缘,本文不支持朝鲜落新妇作为大落新妇异名,其更可能是落新妇的变种或近缘种。腺萼落新妇(Astilbe rubra)不是单系类群。朝鲜半岛和日本分布的“腺萼落新妇”居群与落新妇近缘,且序列高度同源。云南西部、西藏东南部和印度北部的居群则同台湾分布的长果落新妇(A. longicarpa)、阿里山落新妇(A. macroflora)及巴布亚分布的菲律宾落新妇(A. philippinensis)近缘,本文支持Fanchet的观点将朝鲜半岛和日本分布的“腺萼落新妇”处理为落新妇变种或异名。研究不支持单叶落新妇(A. simplicifolia)作为落新妇内的原始类群,而处于较为进化的位置。溪畔落新妇(A. rivularis)种内存在很高的形态和遗传变异,其种内遗传分化与地理分布相对应,该种可能起源于横断山区,并逐渐向西发展。生物地理学分析表明,落新妇属的起源在中新世中期之前(不晚于11.96百万年前),其起源地很可能在亚洲东北部,并通过白领海峡散布到到新大陆。基于分子系统学研究、原始文献和馆藏模式,对落新妇属进行了分类处理。落新妇属共有16种4变种。另有Astilbe indica (印尼爪哇岛)、Astilbe stoliczkai(喜马拉雅西北部)和Astilbe khasiana(印度卡西亚地区)三个种由于缺少标本和文献支持,其分类学地位尚有疑问。新提升狭叶落新妇至种级水平(Astilbe angustifoliolata (H.Hara) W. D. Zhu, H. Sun et J. Wen stat. nov.)。2. 鬼灯檠属的分子系统学与生物地理学研究:利用叶绿体trnL-F、matK及核糖体ITS基因序列扩增片段对鬼灯檠属(Rodgersia, 5-6个种,21个样品)以及近缘类群大叶子属(Astiloboides)、岩白菜属(Bergenia)、槭叶草属(Mekdenia)、独根草属(Oresitrophe)以及北美西部分布的单种属Darmera属进行了分子系统发育研究。分析结果表明:鬼灯檠属为单系类群,该属同传统落新妇族的大叶子属以及北美西部分布的Darmera属关系最为近缘,并构成姊妹群关系。分子系统学结果支持将鬼灯檠属、大叶子属、Darmera属、槭叶草属、独根草属、岩白菜属归并为单系的自然组合群(Darmera Group),并获得了细胞学和类黄酮化学的证据支持。七叶鬼灯檠(Rodgersia aesculifolia Batalin)不是一个单系类群。分布于横断山区云南西部分布的“滇西鬼灯檠(R. aesculifolia var. henrici)”与同样分布式样的羽叶鬼灯檠(R. pinnata)和西南鬼灯檠(R. sambucifolia)聚在一起,滇西鬼灯檠分布区西限的西藏东南部样品则单独聚为一组并同前者构成姊妹群。秦岭和大巴山地区分布的七叶鬼灯檠则与横断山及西藏分布的鬼灯檠属植物构成姊妹群,进而与日本和朝鲜分布的鬼灯檠构成姊妹关系。基于形态学和本文的分子系统学证据,滇西鬼灯檠与七叶鬼灯檠区别明显,本文支持将滇西鬼灯檠重新提升到种级水平(Rodgersia henrici Franchet)。滇西鬼灯檠云南西部分居群与羽叶鬼灯檠(R. pinnata var. pinnata)近缘,且分布区有重叠,自然杂交可能存在于两种重叠分布区居群中。尼泊尔东部和印度锡金邦分布的孤立种尼泊尔鬼灯檠(R. nepalensis)处于系统树的基部,这与该属起源地以及基于传统形态分支系统学和细胞学证据的鬼灯檠属内进化关系相反。这可能是由于其分布区狭小,加之与属内其他种分布区相隔离,遗传基因相对保守,形态特化特征与遗传进化不同步导致。其形态特化特征很可能是对复杂生态环境的适应。除与属内其他种间断分布的狭域种尼泊尔鬼灯檠,鬼灯檠属属内分子系统进化关系与传统的形态学和细胞学证据一致。生物地理学分析表明:鬼灯檠属的起源于亚洲东北部,并通过秦岭和大巴山地区向横断山区、青藏高原和喜马拉雅地区等地迁移,并在横断山区获得发展,成为该属的现代分布中心。隔离分化、种间自然杂交和多倍化等可能在属内植物进化上起了重要作用。本文还对对鬼灯檠属属下分类进行了处理。鬼灯檠属共有6种2变种。
资助项目Astilbe Buch.-Ham. ex D.Don, Rodgersia A.Gray together with Astilboides Engler. blong to the tribe Astilbeae in Saxifragaceae. Rodgersia is a genus of tall, statuesque herb with unique compound leaves. It consists of six or five species, ranging from Japan to east Himalaya through Qinling-Daba mountain areas and southwest China. Astilbe is a well defined genus of Saxifragaceae, with a disjunct distribution between eastern Asia and eastern North America extending westward to northern India and southward to Philippines, Java Island, and New Guinea.Two separate molecular phylogeny studies using multiple DNA sequences were conducted on Astilbe and Rodgersia as well as their allies. 1. Molecular phylogeny of eastern Asian-eastern North American disjunct Astilbe,Phylogenetic analyses were conducted for the eastern Asian-eastern North American disjunct genus Astilbe (Saxifragaceae) and allies using sequences of plastid matK, trnL-trnF, psbA-trnH, and nuclear ribosomal ITS region. A worldwide 65 collections of Astilbe species and allies were sampled from Mainland China, Taiwan, Korea, Japan, Nepal, United States and New Guinea. The monophyly of Astilbe was supported based on analyses of both ITS and plastid data sets. Astilbe is distinct from Rodgersia and Astilboides, but close related to western North American Saxifragopsis. The combination of Astilbe and Saxifragopsis suggested by Soltis with the name of Astilbe Group was supported by our molecular result, as well as morphological and cytological traits. The plastid and the ITS trees differed primarily on the placement of eastern North American Astilbe biternata. Hybridization and subsequent chloroplast capture may best explain the topological incongruence between neulcear and plastid data. Our results supported a hybrid origin of allotetraploid A. biternata with two Asian dimorphism diploid progenitors. Astilbe rivularis var. myriantha (= A. myriantha) was suggested to be elevated to the species level. Without being allied to A. rivularis, A. myriantha was related to taxa with perfect petals. Astilbe rubra was not monopyletic, “A. rubra” recorded from Korea and Japan is grouped together with A. chinensis and A. koreana, while two collections of A. rubra sampled from of western Yunnan of China grouped with A. longicarpa, A. macroflora from Taiwan, and A. philippinensis from Papua of Indonesia. In present analyses, the northestern Asian Astilbe koreana was not supported as synonym of A. grandis, but being closely allied with A. chinensis and “A. rubra”.Astilbe simplicifolia occupied a derived position other than being a primitive position. Molecular result supported regarding it as a new neotenous derivative of A. thunbergii. High genetic variations were first found in Astilbe rivularis, our molecular deriviations were corresponded with biogeographic distributions of A. rivularis complex. Astilbe rivularis might have an origin in Hengduan Mountains and with a development westward to Himalayas and eastward to eastern Hengduan mountains. Dating analysis based on the Bayesian approach with BEAST estimated the age of the crown Astilbe to be 11.96 mya, the divergent time for intercontinental disjunction in Astilbe was inferred to be not earlier than in middle Miocene. Astilbe originated from northeastern Asia with its allies in western North America. The Bering land bridge might be the most likely migration route for origin of the new world Astilbe. Based on present molecular results, as well as original description papers and Type specimen, 16 species and four variaties were recognized in Astilbe. For the other three species: Astilbe indica (recorded from Java, Indonesia), Astilbe khasiana (recorded from Khasia, India), and Astilbe stoliczkai (recoded from northwest Himalaya), the taxonomical status were still questionable. According to morphological and molecular evidences, Astilbe rivularis var. angustifoliolata was raised to sepecies level with the name of Astilbe angustifoliolata (H.Hara) W. D. Zhu, H. Sun et J. Wen stat. nov.2. Molecular phlogeny of Rodgersia and allies,Rodgersia is a well defined genus native to eastern Asia. Molecular phylogenetic analyses were conducted for Rodgersia, as well as its alies Astilboides, Darmera, Oresitrophe, Bergenia, Mukdenia. The sequences of matK, trnL-trnF and nuclear ITS regions were used. The monophyly of Rodgersia was well supported. Rodgersia was closely related to eastern Asian endemic Astilboides and western North American Darmera; the combination of Rodgersia, Astilboides, Darmera, Oresitrophe, Bergenia, and Mukdenia by Soltis with the name of Darmera group was supported. The key taxonomic traits of leave arrangement and pubescence were not suppoted by molecular result, especially for taxa from Hengduan Mountains and Himalayas. Multiple sampled Rodgersia aesculifolia was not monophyly, samples from Hengduan Mountains (R. henrici = R. aesculifolia var. henrici) were nested with R. pinnata and R. sambucifolia, while samples from southeast Tibet (R. henrici = R. aesculifolia var. henrici) form a clade sister to the former taxa. Samples of R. aesculifolia from Qingling and Daba mountains (R. aesculifolia var. aesculifolia = Triditional R. asculifolia) are distinct with all the above. R. aesculifolia var. henrici is distinct from A. aesculifolia var. aesculifolia and is suggested be raised to spcies level again as Rosgersia henrici Franchet. Populations of R. henrici from western Yunnan are grouping with R. pinnata, natural hybridization are supposed to occur. Rodgersia podophylla from Korea and Japan is sister to Chinese Rodgersia. The furthermore study of infraspecific taxonomy of R. aesculifolia is suggested.The relict Rodgersia nepalensis from eastern Nepal branched first in the combined ITS and plastid tree, which is different from evidences of the traditional morphology and cytology. This might due to its narrow distribution disjuct from other species of Rodgersia, low level of gene flow and subsequent conserved genetic system. It may evolved by polyploidy, the spcecialized morphological character of R. nepalensis may be a strategy for ecological tolerance and self-protection. Our molecular phylogeny of Rodgersia is accordant with the former morphological and cytological evidences. Hybridization and polyploidy may play an important role in evolution and speciation in Rodgersia. Rodgersia may origin from northestern Asia and migrated into Hengduan mountains and Himalayas through Qingling and Daba mountains. Based on present molecular results, as well as original description papers and Type specimen, six species and two variaties were recognized in Rodgersia. Rodgersia henrici was recognized in our study, and was supported to be raised to species level again
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文献类型学位论文
条目标识符http://ir.kib.ac.cn/handle/151853/16102
专题昆明植物所硕博研究生毕业学位论文
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朱卫东. 落新妇属和鬼灯檠属的分子系统学及生物地理学研究[D]. 中国科学院研究生院,2010.
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