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中文题名:

 中国及泛喜马拉雅地区滇紫草属(广义)(Onosma L.) 的分类学研究    

姓名:

 何毅    

保密级别:

 公开    

论文语种:

 中文    

学科代码:

 071001    

学科专业:

 植物学    

学生类型:

 博士    

学位:

 理学博士    

学位类型:

 学术学位    

学位年度:

 2019    

校区:

 北京校区培养    

学院:

 生命科学学院    

研究方向:

 种子植物分类学    

第一导师姓名:

 刘全儒    

第一导师单位:

 北京师范大学生命科学学院    

提交日期:

 2019-06-24    

答辩日期:

 2019-05-28    

外文题名:

 A TAXONOMIC STUDY OF THE GENUS ONOSMA L. S. L. (BORAGINACEAE) IN CHINA AND PAN-HIMALAYA    

中文关键词:

 广义滇紫草属 ; 紫草科 ; 分类修订 ; 微形态 ; 分子系统学 ; 泛喜马拉雅 ; 中国    

外文关键词:

 Onosma s.l. ; Boraginaceae ; taxonomic revision ; micromorphology ; molecular phylogeny ; Pan-Himalaya ; China    

中文摘要:
广义滇紫草属(Onosma L. s. l.)隶属于紫草科紫草亚科紫草族蓝蓟亚族(Subtrib. Echiinae A. DC.),全世界约有250余种,主要分布于旧世界温带地区,以地中海及西亚地区为分布中心,向东北延伸至阿勒泰山及西伯利亚,东南进入泛喜马拉雅地区。本研究所着眼的中国及泛喜马拉雅地区涵盖了滇紫草属向东扩散的南北两支,与其分布中心相比,这一地区的各方面研究都相对薄弱:狭义滇紫草属和囊萼紫草属(Maharanga A. DC.)的分合一直存在争议,对于这一类群的野外居群调查十分有限,馆藏标本数量很少,在物种划分、名实考证、属下分类系统上仍存在大量问题。因此,对本研究区域广义滇紫草属植物的全面系统研究既有助于完成《泛喜马拉雅植物志》的编写任务,更对于理解该属的扩散与演化也具有重要意义。 本研究在文献考证的基础上,查阅了国内外33个标本馆的标本及标本照片共3000余份。历时3年,前往云南、四川、甘肃、青海、新疆、西藏等地进行了累计4个月的野外采集,采集到广义滇紫草属植物31种2变种,共计93个居群,980余份标本,对花果形态、叶、花粉等性状进行了重新地评估和总结,完成了对本区滇紫草属植物的分类修订;利用分子系统学研究手段,结合核基因ITS、叶绿体基因trnL-F、rpl32-trnL、rps16、trnS-G以及叶绿体基因组构建了滇紫草属的系统发育关系。得到了以下主要结论: 1)增补和修正了6种滇紫草的花冠形态性状描述;认为前人研究所指出的花冠形状、花药联合情况、花丝基部形态以及花冠基部腺体的毛被对于物种划分确有重要意义,但花冠内面毛被在种内并不稳定,不宜单独作为物种的划分依据;认为花冠颜色除了可以作为物种识别的重要依据还具有系统学意义。 2)滇紫草属植物小坚果的基本形式为角卵形,果实大小、喙长、肩部与基部的凸起程度对于果实形态识别具有重要意义;首次在滇紫草属植物中报道了果实基端具有凸起和雌蕊基具骨质托状结构;果实的表面微形态可以识别出三种类型;果实大小和表面纹饰特征反映出了明显的地理相关性及系统学意义;提供了基于果实形态的检索表。 3)叶表皮毛主要可分为具基盘的长硬毛和不具基盘的糙伏毛两种类型;表皮毛的基盘式样均一,未发现星状基盘类型,对于构建本区物种属下分类系统无法提供有价值的依据;毛被的朝向和颜色、叶缘、叶脉序及叶背颜色对于区分近缘种具有一定的分类学价值;解剖学证据显示昭通滇紫草和小叶滇紫草的叶片均为异面叶。 4)花粉形状可分为扁球形、近球形、长球形及哑铃形等多种,具三孔沟或三合孔沟,等极性或异极性;表面常具有褶皱和瘤突;西亚和中亚类群的花粉几乎全部为梨形,异极性三合孔沟,喜马拉雅类群的花粉则全部为等极性,大部分为三孔沟;聚类分析和主成分分析均表明囊萼紫草的花粉类型与喜马拉雅物种更为接近,二者与于西亚和中亚类群差别明显,花粉形态可以为本属的属下系统划分提供有力的支持。 5)分子系统学研究结果表明:广义滇紫草属是单系类群,可分为地中海-西亚-中亚支和喜马拉雅支两支;喜马拉雅支下可识别出5个支系,囊萼紫草属嵌入其中;分化时间推算喜马拉雅支的起源时间约为中新世早期至晚期,其下各支系的分化时间为晚中新世至上新世早期。 6)确认中国及泛喜马拉雅地区共有广义滇紫草属植物49种1亚种3变种;发现并描述2个新物种:山南滇紫草(O. lhokaensis Y. He & Q.R. Liu)和富蕴滇紫草(O. fuyunensis Y. He & Q.R. Liu);将10个名称处理为新异名:尾苞滇紫草(O. dolichouta W.T. Wang)处理为尖苞滇紫草(O. bracteata Wall.)的异名,并报道了该种在中国的新纪录;壤塘滇紫草(O. liui Kamelin & Popova)处理为马尔康滇紫草(O. maaikangensis W.T. Wang)的异名;丽江滇紫草(O. lijiangensis Y.L. Liu)处理为密花滇紫草(O. conferta W.W. Smith)的异名;川西滇紫草(O. mertensioides I.M. Johnston)、细茎滇紫草(O. tenuicaulis Riedl)、察隅滇紫草(O. zayuensis Y.L. Liu)和乡城滇紫草(O. xiangchengensis W.T. Wang)处理为德钦滇紫草(O. wardii (Smith) I.M. Johnston)的异名;宽叶丛茎滇紫草(O. waddellii f. latifolia W.T. Wang)处理为腺花滇紫草(O. adenopus I.M. Johnston)的异名;匍枝滇紫草(O. brachylina (I.M. Johnston) I.M. Johnston)处理为西藏滇紫草(O. waltonii Duthie)的异名;长花滇紫草(O. hookeri var. longiflora Duthie ex Stapf)处理为细花滇紫草(O. hookeri C.B. Clarke)的异名;为尖苞滇紫草和西藏滇紫草指定了选模式;确认昭苏滇紫草(O. echioides L.)和单茎滇紫草(O. simplicissima L.)在中国没有分布;提出了新的属下分类系统,将广义滇紫草属分为2个亚属:原滇紫草亚属(subgen. Protonosma (M. Popov) Y. He)和滇紫草亚属(subgen. Onosma),原滇紫草亚属下分5个组:囊萼紫草组(sect. Maharanga (A. DC.) Gurke.)、狭叶组(sect. Angustifoliae Y. He)、长苞组(sect. Bracteatae Y. He)、原滇紫草组(sect. Protonosma M. Popov)和雅谷组(sect. Jacotianae Y. He);滇紫草亚属下分1组:滇紫草组(sect. Onosma)。
外文摘要:
Onosma L. s.l. (Boraginaceae: Subtrib. Echiinae A. DC.), consisting more than 250 species, mainly distributed in the temperate regions of the old world. This genus has its center of distribution in West Asia (Iran and Turkey), extending northeast to Altay Mountains and Siberia, and southeast into the Pan-Himalaya region. This study in China and Pan-Himalaya region covered both two branches, comparing with its diversity center, the taxonomic study of Onosma species from this area is relatively weak in most aspects: The relationship between Onosma l.l. and Maharanga A. DC. is still controversial; Field survey and amount of specimens are seriously insufficient; The nomenclature, species identification and infrageneric classification have been far from satisfactory. Therefore, a comprehensive taxonomic revision of Onosma in this region will contribute to the compilation of Flora of Pan-Himalaya, as well as a better comprehension of the migration and evolution of this genus. Based on literature examination, the author examined more 3000 specimens (including high resolution specimen pictures) from 33 herbariums all over the world. Type locations in Yunnan, Sichuan, Gansu, Qinghai, Xinjiang and Xizang Province have been visited in four months’ field trips during the past three years, more than 980 specimens including 93 populations of 31 species 2 varities have been collected. Morphology characters of flower, nutlet, leaf indumentum and pollen have been carefully revalued and summarized, contributed to a comprehensive taxonomic study of the genus Onosma s.l. Phylongeny trees based on nuclear gene ITS, chloroplast gene trnL-F, rpl32-trnL, rps16, trnS-G and chloroplast genome have been built. The results are summarized as follows: 1)Supplement and correct morphology characters of flowers of 6 Onosma species; The shape of corolla, nature of anthers and filaments, indumentum of nectary are significant to species identification, but indumentum inside corolla is not a stable character among different populations; Color of corolla is not only useful to identy species, but also has systematic significance. 2)The shape of nutlet of Onosma species is usually trullate; The size of nutlet, length of beak, projection of shoulder and base are significant characters to identify nutlets; Stipitate hilum and gynobase with four cartilaginous cushion-like lobes are first reported in Onosma; Three micromorphology types of nutlet surface can be recongnized; A close correlation among size, surface ornamentation and geographical distribution are found; Nutlet morphology show strong systematic significance. A key based on nutlet morphology is provided.3)There are two types of leaf indumentum: long bristle surrounded by a conspicuous basal tubercle and shorter strigose without basal tubercle. All species share the uniformity of leaf indumentum, the shape of tubercle cannot provide evidence to section division; The direction and distribution of indumentum, the nature of leaf margin, color and venation could be used to identify a few closely-related species; Mesophyll of O.cingulata and O. sinica are reported to be dorsiventral. 4)Pollen shape varies from suboblate, oblate-spheroidal, prolate-spheroidal, subprolate and prolate, 3-colporate or 3-syncolporate, isopolar or heteropolar; Ornamentation is always regulate and scabrate; Pollen of most species from West and Middle Asia are pear-shaped, 3-syncolporate and heteropolar,while speices from Himalaya region are isopolar, most 3-colporate; UPGMA and PCA anylaysis shows that pollens of Maharanga and other Himalayan Onosma s.l. share more similarities than West and Middle Asian species. Pollen morphology shows strong systematic significance: 5)Molecular phylogenetic study shows that Onosma L. s.l. is monophyly, which can be divided into two large clades: Mediterranean-West Asia-Middle Asia clade and Himalaya clade;Within the Himalaya clade, five groups can be recongnized, including Maharanga; Divergence time estimate suggests that the early radiation of Himalaya clade begins at early to late Miocene and the diversification within this clade starts during late Miocene and early Pliocene. 6)There are 49 species, 1 subspecies and 3 varieties of Onosma s.l. in China and Pan-Himalaya region; 2 species are found new to science, O. lhokaensis Y. He & Q.R. Liu and O. fuyunensis Y. He & Q.R. Liu; 10 names are treated as new synonyms: O. dolichouta W.T. Wang is treated as the synonym of O. bracteata Wall.; O. liui Kamelin & Popova to O. maaikangensis W. T. Wang; O. lijiangensis Y.L. Liu to O. conferta W.W. Smith; O. mertensioides I.M. Johnston, O. tenuicaulis Riedl, O. zayuensis Y.L. Liu and O. xiangchengensis W.T. Wang to O. wardii (Smith) I.M. Johnston; O. waddellii f. latifolia W.T. Wang to O. adenopus I.M. Johnston; O. brachylina (I.M. Johnston) I.M. Johnston to O. waltonii Duthie; O. hookeri var. longiflora Duthie ex Stapf to O. hookeri C.B. Clarke; Designated lectotypes for O. bracteata and O. waltonii. Confirm that O. echioides L. and O. simplicissima L. are not distributed in China; Two subgenus are proposed: subgen. 1. Protonosma (M. Popov) Y. He with 5 sections: sect. Maharanga (A. DC.) Gurke., sect. Angustifoliae Y. He, sect. Bracteatae Y. He, sect. Protonosma M. Popov and sect. Jacotianae Y. He ; subgen. 2. Onosma with 1 section: sect. Onosma.
参考文献总数:

 219    

作者简介:

 本人经过研究生期间的科研锻炼,积累了丰富的野外工作经验,掌握了植物区系学、植物分类学与分子系统学研究的相关知识和技术。硕士期间通过文献整理和野外采集(1285号标本),建立了涵盖关山及邻近山地维管植物区系3400余种的名录数据库,开展了关山维管植物区系组成、地理成分和区系比较的研究,发表中文核心期刊一作文章1篇,报道了珍稀濒危植物双蕊兰在黄土高原的新分布;博士论文研究首次建立了中国及泛喜马拉雅地区的广义滇紫草属植物的属下分类系统。提出了1个新亚属(原滇紫草亚属)、3个新组(狭叶组、长苞组、雅谷组)、2个新亚组(旱谷亚组、栎丛亚组);发表了滇紫草属2个新种:山南滇紫草(Onosma lhokaensis)和富蕴滇紫草(Onosma fuyunensis);提出了10个新异名;报道了尖苞滇紫草在中国的新分布;在紫草亚科中首次利用叶绿体基因组数据对广义滇紫草属植物进行了分子系统学研究,绘制了首个紫草亚科物种的叶绿体基因组图谱。发表SCI一作论文3篇,2作论文3篇。    

馆藏地:

 图书馆学位论文阅览区(主馆南区三层BC区)    

馆藏号:

 博071001/19003    

开放日期:

 2020-07-09    

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