青藏高原广义黑瘤衣属地衣分类及系统发育研究
艾敏
导师王立松
关键词系统发育分析,物种界定,模式标本原产地,鳞饼衣属 Phylogenetic analyses, species identification, type localities, Dimelaena
摘要广义黑瘤衣属(Buellia s.lat.)(粉衣目Caliciales,粉衣科Caliciaceae)是地衣型真菌中物种数量最多的属之一,全球约400种。该属地衣外形较小,但形态各异,分布广泛。前人依据外部形态、解剖特征、地衣特征化合物对其进行分类。截至目前,先后有16个属被分离出来。然而,只有四胞衣属(新拟)Tetramelas,喜瘤衣属Amandinea,多瘤胞属Diplotomma等少数属有分子数据支持,大多数物种缺乏分子数据,不能明确其系统位置。由于该属物种繁多、分类困难、分子数据少,目前鲜有文章探讨该属系统发育关系。本研究依托第二次青藏高原科学考察项目,全面调查青藏高原主体区域(喜马拉雅区;亚洲水塔区;祁连山–阿尔金山区;横断山高山峡谷区),获得1000余份相关研究标本。此外,通过多次野外补点采集,国内外标本馆馆藏标本借阅,极大程度地丰富了研究材料。本论文共获得DNA序列900余条,其中760余条纳入分析,基于多基因(nrITS-nrLSU-mtSSU-β-tubulin-RPB2)联合矩阵,对广义黑瘤衣属进行分子系统发育分析,并结合形态、解剖、化学、生态综合对黑瘤衣属物种开展分类研究,取得如下进展: 1. 澄清了青藏高原模式标本原产地物种12个 在400多个黑瘤衣属物种中,中国有记录物种数60余个,18个物种的模式标本采自青藏高原地区,其中14个物种仅在中国有报道。本研究通过多次原产地野外采集,与模式标本,原始描述对照,收集并鉴定了18个物种中的12个物种,并对其进行详细描述,种间讨论,提供特征图片和检索表及给出其在黑瘤衣属中的系统位置。本论文首次澄清这些原产地物种,为这些物种的鉴定建立了标杆。 2. 新发现6个属级系统发育分支 基于五基因联合矩阵构建的粉衣科系统发育树,初步解析了黑瘤衣属及其相关属的系统演化关系。系统发育树显示广义黑瘤衣属共形成了三个大的分支(Clades Ⅰ–Ⅲ),其中Clade Ⅰ包含物种数最多,拓扑结构稳定,关系解析最为清晰,被作为研究的着手点。Clade Ⅰ下共发现7个属级分支,其中四胞衣属为早期成立的属,其单系性得到支持,另外6个属级分支,因需要对其进行更深层次的分析才能确定分类地位,本研究暂把这6个属级分支处理为groups(group A–C,E–G),这些groups在生境,形态特征,特征化合物等方面均有明显差异,通过进一步研究,有希望提升为属。本论文将提供以上groups的详细信息及代表物种的特征图片。 3. 澄清了A group和F group物种组成 基于系统发育、形态、化学研究证据,梳理了A group(Buellia polita-group)共3个种级分支,其中新种2个,外部形态和解剖特征是A group物种划分的重要依据。F group(Buellia subarmeniaca-group)共5个种级分支,其中新种2个,特征化合物差异是F group物种划分的关键特征。本论文提供了以上8个物种的详细描述,种间讨论,化学信息,特征图片和检索表,以便日后物种鉴定。 4. 修订鳞饼衣属Dimelaena含usnic acid物种三个 鳞饼衣属为黑瘤衣属的姊妹群,早期被误放于蜈蚣衣科当中,本研究获得该属模式种模式标本采集地的分子数据,基于三基因联合矩阵(nrITS-nrLSU-mtSSU)构建了系统发育树,证实了它隶属于粉衣科。此外探讨了属模式种多样的化学型不能作为物种划分依据,但形态差异同几个单系分支具有对应关系,可以作为物种划分依据。本研究修订了鳞饼衣属中含usnic acid物种3个,包括1个新种和1个新组合,并重新界定了鳞饼衣属模式种范围。; Buellia s.lat. is one of the largest genera of lichenized fungi, containing about 400 species. Species within Buellia are usually small in size, but have diverse morphology and are widely distributed. Previous taxonomic studies by several lichenologists were based on external morphology, anatomy and chemistry. So far, a total of 16 genera have been segregated from Buellia s.lat. However, only a few of these genera, such as Tetramelas, Amandinea and Diplotomma, have been validated as monophyletic clades by molecular data. Most of the species lack molecular data, so their phylogenetic positions are still unclear. Due to the large number of species in Buellia, difficulty in classification and lack of molecular data, phylogenetic relationships in this genus are rarely discussed. With the support of the Second Tibetan Plateau Scientific Expedition and Research Program (STEP), the main areas of the Qinghai-Tibet Plateau were comprehensively surveyed and more than 1,200 specimens of this genus were obtained. With the further addition of fresh samples collected from multiple field trips and specimens borrowed from domestic and foreign herbaria, the available research materials were expanded. Over 900 DNA sequences were generated in this study, of which over 760 were analyzed. Based on multi-gene (nrITS-nrLSU-mtSSU-β-tubulin-RPB2) matrix, molecular phylogenetic analyses of Buellia s.lat. were carried out. Combined with morphological, anatomical, chemical, molecular and ecological characteristics, comprehensive classifications of Buellia species were carried out. The main results are summarized below. 1. Identification of 12 species from their type localities Among more than 400 species of Buellia s.lat., over 60 species were recorded in China. The type specimens of 18 of these species were collected from the Qinghai-Tibet Plateau, with 14 species reported solely from China. In this study, 12 of those species were collected and identified through multiple field collections, compared with type specimens and original descriptions. Detailed descriptions, comparisons between species, characteristic pictures and a key to these 12 species were provided, in order to facilitate their future identification and use of species. 2. Discovery of 6 well-supported genus-level clades A phylogenetic tree of Caliciaceae was built based on 5-gene concatenated matrix; clarifying the relationships between the genus Buellia s.lat. and its related genera. The results indicated that the genus Buellia s.lat. formed three main clades (Clade Ⅰ–Ⅲ). Of these, Clade Ⅰ contains the largest number of species, with stable topological structure and clear relationships. A total of 7 genus-level clades were formed within Clade Ⅰ: Tetramelas is an established genus, which forms a monophyletic clade; the other 6 genus-level clades need further analysis to determine their taxonomic status, so they are treated as groups (group A–C, E–G). The 6 genus-level clades are distinctively different in th
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/151853/75160
专题昆明植物所硕博研究生毕业学位论文
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艾敏. 青藏高原广义黑瘤衣属地衣分类及系统发育研究[D]. 中国科学院大学,2022.
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