中国白鬼伞族真菌的分类及该族的分子系统学研究
其他题名Study on Taxonomy of Leucocoprineae (Agaricales) from China and Molecular Phylogeny of the Tribe
葛再伟
导师杨祝良
学位专业植物学
关键词伞菌科 白鬼伞族 分类学 系统发育 分布
摘要  通过对白鬼伞族Leucocoprineae真菌进行补充采集和对中国主要标本馆(GDGM、HKAS、HMAS、HMJAU、MHSU)馆藏标本及部分国外标本馆相关标本的借阅,以保存较好的白鬼伞族真菌标本作为研究对象,对盖表鳞片的结构、褶缘囊状体和担孢子的形状、大小及特性作深入、细致的形态解剖研究,首次较为系统、深入地对中国的白鬼伞族真菌进行了分类和修订,纠正了大量鉴定错误,澄清了我国此类真菌分类中存在的一些混乱。主张将丝盖环柄菇属Sericeomyces作为一个独立的属来处理,并对该属的概念进行了重新界定。通过建立一新组,完善了大环柄菇属Macrolepiota的属下分类系统。结合形态分类研究结果,基于内转录间隔区(ITS)、部分编码核糖体大亚基(LSU)、部分编码线粒体小亚基(mtSSU)等3个片段的DNA核苷酸序列,对白鬼伞族进行了分子系统学研究,探讨了白鬼伞族的系统亲缘关系。此外,对研究鉴定出的中国白鬼伞族真菌的地理分布进行了初步总结,提出了西南地区和华南-东南地区是中国白鬼伞族真菌的物种多样性中心。并对三个热带分布的白鬼伞族真菌的分布式样进行了分析及探讨。主要研究结果如下:   根据外部形态和显微结构特征,鉴定确认分布在我国的白鬼伞族真菌有6属39种,存疑种1种。其中绿褶托菇属Clarkeinda 1种;大环柄菇属6种;青褶伞属Chlorophyllum 5种;白环蘑属Leucoagaricus 14种;白鬼伞属Leucocoprinus 6种;丝盖环柄菇属7种。在鉴定出的39个种中,包括5个新种、3个新组合和7个中国新记录种。其中,新种包括:球孢青褶伞Chl. sphaerosporum Z. W. Ge & Zhu L. Yang、东方裂皮大环柄菇M. orientiexcoriata Z. W. Ge & Zhu L. Yang、拟晶缘囊体丝盖环柄菇S. subcrystallifer Z. W. Ge & Zhu L. Yang、拟丁香紫丝盖环柄菇S. subpurpureolilacinus Z. W. Ge & Zhu L. Yang及平顶丝盖环柄菇S. truncatus Z. W. Ge & Zhu L. Yang;新组合为:滴泪柄白环蘑La. lacrymans (T.K.A. Kumar & Manim.) Z. W. Ge & Zhu L. Yang、灰褐丝盖环柄菇S. brunneocanus (J. F. Liang & Zhu L. Yang) Z. W. Ge & Zhu. L. Yang、红鳞丝盖环柄菇S. rubrotinctus (Peck) Z. W. Ge & Zhu L. Yang;中国新记录种为:球盖青褶伞Chl. globosum (Mossebo) Vellinga、红变青褶伞Chl. hortense (Murrill) Vellinga、拟珠鸡白环蘑La. holospilotus (Berk & Broome) Bon、滴泪柄白环蘑、翘鳞白环蘑La. nympharum (Kalchbr.) Bon、浅鳞白鬼伞Lc. cretaceus (Bull.: Fr.) Locq.、丝盖环柄菇S. serenus (Fr.) Bon & Boiffard。对各个种进行了显微形态绘图及描述,并对各个种的鉴别特征及其与相近种的区别进行了讨论,提出了族下分属及属下分种的检索表。   根据外部形态和显微结构特征及分子系统学证据,结合大量的相关文献以及现有属下分类系统,对大环柄菇属的属下分类系统提出了新建议:在Bon提出的两个组(sect. Macrolepiota 和 sect. Macrosporae)的基础上,建立了一个新组Macrolepiota section Volvatae Z. W. Ge & Zhu L. Yang。提出了分组及组下的分种检索表。   丝盖环柄菇属的成员在菌盖表皮结构及担孢子形状等形态特征方面与白环蘑属的差异明显,应从白环蘑属中独立出来。除包括那些盖表及菌柄无明显色素的种类外,丝盖环柄菇属还应包括具有以下特征的种:菌盖表面并非白色,但菌盖表皮是由长形的、近不分化的、呈平伏状或辐射状排列的菌丝组成,它们的担孢子为卵形至杏仁形、孢子近顶端缢缩变尖,无芽孔。原置于白环蘑属中的红染白环蘑等一些种,除了菌盖表皮有颜色外,与丝盖环柄菇属的成员在菌盖表皮结构及担孢子形状等形态特征方面并无本质区别,应转入丝盖环柄菇属中。本文对该属的特征和范围进行了修订。   基于三个基因片段ITS、LSU和mtSSU序列,运用最大简约法和贝叶斯法对白鬼伞族进行了系统发育分析。研究结果表明白鬼伞族不是一个单系类群。伞菌族的模式属伞菌属Agaricus及其近缘属Micropsaliota都聚在白鬼伞族内,使白鬼伞族成为一个并系类群。在形态学上所界定的白鬼伞族下几属中,白环蘑属和白鬼伞属为一复合类群,共同形成一个单系;而大环柄菇属、青褶伞属及修订后的丝盖环柄菇属较为自然,各自形成单系类群;伞菌属及其近缘属为青褶伞属的姐妹群。分子系统发育分析支持将丝盖环柄菇属从白环蘑属中独立出来,也支持将大环柄菇属分为三个组处理。   通过对研究鉴定出的中国白鬼伞族真菌的地理分布进行整理和统计分析,初步得出了我国白鬼伞族真菌的地理分布特点。根据现有资料,白鬼伞族真菌在中国的分布范围较为广泛,西南地区和华南-东南地区是中国白鬼伞族真菌的物种多样性中心。对同是热带分布的长柄大环柄菇、球盖青褶伞及绿褶托菇的世界分布图进行了绘制,并对三者的分布式样进行了分析及探讨。;   Based on extensive field investigation and materials from main mycological herbaria from China, including GDGM, HKAS, HMAS, HMJAU, MHSU, as well as putative collections deposited in abroad herbaria such as BPI and FH, taxonomy and molecular phylogeny of the tribe Leucocoprineae from China were studied. Sericeomyces has been recognized as a separate genus and the infrageneric system within the genus Macrolepiota was improved by introducing a new section, M. sect. Volvatae. Species cited in literature but turned out to be misidentified were discussed and corrected. Based on sequence data from the nuclear ribosomal large subunit gene (nLSU), the internal transcribed spacer region of the nuclear ribosomal array (ITS), and the mitochondrial ribosomal small subunit gene (mtSSU), phylogenetic trees of Leucocoprineae were reconstructed and systematic positions of genera within the tribe and relationships among the genera were discussed. Furthermore, distributions of Leucocopeineae within China were supplemented and discussed with an emphasis on tropical distributed exemplars. Southwestern China and South-Southeastern China are two species diversity centers as suggested by the current data. The main results of the dissertation were summarized as follows:   Specimens of putative Leucocoprineae from China collected by the author and by colleagues were examined. As a result, thirty-nine species were identified which belong to six genera of this tribe, including five species within Chlorophyllum, one species within Clarkeinda, fourteen species within Leucoagaricus, six species within Leucocoprinus, six species within Macrolepiota, and seven species within Sericeomyces. All identified species were described and illustrated in detail, and differences between similar species are provided and discussed. A key to the genera of the Leucocoprineae and keys to the species belong to the six studied genera in the tribe were provided.   Among the thirty-nine identified species, five are new to science and seven are new to China. The new taxa are Chl. sphaerosporum Z. W. Ge & Zhu L. Yang, M. orientiexcoriata Z. W. Ge & Zhu L. Yang, S. subcrystallifer Z. W. Ge & Zhu L. Yang, S. subpurpureolilacinus Z. W. Ge & Zhu L. Yang, and S. truncatus Z. W. Ge & Zhu L. Yang; the new records to China are Chl. globosum (Mossebo) Vellinga, Chl. hortense (Murrill) Vellinga, La. holospilotus (Berk & Broome) Bon, La. lacrymans (T.K.A. Kumar & Manim.) Z.W. Ge & Zhu L. Yang, La. nympharum (Kalchbr.) Bon, La. serenus (Fr.) Bon & Boiffard and Lc. cretaceus (Bull.: Fr.) Locq. Besides, three new combinations, namely La. lacrymans (T.K.A. Kumar & Manim.) Z. W. Ge & Zhu L. Yang, S. brunneocanus (J. F. Liang & Zhu L. Yang) Z. W. Ge & Zhu. L. Yang and S. rubrotinctus (Peck) Z. W. Ge & Zhu L. Yang, are proposed.   With regard to the infrageneric classification of Macrolepiota, section Macrolepiota and section Macrosporae were accepted, and a new section, M. section Volvatae Z. W. Ge & Zhu L. Yang, is proposed to accommodate the volvatae Macrolepiota species based on macro- and micro-morphological characters.   Morphological characters support the recognition of Sericeomyces as a separate genus that differs from Leucoagaricus. The structure of pileal surface of Sericeomyces is an epicutis composed by undifferentiated repent hyphae, while those of Leucoagaricus, a heterogeneous group, are much more diverse; basidiospores of Sericeomyces representatives are constantly without a germ pore and amygdaliform-shaped with an apical papilla, while some species of the Leucoagaricus bear a germ pore and the shapes of the spores are also more diverse. Besides the species with scales typically without pigments, species previously placed in Leucoagaricus but with an epicutis pileal structure and amygdaliform-shaped spores with an apical papilla, such as La. rubrotinctus, should be transferred to Sericeomyces in spite of their colored pileal surface.   Current classifications of the Leucocoprineae were evaluated by means of molecular phylogeny. Parsimony and Bayesian analyses were performed to reconstruct the phylogenetic tree of Leucocoprineae based on sequence data from ITS, LSU, and mtSSU. Results of the analyses show that Leucocoprineae is not monophyletic, with Agaricus, the type genus of tribe Agariceae, and its allies derived from the Leucocoprineae clade and formed the sister clade to Chlorophyllum. With regard to the monophyly of genera circumscribed by morphology within the Leucocoprineae, Sericeomyces recognized as a separate genus got strong statistic supports from both parsimony and Bayesian analyses; revised genera Chlorophyllum and Macrolepiota formed monophyletic group of their own respectively, while genera Leucoagaricus and Leucocoprinus are shown to be artificial, with species of either genus shifted from one to another. Phylogenetic study of Macrolepiota based on ITS data support the recognition of three sections within it. The new section, M. section Volvatae, proposed to contain the volvatae species of Macrolepiota received strong supports from both parsimony and Bayesian analyses.   The geographic distribution of Leucocoprineae within China were summarized and analyzed. Leucocoprineae has a wide distribution in China. Southwestern China and South-Southeastern China are two species diversity centers as suggested by the current data. Distribution map of C. trhacodes, Chl. globosum, as well as M. dolichaula was protracted, and their biogeographic patterns were discussed.
语种中文
2008-01-24
学位授予单位中国科学院昆明植物研究所
文献类型学位论文
条目标识符http://ir.kib.ac.cn/handle/151853/262
专题昆明植物所硕博研究生毕业学位论文
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葛再伟. 中国白鬼伞族真菌的分类及该族的分子系统学研究[D]. 昆明植物研究所. 中国科学院昆明植物研究所,2008.
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