中国环柄菇属的分类学及该属的分子系统学研究--兼论冠状环柄菇的群体遗传学
其他题名Study on Taxonomy of Lepiota from China and Molecular Phylogeny of the Genus with Notes on Population Genetics of L. cristata
梁俊峰
学位类型博士
导师杨祝良
2007-05-28
学位授予单位中国科学院昆明植物研究所
学位授予地点昆明植物研究所
学位专业植物学
关键词环柄菇属 分类学 分子系统学 物种概念 形态种 系统发育种 基因流 地理遗传模式
摘要本文通过对中国环柄菇属(Lepiota)外形、显微结构特征和基于ITS、LSU、IGS及mtSSU等4个基因片段的分子系统学研究,探讨了环柄菇属属下分类系统及系统演化关系。利用ITS、LSU、IGS、mtSSU、EF1a和BIP等6个基因片段对绿化白环蘑(Leucoagaricus viriditinctus)复合类群(过去置于环柄菇属)进行了分子生物学研究,探讨了该复合群形态种与系统发育种的关系。利用ITS、IGS和mtSSU等3个基因片段对冠状环柄菇(Lepiota cristata)进行了群体遗传学研究,讨论了冠状环柄菇种群内遗传多样性与生物地理的关系。主要的研究结果和结论如下: 1、分类学与系统学研究 根据对中国、斯里兰卡、欧洲和北美环柄菇属标本(含部分种的模式标本)的研究,共鉴定出环柄菇属(Lepiota)36个种,白环蘑属(Leucoagaricus)6个种,并进行了绘图及描述。其中包括16个新种,即:赭鳞环柄菇(L. ochraceosquamea J. F. Liang & Zhu L. Yang)、宽囊环柄菇(L. ampliocystidia J. F. Liang & Zhu L. Yang)、近梭孢环柄菇(L. submagnispora J. F. Liang & Zhu L. Yang)、暗褐环柄菇(L. fusco-brunnea J. F. Liang & Zhu L. Yang)、窄孢环柄菇(L. contractispora J. F. Liang & Zhu L. Yang)、近栗色环柄菇(L. subcastanea J. F. Liang & Zhu L. Yang)、假冠状环柄菇(L. pseudocristata H. C. Wang ex J F. Liang & Zhu L. Yang)、浅赭环柄菇(L. pallidiochracea J. F. Liang & Zhu L. Yang)、巨孢环柄菇(L. megaspora J. F. Liang & Zhu L. Yang)、窄囊环柄菇(L. angusticystidia J. F. Liang & Zhu L. Yang)、红褐环柄菇(L. rubro-brunnea H. C. Wang ex J F. Liang & Zhu L. Yang)、褐绒环柄菇(L. brunneosquamula J. F. Liang & Zhu L. Yang)、滴孢环柄菇(L. stillispora J. F. Liang & Zhu L. Yang)、变绿白环蘑(La. viridianus J. F. Liang & Zhu L. Yang)、变蓝白环蘑(La. azurescens J. F. Liang & Zhu L. Yang)和灰褐白环蘑(La. brunneocanusJ. F. Liang & Zhu L. Yang);2个新变种,即细环柄菇黑鳞变种(L. clypeolaria var. nigrosquamosa H. C. Wang ex J F. Liang & Zhu L. Yang)和近栗色环柄菇双孢变种(L. subcastanea var. bispora J. F. Liang & Zhu L. Yang);2个新组合,即:冠状环柄菇大孢变种[L. cristata var. macrospora(Zhu L. Yang)J. F. Liang & Zhu L. Yang]和绿化白环蘑[La. viriditinctus(Berk. & Broome)J. F. Liang & Zhu L. Yang];5个新记录种,即:糙鳞环柄菇(L. aspericeps Murrill)、库勒环柄菇(L. kuehneriana Locq.)、褐紫环柄菇(L. brunneolilacea Bon & Boiffard)、锥鳞环柄菇(L. jacobi Vellinga & Knudsen)和娇柔白环蘑[La. tener(P.D. Orton)Bon]。 根据外部形态和显微结构特征,将环柄菇属划分为环柄菇组(sect. Lepiota)、距孢环柄菇组(sect. Stenosporae)、卵孢环柄菇组(sect. Ovisporae)、冠盖环柄菇组(sect. Cristatae)和刺鳞环柄菇组(sect. Echinatae)5个组,给出了分组及组下的分种检索表,并给出了其近缘属白环蘑属部分种的分种检索表。 2、分子系统学研究 通过4个基因片段ITS、LSU、IGS和mtSSU,运用最大简约法、贝叶斯法、T-PTP测验、部分同质性测验和遗传距离分析方法对环柄菇属进行了系统发育分析。研究结果表明目前基于形态特征的分类系统尚不完善,5个组均非单系,并对其盖表鳞片结构及担孢子的演化进行了推断。认为盖表结构由栅状依次向子实层状、念珠状演化;担孢子从企鹅形和纺锤形向椭圆形演化,再演化为麦角形或近三角形和近圆柱形。 3、形态种与系统发育种的关系 通过ITS、LSU、IGS、mtSSU、EF1a和BIP等6个基因片段对受伤或干燥后变蓝绿色的“绿化白环蘑”进行了研究,初步探讨了环柄菇类真菌的形态种与系统发育种的关系。研究结果表明,绿化白环蘑中包含了5个系统发育种,也即一个形态种可能包含一个或多个系统发育种。对环柄菇类真菌中不同基因片段的演化速率进行了比较和推断,显示出如下的演化速率:慢 LSU → mtSSU → ITS →(EF1a、BIP)→ IGS 快。 4、冠状环柄菇的群体遗传学及生物地理的关系研究 运用ITS、IGS和mtSSU等3个基因片段对冠状环柄菇进行了群体遗传学研究,初步探讨了冠状环柄菇种群遗传多样性与生物地理的关系。在冠状环柄菇内,不同地理群体之间存在长距离的基因流,而且这种基因流非常普遍,因此在该种群体间无明显的地理遗传变异式样。白色的担孢子有可能进行长距离的传播,这与前人所推测的白色担孢子不宜作长距离传播的结论相反。根据群体遗传学研究结果,发现在冠状环柄菇复合群中,隐藏着一个有别于冠状环柄菇的新种即假冠状环柄菇,同时发现近亲环柄菇大孢变种(L. fraterna var. macrospora)属于冠状环柄菇种下的一个特殊类群。
其他摘要In this thesis, the macromorphological and micromorphological characters of Lepiota from China and molecular phylogeny based on ITS, LSU, IGS and mtSSU sequences of the genus were studied. The infrageneric system and phylogeny within the genus were discussed. The relationships of morphological and phylogenetic species in the complex Leucoagaricus viriditinctus, previously placed in Lepiota, were elucidated based on ITS, LSU, IGS, mtSSU, EF1a and BIP sequences data. Genetic diversity and biogeography of L. cristata were inferred from ITS,IGS and mtSSU sequences. The main results are summarized as follows: 1. Taxonomy and systematics Specimens (including some types) of Lepiota (abbreviated as L. here) and Leucoagaricus (abbreviated as La. here) from China, Sri Lanka, Europe and North America were examined. As a result, thirty-six species of Lepiota and six species of Leucoagaricus were identified, of which sixteen species and two varieties are new to science, namely, L. ochraceosquamea J. F. Liang & Zhu L. Yang, L. ampliocystidia J. F. Liang & Zhu L. Yang, L. submagnispora J. F. Liang & Zhu L. Yang, L. fusco-brunnea J. F. Liang & Zhu L. Yang, L. contractispora J. F. Liang & Zhu L. Yang, L. subcastanea J. F. Liang & Zhu L. Yang, L. pseudocristata H. C. Wang ex J F. Liang & Zhu L. Yang, L. pallidiochracea J. F. Liang & Zhu L. Yang, L. megaspora J. F. Liang & Zhu L. Yang, L. angusticystidia J. F. Liang & Zhu L. Yang, L. rubro-brunnea H. C. Wang ex J F. Liang & Zhu L. Yang, L. brunneosquamula J. F. Liang & Zhu L. Yang, L. stillispora J. F. Liang & Zhu L. Yang, La. viridianus J. F. Liang & Zhu L. Yang, La. azurescens J. F. Liang & Zhu L. Yang, La. brunneocanus J. F. Liang & Zhu L. Yang, L. clypeolaria var. nigrosquamosa H. C. Wang ex J F. Liang & Zhu L. Yang and L. subcastanea var. bispora J. F. Liang & Zhu L. Yang. Five species, viz. L. aspericeps Murrill, L. kuehneriana Locq., L. brunneolilacea Bon & Boiffard, L. jacobi Vellinga & Knudsen and La. tener (P.D. Orton) Bon, are new to China. Two new combinations, L. cristata var. macrospora (Zhu L. Yang) J. F. Liang & Zhu L. Yang and La. viriditinctus (Berk. & Broome) J. F. Liang & Zhu L. Yang, are proposed. Based on morphological and anatomical characters, Lepiota was divided into five sections, namely, sect. Lepiota, sect. Stenosporae (Lange) Kühner, sect. Ovisporae (Lange) Kühner, sect. Cristatae (Kühner ex Wass.) Bon and sect. Echinatae Fayod. A key to the sections of the genus and keys to the species studied in each section were provided. Additionally, a key to a few species in an allied genus Leucoagaricus was provided. 2. Molecular phylogeny In virtue of some analytic methods such as Most Parsimony analysis, Bayesian analysis, T-PTP (the topology-dependent permutation tail probability) test, PH (the partition homogeneity) test and Genetic distances analysis (HKY85), data sets of ITS, LSU, IGS and mtSSU sequences of Lepiota were analyzed. It was revealed that all five sections based on morphology are polyphyletic, thus the existing infrageneric morphology-based system is imperfect. The evolutiona pileus and basidiospores were speculated. The evolutional approach of pileus might be from trichoderm to hymeniform to inflated elements in chains, while the evolution of basidiospores might be from penguin-shaped or fusiform forms to ellipsoid to triangular or subcylindrical ones. 3. Relationships of morphological and phylogenetic specie Relationships of morphological and phylogenetic species in La. viriditinctus complex were inferred from ITS, LSU, IGS, mtSSU, EF1a and BIP sequences data. The results showed that La. viriditinctus complex consist of five phylogenetic species, and thus, a morphological species might cover one or more phylogenetic species in lepiotaceous fungi. Evolutional rates of ITS, LSU, IGS, mtSSU, EF1a and BIP genes in the genus were compared and founded that the rates of these genes are faster and faster from the first to the last, namely, LSU → mtSSU → ITS →(EF1a, BIP)→ IGS. 4. Population genetics and biogeography of L. cristata Genetic diversity and biogeography of L. cristata were inferred from ITS, IGS and mtSSU sequences. It was revealed that there was sufficient evidence supporting long-distance gene flows among geographic populations and such gene flows are quite common. The gene flow could contribute to the lack of an overall geographic pattern of genetic variation in L. cristata populations. The white basidiospores might be dispersed for long-distance. It was contrary to the former result that white basidiospores were not suitable for long distance dispersal. Population genetics studies revealed a new cryptic species, L. pseudocristata, and a new combination, Lepiota cristata var. macrospora.
页数214
语种中文
文献类型学位论文
条目标识符http://ir.kib.ac.cn/handle/151853/168
专题昆明植物所硕博研究生毕业学位论文
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梁俊峰. 中国环柄菇属的分类学及该属的分子系统学研究--兼论冠状环柄菇的群体遗传学[D]. 昆明植物研究所. 中国科学院昆明植物研究所,2007.
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