中国杯伞科的系统发育与分类研究

	中国杯伞科的系统发育与分类研究
	何正蜜
导师	杨祝良
关键词	广义杯伞，杯伞科，金钱菌属，单拷贝同源直系基因，毒蕈碱 Clitocybe s.l., Clitocybaceae, Collybia, single-copy gene, muscarine
摘要	杯伞科Clitocybaceae隶属于蘑菇目Agaricales，子实体大多呈杯伞状，具有重要的生态和经济价值。该科真菌物种多样，分布广泛，但种间亲缘关系复杂不明，整个系统发育框架需重建，各个种的形态特征不易把握，是蘑菇目中分类最为困难的类群之一。该科既有常见的食用菌，也有含神经毒素毒蕈碱的毒菌，但哪些种含毒蕈碱，它们在该科中的系统位置如何等问题同样有待解决。本研究基于分子系统发育、外观形态、显微结构、生境生态和毒素成分对中国杯伞展开了研究，主要结果如下： 1. 基于ITS、多基因ITS-LSU-TEF1-RPB1-RPB2-ATP6以及全基因组单拷贝同源直系基因序列分析，对杯伞科进行了系统发育重建，结果表明：① 杯伞科为单系类群，包括9个属级分支：金钱菌属Collybia（重新界定）、白丽蘑属Leucocalocybe（重新界定）、大孢杯伞属Macrosporocybe（新拟）、肉色香蘑属Carneolepista（新拟）、假离褶伞属Pseudolyophyllum（亚属升属）、狭义香蘑属Lepista s.s.、树状金钱菌属Dendrocollybia、囊泡杯伞属Singerocybe和狭义杯伞属Clitocybe s.s.。本研究对以上属进行了描述并编写了该科分属检索表；② 金钱菌属划分为8个Clade，孢子的聚离、生长基质和毒蕈碱的有无可作为各Clade的区分特征；③ 在本研究中，杯伞科有62个种级分支，其中本研究建立了15个新种（金钱菌属11种，杯伞属2种，大孢杯伞属和假离褶伞属各1种），新组合17种（金钱菌属6种，白丽蘑属5种，假离褶伞属5种，肉色香蘑属1种），未命名金钱菌2种，并提供了我国金钱菌的分种检索表。 2. LC-MS/MS检测分析表明：① 在杯伞科中，毒蕈碱毒素仅分布于金钱菌属中；② 在金钱菌属中，仅位于基部的蓝绿金钱菌C. odora（Clade VIII）不含毒蕈碱；③ 毒蕈碱在金钱菌属中呈单系分布，为有毒金钱菌的一个共衍征。 3. 研究发现，有的杯伞状真菌与杯伞科成员并无亲缘关系。根据外观形态、显微结构观察及多基因分子系统发育分析所得证据，① 建立了蜡伞科Hygrophoraceae的一个新属——灰盖杯伞属Spodocybe，发表新种皱灰盖杯伞S. rugosiceps和双孢灰盖杯伞S. bispora；② 明确了蜡伞科的单系性，提出了一个新的亚科——拱顶菇亚科Cuphophylloideae，该亚科包括4个属，分别为拱顶菇属Cuphophyllus、棒柄杯伞属Ampulloclitocybe、喙囊圆顶伞属Cantharocybe和灰盖杯伞属。 4. 基于形态学和分子系统学证据，本文还研究了其他4属子实体为杯伞状的真菌。① 报道了我国邦氏杯伞属Bonomyces的1个新种（细鳞邦氏杯伞B. squamulosus）和哈氏杯伞属Harmajaea的1个新记录种（哈珀杯伞H. harperi），此二属隶属于假杯伞科Pseudoclitocybaceae；② 描述了我国漏斗伞属Infundibulicybe的7个种，含2个新种（双孢漏斗伞I. bispora和椭孢漏斗伞I. ellipsospora），并提供了该属分种检索表；③ 报道了我国假香蘑属Notholepista的1个新种（空柄假香蘑N. fistulosa），此属科级系统位置不清，有待继续研究。; The family Clitocybaceae in order Agaricales has both important ecological and economic values, whose fruitbodies are mostly clitocyboid. This family has a high species diversity and wide distribution, but it is one of the most difficult taxa to be classified in Agaricales because it has complex interspecific phylogenetic relationships and untypical interspecific morphological characteristics, and its whole phylogenetic framework of the family needs to be rebuilt. In this family, there are common edible fungus, but also neurotoxic fungus containing muscarine. The problems which species contain muscarine and where are their phylogenetic positions in the family are also needed to be resolved. In this context, our study on the taxonomy of clitocyboids is carried out on the basis of macromorphology, microstructure, molecular phylogeny and toxin. The main results are as follows: 1. The phylogenetic analyse based on ITS, multi-gene ITS-LSU-TEF1-RPB1-RPB2-ATP6 and genomic single-copy genes suggest that (i) the family Clitocybaceae is monophyletic, consisted of emended Collybia and Leucocalocybe, Macrosporocybe gen. nov., Carneolepista gen. nov., Pseudolyophyllum stat. nov., Lepista s.s, Dendrocollybia, Singerocybe and Clitocybe s.s., and a key to the genera in family Clitocybaceae is provided; (ii) The genus Collybia is divided into eight clades and the conditions of spores (single or in tetrad), substrates and presence and absence of muscarine can be the diagnostic features for clades; (iii) 62 specific branches are found, of which 11 Collybia, 2 Clitocybe, 1 Macrosporocybe and 1 Pseudolyophyllum are new species, 6 Collybia, 5 Leucocalocybe, 5 Pseudolyophyllum and 1 Carneolepista are new combination, 2 Collybia are unnamed speice; besides, a key to Chinese Collybia spp. is provided. 2. The LC-MS/MS detection shows that (i) in Clitocybaceae, muscarine is only distributed in genus Collybia; (ii) in Collybia, only the basal species C. odora (Clade VIII) does not produce muscarine; (iii) muscarine-producing Collybia spp. cluster in a monophyletic clade, indicating that muscarine is a synapomorphy of poisonous collybias. 3. According to research findings, some clitocyboid fungi are unrelated with Clitocybaceae. Using evidences of macromorphology, microstructure and molecular phylogeny, (i) a new genus Spodocybe and two new species in family Hygrophoraceae, namely S. rugosiceps and S. bispora, are established; (ii) the monophyly of Hygrophoraceae is confirmed and a new subfamily made up of Cuphophyllus, Ampulloclitocybe, Cantharocybeis and Spodocybe is proposed. 4. Based on mophylogical and phylogenetic evidence, another four clitocyboid genera has also been studied. (i) Bonomyces squamulosus sp. nov. is proposed and Chinese Harmajaea harperi is reported for the first time, which are both placed in Pseudoclitocybaceae. (ii) Seven Infundibulicybe species are described, including two new species, namely I. bispora and I. ellipsospora, and a key to Chinese Infu
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/151853/75102
专题	昆明植物所硕博研究生毕业学位论文
推荐引用方式 GB/T 7714	何正蜜. 中国杯伞科的系统发育与分类研究[D]. 中国科学院大学,2022.

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