摘要
本研究分为两大部分。第一部分采用现代真菌分类学技术，对源于中国华南地区的斑痣盘菌目（Rhytismatales）菌物进行了物种多样性的研究，从形态解剖学特征、个体发育学、地理分布、生态生境等多个方面入手，对这一地区该目成员进行了较为系统全面的研究。第二章运用分子生物学技术，对采自中国本土的斑痣盘菌目成员进行系统发育学研究，主要基于rDNA-ITS和nLSU rDNA单独序列和双基因联合序列构建斑痣盘菌目系统发育树来探讨该目科间、属间、种间及种内的亲缘关系。
通过对自行采集和安徽农业大学森林菌物标本室（AAUF）保存的华南地区干标本进行逐号检查，共获斑痣盘菌标本70余号，涉及寄主近40种。基于Darke（r1967）、Cannon&Minter（1983，1986）、Johnston（1986）、Spooner（1991）、Kirk et al.（2001、2008）等国外知名真菌学家提出的相关分类原则，将所有标本进行鉴定与整理，共查明斑痣盘菌目1科9属29种，其中包括新种3个，分别为银杉散斑壳（Lophodermium cathayae）、简丝特里尔盘菌（Terriera simple）和络石特里尔盘菌（T. trachelosperi）；中国新纪录种1个，即杜鹃隔孢缝壳（Lophomerum rhododendri）；中国大陆新纪录种1个，为短特里尔盘菌（Terriera brevis）。对3新种作了详尽的形态学描述并提供了点线合成图或显微照片；对两个新纪录种绘制了子囊、子囊孢子及侧丝的点线合成图；对其余各种作了简要的形态学特征的描述与讨论。同时，编制了中国华南地区斑痣盘菌目分种检索表，以便于简明扼要地区分该区各种。
在对华南地区斑痣盘菌目成员物种多样性形成的原因分析中，主要从寄主、个体发育特征和地理分布等三个方面展开。结果表明源于华南地区的29种斑痣盘菌寄主范围广泛，涉及多种植物，其中以裸子植物居多，特别是松属（Pinus）植物，本次查明的松树上共有9种斑痣盘菌。有些种有较广泛的寄主范围，具有较弱的寄生性；但也有些种具有较强的寄主专化性，甚至只寄生于一种植物上，如银杉散斑壳（L.cathayae）。从真菌保护学意义上来说，银杉散斑壳虽然是导致银杉落针病的病原菌，但是它也很可能会随着“活化石”——银杉（Cathaya argyrophylla）的灭绝而消失，是世界菌物库的损失。同时，地理环境对于斑痣盘菌个体差异性也有很大的影响。在华南地区采集的这29个种中，大部分种分布范围广泛，在世界大多数温带和热带地区均有分布；但也存在目前仅报道于中国华南地区的斑痣盘菌，如银杉散斑壳（L.cathayae）和广西散斑壳（L. guangxiense）。另外，斑痣盘菌科（Rhytismataceae）是该区的优势科，其中散斑壳属、齿裂菌属和特里尔盘菌属是该区的优势属。各属、种阶元的分类主要依据子囊果外部形态和内部结构的特征，如子囊果、子囊、子囊孢子的形状，侧丝顶端的形状，子座覆盖层与基部层毗邻处的组织结构，唇细胞的有无以及分生孢子器、分生孢子的形状和线纹多少等。
第二部分对来源于全国各地，特别是长江流域及其以南地区的斑痣盘菌目成员进行了系统发育学的研究。采用CTAB法提取供试菌株的DNA，通过PCR扩增及测序的方法，基于rDNA-ITS序列、28S rDNA序列和二者的联合序列，以柔膜菌目的Leotialubrica为外群，分别构建了斑痣盘菌目系统发育树；并根据rDNA-ITS序列分析对银杉散斑壳进行了分子确认。结果表明，在斑痣盘菌目中，同一科的种、同一属的种、同一个种通常会优先聚在一起，并具有较高的自举支持率。从而验证了利用基于表型性状的现代菌物分类技术对斑痣盘菌目成员进行的分类具有重要的意义和很高的参考价值。但是，也有个别在表型上分属于两个不同属、甚至不同科的种被聚在了一起，说明利用表型分类技术获得的分类研究结果不一定都是绝对可靠的，有时必须结合分子生物学技术对部分类群作进一步鉴定与分类。
研究结果还表明，寄主、地理分布、海拔高度、气候环境等因素与斑痣盘菌目成员之间的亲缘关系研究有很大的关联性。大部分寄生在针叶树和阔叶树上的斑痣盘菌科成员与生长于落叶层或林地上的地锤菌科（Cudoniaceae）成员总是分属两大分支，尤其在ITS+28S rDNA的联合序列构建的系统发育树中二科的界限更为明晰。在基于28S rDNA序列建立的系统发育树中，源自于国外（欧洲、北美、非洲）的斑痣盘菌形成了一个独立的分支。在rDNA-ITS序列构建的系统发育树中，有一分支聚集的95%以上的Rhytismatales成员均来自于湖北神农架；而在ITS+28S rDNA的联合序列构建的系统发育树中，采自于安徽黄山或者天堂寨的悬钩子皮下盘菌（Hypoderma rubi）和采自湖北神农架的H. rubi无法聚在一起。因此，我们认为，在研究菌物系统发育学的过程中，仅仅依赖双基因甚至更多基因联合序列的分析还是不够的，只有结合菌物形态学、个体发育、培养性状、寄主（基物）范围及地理分布、生境及获养方式等表型性状进行全面地综合分析，才能获得较为科学、合理的研究结果。
This study includes two parts. The species diversity of Rhytismatales from South Chinawas researched systematically and comprehensively in the aspects of morphological andanatomical characters, ontogeny, geographic distribution and ecology using moderntechniques for fungal taxonomy in the first part. Based on rDNA-ITS, nLSU rDNAsequences and their joint sequences, phylogeny of Rhytismatales members from nativeplants was researched using molecular biotechnology and the relationship of interfamily,intergenera, interspecies and intraspecies were explored by establishing phylogenetic treesin the second part.
More than70specimens which were collected from South China and deposited inthe Reference Collection of Forest Fungi of Anhui Agricultural University, China (AAUF)were examined. According to the taxonomic principles by Darker (1967), Cannon&Minter (1983,1986), Johnston (1986), Spooner (1991), Kirk et al.(2001),1family9genera29species of Rhytismatales were identified. Among them, Lophodermiumcathayae, Terriera simplex and T. trachelosperi were new species; Lophomerumrhododendri was a new record to China; Terriera brevis was a new record to Chinesemainland. Morphologic descriptions and point and line integrated illustrations ormicrograph were given for3new species; illustrations of asci, ascospores and paraphyseswere provided for2newly recorded species; and simple descriptions and discussions ofmorphologic characteristics were given for other species. Keys were compiled foridentification of all species.
The reasons of species diversity of Rhytismatales were analyzed in3aspects: host,the characteristics of ontogeny and distributions. The results showed that: the host rangesof Rhytismatales were broad that involved many kinds of plants. Among these plants,gymnosperm was predominant, especially Pinus sp. And in this study,9species ofRhytismatales were collected on pines. However, there were some species having hostspecificity, for example L. cathayae. In a sense of fungal conservation, though L. cathayaecaused the needle blight of Cathaya argyrophylla, this pathogen would disappear followingthe extinction of the “living fossil”, which would be the loss of fungus resource in theworld. Meanwhile, geography had a great effect on the phylogeny of Rhytismatales. Mostof the species being collected in South China had a wide distribution, but there were alsosome species only being seen in South China, for example, L. cathayae and L. guangxiense. In addition, Rhytismataceae was dominating family in this area; and Lophodermium,Coccomyces and Terriera were dominating genera. The identifications of intergenera orinterspecies were showed based on the features of external morphology and internalstructures of ascoma, such as shapes of ascoma, asci, ascospore, paraphyses tips, texturalstructures between covering and basal stroma, absence or presence of lip cells, shapes ofconidioma and conidia, quantity of zone lines and so on.
In the second part, the phylogeny of Rhytismatales from various areas of China，especially Yangtze River and its south region was studied. Genomic DNA from isolates oftested strains was extracted from mycelia by the method of CTAB. ITS and28S regionswere amplified by the polymerase chain reaction (PCR). The phylogenetic trees wereestablished based on the sequences of rDNA-ITS, nLSU rDNA and ITS+28S rDNA,selecting Leotia lubrica as outgroup. The classification status of L. cathayae wasconfirmed based on some sequences of rDNA-ITS. The results showed that: species in thesame family or genus and the same species were usually clustered into a group with strongbootstraps, which verified the modern techniques based on phenotypic traits for fungaltaxonomy had important significance and high reference value to the classification ofRhytismatales. However, some species being placed in different genera, even differentfamily with different morphology could also be clustered together, which showed that thetaxonomic results based on classical classification may be limited, and must be combinedwith molecular biotechnology.
From the phylogenetic trees, we know that host, geography, altitude, climate andenvironment were related to the relationship of Rhytismatales. Rhytismataceae on conifersand broad leaves and Cudoniaceae on duff were separated into two clades, especially in thetree based on ITS+28S rDNA sequences. In28S-rDNA tree, species from abroad wereclustered together. In rDNA-ITS tree, one group included R. members, of which beyond95%from Shennongjia. In the two-gene phylogenetic tree, Hypoderma rubi fromShennongjia (Hubei Province) couldn’t be clustered with the same species fromHuangshan Scenic Spot and Tiantangzhai (Anhui Province). As a result, it’s not enough tostudy the phylogeny of fungus only by the comprehensive analysis based on two-ormulti-gene. The scientific and reasonable results will be achieved if combining molecularbiotechnology with phenotypic traits of fungi.
引文
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