星盾炱科、痂囊腕菌科和多腔菌科真菌的分类学研究
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摘要
真菌是自然界中一类重要的微生物资源，在食品生产、新药开发和生物防治等领域，具有显著的生态与经济价值。真菌中的座囊菌纲真菌，是真菌中具有重要经济价值的一类真菌，包括了许多重要的植物病原真菌。其中，痂囊腔菌属可侵染果木表面而形成疮痂状病斑，引起柑橘、葡萄、梨、甘薯、洋扁豆、刀豆等几十种经济林木与农作物致病。
关于座囊菌纲真菌的分类学研究，主要集中在一些新种、新记录种的报道研究上，对各科、属分类地位的描述和总体评价很少，系统集中地研究各个属的模式标本未见报道。模式标本存在很多问题，诸如标本太老、标本已经在二战中毁坏和损坏、其形态特征不明显、有些标本属名种名混乱不清楚等等各种各样的问题。因此，座囊菌纲真菌的资源调查、系统分类和系统发育分析对于座囊菌纲真菌资源的系统收集与保存、物种鉴定及生物多样性研究意义重大。
目前，星盾炱科真菌在座囊菌纲中分类地位尚未确定，痂囊腔菌科和多腔菌科暂被归于座囊菌纲多腔菌目。本文主要对座囊菌纲星盾炱科、痂囊腔菌科和多腔菌科真菌进行了形态分类学研究，结合相关文献的分子生物学研究，依据最新的分类大纲，通过形态显微结构的观察，探讨了星盾炱科、痂囊腔菌科和多腔菌科共52属的分类地位，主要的研究结果和结论如下：
本研究依据盾状囊壳表生，圆形（从外部看）或线性，具辐射状的星形开裂或者狭长的纵裂状开口，从中心孔口到边缘辐射，子囊8-孢型，球形至亚球形，子囊的排列方式是垂直于基部排列，子囊孢子椭圆或圆桶形，透明至棕色，大部分是2-孢型，大部分成熟后变成棕色，将Allothyrium属、Aphanopeltis属、Asterina属、Batistinula属、Cirsosia属、Halbania属、Lembosia属、Meliolaster属、Neostomella属、Parasterinopsis属、Prillieuxina属、Trichasterina属、Uleothyrium属、Viegasia属、Yamamotoa属归于星盾炱科。
依据盾状囊壳具不规则的裂口开裂方式，在寄主表面不形成表生菌丝及附属枝，将Echidnodes属，Lembosina属、Morenoina属、Thyriopsis属归于管腔菌科。
依据子囊果中心处具孔口，盾状囊壳由互相交织的细胞组成，在盾状囊壳内子囊倾斜排列和圆柱形的子囊具短柄，将Anariste属归于微盾壳科。
依据子囊果不具假侧丝，以及亚表皮的子囊果，具圆形的孔口，倒棍棒形的子囊，目前将Lembosiopsis归于球腔菌科。
Asterodothis具星盾炱科真菌的特征，但是其盾状囊壳细胞的排列方式，无色的以及两端细胞不是等球形的子囊孢子，不同于星盾炱科真菌的特征，不能确定将其归于星盾炱科，暂将Asterodothis归于星盾炱科不确定地位的属。
依据其菌丝形成黑色的，圆形的斑块区域和具线形的裂口，大多具碳质化，具多腔盾菌科的特征，暂将Leveillella属、Placosoma属归于多腔盾菌科。
分子生物学研究显示Aulographina属聚在Teratosphaeriaceae科中，依据形态学分类特征结合分子生物学研究，暂将Aulographina属归于Teratosphaeriaceae科。
对于所借到的标本存在形态特征不明显和一些标本的子囊和子囊孢子未成熟，以及有些标本不是模式标本和标本的名称不准确，将Asterolibertia属、Asterotexis属、Dothidasteromella属、Echidnodella属、Eupelte属、Macowaniella属、Parasterinella属、Petrakina属、Placoasterella属、Placoasterina属、Symphaster属、Trichamelia属归于有疑问的属，进一步的分类学研究来确定这些属准确的分类地位。
依据子囊果寄生于叶片，形成病斑和植物炭疽病害，子囊果呈斑点状囊壳，白色偶呈褐色，切片后呈数个子囊腔室散生于子座内，数个子囊散生于每个子囊腔室内；子囊球形至亚球形；子囊孢子无色，砖隔孢子，具横隔膜，将Butleria属、Elsinoe属、Micularia属、Stephanotheca属仍归于痂囊腔科。由于模式标本没有借到，文献中报道Hemimyriangium属和Molleriella属具痂囊腔菌科真菌的典型特征，目前不能确定其准确的分类地位，暂将Hemimyriangium属、Molleriella属归于痂囊腔菌科中有疑问的属。暂将Hyalotheles属归于座囊菌纲不确定地位的属。依据活体寄生于寄主植物，将Beelia属归于刺盾炱科。依据其盾状囊壳具小盾壳科的典型特征，子囊果有辐射状排列的细胞组成，黑色的中心区域是子囊发育部位，将Saccardinula属归于小盾壳科。依据其寄生于植物叶片，基部细胞与寄主细胞融合在一起，梭型至圆柱形的子囊非常特别，将Xenodium属归于有疑问的属。
依据其子座由数个子囊果聚生在一起，呈突出状排列于寄主茎秆上，革质的，具扁平的顶部，子囊球形至亚球形，双囊壁，具细小的短柄或不具短柄，子囊孢子不规则排列，椭圆形至卵形，褐色，砖隔状，将Anhellia属、Myriangium属仍归于多腔菌科。标本没有借到，文献中报道Diplotheca属和Eurytheca属具痂囊腔菌科真菌的典型特征，目前不能确定其准确的分类地位，暂将Diplotheca属、Eurytheca属归于多腔菌科有疑问的属。
新鲜标本采集自高黎贡山国家公园，依据表生菌丝形成叶菌体，呈喷泉状排列和其子囊果形状类似铜钱币状，中心具孔口，重寄生在其它的子囊菌真菌中，菌丝通常会感染寄生的真菌和地衣形成叶菌体，结合基因测序分析结果，暂时将此新鲜标本鉴定为Trichothyrium sp.。依据子囊果具Y-形裂口，子囊亚球形至椭圆形，符合Lembosiella属的特征，所采集到的标本寄主为Menispermacea，与模式种的寄主不同，由于Lembosiella属是单模式属，对于采集到的标本进行形态学研究，其特征与模式种Lembosiella Polyspora(Pat.) Sacc.,的特征相似，而采集的标本寄主植物不同于模式种的寄主植物，将两份新鲜标本鉴定为Lembosiella sp.和Lembosiella sp.。依据子囊果具线状裂口，子囊球形至亚球形，独特之处在于子囊遇碘化钾试剂变蓝，暂时将此新鲜标本鉴定为Aulographum sp.。
Fungi is an ecological group, with high diversity. Fungi palys an important role inecosystem. They can decompose organinc substrata in the environment. Dothideomycetes is alarge class within fungi, which causes a lot of plant disease. Fungi have been classifiedaccoding to morphological characters for a long time; and few taxa have been classfied relyingon molecular data.
There are some problems in type species: such as specimen old, specimen in badcondition, the detail of specimen unequal to the morphological character, the latin nameunfixed by the scientist.
Asterinacee family belongs to Dothideomycetes uncertain family. Elsinoaceae andMyringiaceae family belong to the order Myriangiales.
Based on the examinations of generic types of5families of Asterinaceae、Elsinoaceae andMyriangiaceae, we revised these3families taxonomy based on the morphological characters.The conclusion is as follows:
The family Asterinaceae are reduced into fifteen genus: Allothyrium, Aphanopeltis,Asterina, Batistinula, Cirsosia, Halbania, Lembosia, Meliolaster, Neostomella, Parasterinopsis,Prillieuxina, Trichasterina, Uleothyrium, Viegasia, Yamamotoa; based on the character ofthyriothecium with linear or stellate fissures, asci8-spored, subglobose to globose, andascospores hyaline, brown when mature.
The four genus are transferred into the family Aulographaceae: Echidnodes, Lembosina,Morenoina, Thyriopsis, based on the characters of thyriotheciumwith large fissures, unforminghyphopodia on the mycelium.
The genus Anariste are transferred into the family Micropeltidaceae; based on thecharacter of thyriothecium with ostiolate, composed of cells complex together and with asciclined inside thyriothecium.
The genus Lembosiopsis are transferred into the family Mycosphaerellaceae;
The genus Asterodothis are placed into the doubtful genus within Asterinaceae.
The two genus are placed into the family Parmulariaceae: Leveillella, Placosoma; thegenus Aulographina are transferred into the family Teratosphaeriaceae, with character of themycelium composed black areas on the surface and hardly removed from host surface and withhyphopodia absent.
The twelve genus are placed into doubtful genus: Asterolibertia, Asterotexis,Dothidasteromella, Echidnodella, Eupelte, Macowaniella, Parasterinella, Petrakina,Placoasterella, Placoasterina, Symphaster, Trichamelia, based on specimen in bad condition,specimen unloaned from herbarium, morphological chararacter unequall to the literaturedescription.
The family Elsinoaceae reduced to four genus: Butlreia, Elsinoe, Micularia,Stephanotheca; based ont the morpholgical character of forming scab on the host surface, asciglobose to subglobose, ascospres hyaline to brown, occasionally muriform. The two genus areplaced into doubrful genus: Hemimyriangium, Molleriella. The two genus are transferred intoDothideomycetes incertae sedis: Hyalotheles, Xenodium; the genus Beelia are transferred intoChaetothyriaceae; the genus Saccardinula are transferred into Microthyriaceae. Xenodium isplaced into doubtful gnera, based on the asci cylindrical, ascospore cylindrical and themorphological character could not included into any familyof Dothideomycetes.
The family Myriangiaceae reduced into two genus: Anhellia, Myriangium, with charactersof the ascostromata composed of multi-locule, erumpent to the host surface with narrow upside,asci globose to subglobose, ascospores irregularly arranged inside asci, brown and muriform.The two genus Diplotheca, Eurytheca are placed into doubtful genus within Myriangiaceae,based on the specimen unloaned from herbarium.
Four fresh collection were obtained from Gaoligonshan in Yunnan province. Based on themorphological characters with molecular data, we collected four specimens and identified asLembosiella sp., Lembosiella sp., Aulographum sp. and Trichothyrium sp. The host of the typespecies Lembosiella is different from the host of our collection, based on this, we identified as uncertain species. Aulographum sp is unique because being blue when dyed in Melzer’sreagent.
关于座囊菌纲真菌的分类学研究，主要集中在一些新种、新记录种的报道研究上，对各科、属分类地位的描述和总体评价很少，系统集中地研究各个属的模式标本未见报道。模式标本存在很多问题，诸如标本太老、标本已经在二战中毁坏和损坏、其形态特征不明显、有些标本属名种名混乱不清楚等等各种各样的问题。因此，座囊菌纲真菌的资源调查、系统分类和系统发育分析对于座囊菌纲真菌资源的系统收集与保存、物种鉴定及生物多样性研究意义重大。
目前，星盾炱科真菌在座囊菌纲中分类地位尚未确定，痂囊腔菌科和多腔菌科暂被归于座囊菌纲多腔菌目。本文主要对座囊菌纲星盾炱科、痂囊腔菌科和多腔菌科真菌进行了形态分类学研究，结合相关文献的分子生物学研究，依据最新的分类大纲，通过形态显微结构的观察，探讨了星盾炱科、痂囊腔菌科和多腔菌科共52属的分类地位，主要的研究结果和结论如下：
本研究依据盾状囊壳表生，圆形（从外部看）或线性，具辐射状的星形开裂或者狭长的纵裂状开口，从中心孔口到边缘辐射，子囊8-孢型，球形至亚球形，子囊的排列方式是垂直于基部排列，子囊孢子椭圆或圆桶形，透明至棕色，大部分是2-孢型，大部分成熟后变成棕色，将Allothyrium属、Aphanopeltis属、Asterina属、Batistinula属、Cirsosia属、Halbania属、Lembosia属、Meliolaster属、Neostomella属、Parasterinopsis属、Prillieuxina属、Trichasterina属、Uleothyrium属、Viegasia属、Yamamotoa属归于星盾炱科。
依据盾状囊壳具不规则的裂口开裂方式，在寄主表面不形成表生菌丝及附属枝，将Echidnodes属，Lembosina属、Morenoina属、Thyriopsis属归于管腔菌科。
依据子囊果中心处具孔口，盾状囊壳由互相交织的细胞组成，在盾状囊壳内子囊倾斜排列和圆柱形的子囊具短柄，将Anariste属归于微盾壳科。
依据子囊果不具假侧丝，以及亚表皮的子囊果，具圆形的孔口，倒棍棒形的子囊，目前将Lembosiopsis归于球腔菌科。
Asterodothis具星盾炱科真菌的特征，但是其盾状囊壳细胞的排列方式，无色的以及两端细胞不是等球形的子囊孢子，不同于星盾炱科真菌的特征，不能确定将其归于星盾炱科，暂将Asterodothis归于星盾炱科不确定地位的属。
依据其菌丝形成黑色的，圆形的斑块区域和具线形的裂口，大多具碳质化，具多腔盾菌科的特征，暂将Leveillella属、Placosoma属归于多腔盾菌科。
分子生物学研究显示Aulographina属聚在Teratosphaeriaceae科中，依据形态学分类特征结合分子生物学研究，暂将Aulographina属归于Teratosphaeriaceae科。
对于所借到的标本存在形态特征不明显和一些标本的子囊和子囊孢子未成熟，以及有些标本不是模式标本和标本的名称不准确，将Asterolibertia属、Asterotexis属、Dothidasteromella属、Echidnodella属、Eupelte属、Macowaniella属、Parasterinella属、Petrakina属、Placoasterella属、Placoasterina属、Symphaster属、Trichamelia属归于有疑问的属，进一步的分类学研究来确定这些属准确的分类地位。
依据子囊果寄生于叶片，形成病斑和植物炭疽病害，子囊果呈斑点状囊壳，白色偶呈褐色，切片后呈数个子囊腔室散生于子座内，数个子囊散生于每个子囊腔室内；子囊球形至亚球形；子囊孢子无色，砖隔孢子，具横隔膜，将Butleria属、Elsinoe属、Micularia属、Stephanotheca属仍归于痂囊腔科。由于模式标本没有借到，文献中报道Hemimyriangium属和Molleriella属具痂囊腔菌科真菌的典型特征，目前不能确定其准确的分类地位，暂将Hemimyriangium属、Molleriella属归于痂囊腔菌科中有疑问的属。暂将Hyalotheles属归于座囊菌纲不确定地位的属。依据活体寄生于寄主植物，将Beelia属归于刺盾炱科。依据其盾状囊壳具小盾壳科的典型特征，子囊果有辐射状排列的细胞组成，黑色的中心区域是子囊发育部位，将Saccardinula属归于小盾壳科。依据其寄生于植物叶片，基部细胞与寄主细胞融合在一起，梭型至圆柱形的子囊非常特别，将Xenodium属归于有疑问的属。
依据其子座由数个子囊果聚生在一起，呈突出状排列于寄主茎秆上，革质的，具扁平的顶部，子囊球形至亚球形，双囊壁，具细小的短柄或不具短柄，子囊孢子不规则排列，椭圆形至卵形，褐色，砖隔状，将Anhellia属、Myriangium属仍归于多腔菌科。标本没有借到，文献中报道Diplotheca属和Eurytheca属具痂囊腔菌科真菌的典型特征，目前不能确定其准确的分类地位，暂将Diplotheca属、Eurytheca属归于多腔菌科有疑问的属。
新鲜标本采集自高黎贡山国家公园，依据表生菌丝形成叶菌体，呈喷泉状排列和其子囊果形状类似铜钱币状，中心具孔口，重寄生在其它的子囊菌真菌中，菌丝通常会感染寄生的真菌和地衣形成叶菌体，结合基因测序分析结果，暂时将此新鲜标本鉴定为Trichothyrium sp.。依据子囊果具Y-形裂口，子囊亚球形至椭圆形，符合Lembosiella属的特征，所采集到的标本寄主为Menispermacea，与模式种的寄主不同，由于Lembosiella属是单模式属，对于采集到的标本进行形态学研究，其特征与模式种Lembosiella Polyspora(Pat.) Sacc.,的特征相似，而采集的标本寄主植物不同于模式种的寄主植物，将两份新鲜标本鉴定为Lembosiella sp.和Lembosiella sp.。依据子囊果具线状裂口，子囊球形至亚球形，独特之处在于子囊遇碘化钾试剂变蓝，暂时将此新鲜标本鉴定为Aulographum sp.。
Fungi is an ecological group, with high diversity. Fungi palys an important role inecosystem. They can decompose organinc substrata in the environment. Dothideomycetes is alarge class within fungi, which causes a lot of plant disease. Fungi have been classifiedaccoding to morphological characters for a long time; and few taxa have been classfied relyingon molecular data.
There are some problems in type species: such as specimen old, specimen in badcondition, the detail of specimen unequal to the morphological character, the latin nameunfixed by the scientist.
Asterinacee family belongs to Dothideomycetes uncertain family. Elsinoaceae andMyringiaceae family belong to the order Myriangiales.
Based on the examinations of generic types of5families of Asterinaceae、Elsinoaceae andMyriangiaceae, we revised these3families taxonomy based on the morphological characters.The conclusion is as follows:
The family Asterinaceae are reduced into fifteen genus: Allothyrium, Aphanopeltis,Asterina, Batistinula, Cirsosia, Halbania, Lembosia, Meliolaster, Neostomella, Parasterinopsis,Prillieuxina, Trichasterina, Uleothyrium, Viegasia, Yamamotoa; based on the character ofthyriothecium with linear or stellate fissures, asci8-spored, subglobose to globose, andascospores hyaline, brown when mature.
The four genus are transferred into the family Aulographaceae: Echidnodes, Lembosina,Morenoina, Thyriopsis, based on the characters of thyriotheciumwith large fissures, unforminghyphopodia on the mycelium.
The genus Anariste are transferred into the family Micropeltidaceae; based on thecharacter of thyriothecium with ostiolate, composed of cells complex together and with asciclined inside thyriothecium.
The genus Lembosiopsis are transferred into the family Mycosphaerellaceae;
The genus Asterodothis are placed into the doubtful genus within Asterinaceae.
The two genus are placed into the family Parmulariaceae: Leveillella, Placosoma; thegenus Aulographina are transferred into the family Teratosphaeriaceae, with character of themycelium composed black areas on the surface and hardly removed from host surface and withhyphopodia absent.
The twelve genus are placed into doubtful genus: Asterolibertia, Asterotexis,Dothidasteromella, Echidnodella, Eupelte, Macowaniella, Parasterinella, Petrakina,Placoasterella, Placoasterina, Symphaster, Trichamelia, based on specimen in bad condition,specimen unloaned from herbarium, morphological chararacter unequall to the literaturedescription.
The family Elsinoaceae reduced to four genus: Butlreia, Elsinoe, Micularia,Stephanotheca; based ont the morpholgical character of forming scab on the host surface, asciglobose to subglobose, ascospres hyaline to brown, occasionally muriform. The two genus areplaced into doubrful genus: Hemimyriangium, Molleriella. The two genus are transferred intoDothideomycetes incertae sedis: Hyalotheles, Xenodium; the genus Beelia are transferred intoChaetothyriaceae; the genus Saccardinula are transferred into Microthyriaceae. Xenodium isplaced into doubtful gnera, based on the asci cylindrical, ascospore cylindrical and themorphological character could not included into any familyof Dothideomycetes.
The family Myriangiaceae reduced into two genus: Anhellia, Myriangium, with charactersof the ascostromata composed of multi-locule, erumpent to the host surface with narrow upside,asci globose to subglobose, ascospores irregularly arranged inside asci, brown and muriform.The two genus Diplotheca, Eurytheca are placed into doubtful genus within Myriangiaceae,based on the specimen unloaned from herbarium.
Four fresh collection were obtained from Gaoligonshan in Yunnan province. Based on themorphological characters with molecular data, we collected four specimens and identified asLembosiella sp., Lembosiella sp., Aulographum sp. and Trichothyrium sp. The host of the typespecies Lembosiella is different from the host of our collection, based on this, we identified as uncertain species. Aulographum sp is unique because being blue when dyed in Melzer’sreagent.
引文
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