云南地区烟煤菌形态学与分子系统学的研究
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摘要
烟煤菌是指一类具有暗色菌丝及孢子，以蚜虫、粉虱、粉蚧等害虫分泌的蜜露或特定植物叶片分泌的溢泌物为营养的真菌。烟煤菌对寄主植物不直接产生危害，但是它覆盖在叶、枝、花、果等植物组织的表面，影响植物的光合作用、观赏功能以及花、果的品质，降低其商业价值。烟煤菌可以食用，生产氯化物过氧化物酶和细胞壁降解酶，次生代谢物具有较好的抗肿瘤，抗菌活性成分，可以为生物制药和生物农药的研发奠定基础。资源的调查和系统分类研究是防治烟煤菌和利用烟煤菌的基础。云南是动物、植物的王国，生物多样性极其丰富，但烟煤菌系统的资源调查和分类学研究还未见报道。本研究的目的是调查云南烟煤菌的资源，研究烟煤菌的形态和分子系统学，为烟煤菌的防治和利用提供科学依据和理论基础。
本研究从云南27个州县上调查采集烟煤菌资源，鉴定并分离培养烟煤菌的种类；通过ITS、18S和28S序列的分析，构建系统发育树，使烟煤菌的分类系统更趋于自然的分类系统，主要结果如下：
1、采集样本449份，保存干标本96份、分离获得纯培养物62份，鉴定出真菌属于8个属，24种。
2、对烟煤菌做了系统的形态学和分子系统学研究，列出了煤炱科、刺盾炱科和Trichomeriaceae烟煤菌所在属的属征，并详细描述了所采集到的种类。联合ITS、18S和28S序列建立系统发育树，结合形态特征，分别讨论了三个科系统发育及分类地位。
3、描述了烟煤菌14个新种和4个新纪录种，其中煤炱科四个新种：Leptoxyphiumglochidion，Capnodium nonturgidum，C. lagerstroemiae，Phragmocapnias chinensis；3个新纪录种：L. kurandae，P. siamensis，P. Asiticus。刺盾炱科6个新种：Phaeosaccardinulamultiseptata，Ph. dendrocalami，Ph. yunnanensis，Ph. dalbergiae，Cyphellophora schimae，Cy. jingdongensis；1个新纪录种：Ceramothyrium thailandicum。Trichomeriaceae的4个新种：Trichomerium yunnanense，T. multisetariae，T. syzygii，T. Murrayae，通过形态学和分子系统学研究确立了这些种的分类地位以及与其相似种的区别。
4、修订了Capnodium. citri的异名：现在分类学上把C. salicinum作为C. citri的异名，以C. citri作为现在的名字（Index Fungorum2014），但是根据本研究系统发育树来看，两者的亲缘关系较远，并不能作为异名使用。
5、增加了烟煤菌的形态特征：补充了煤炱科煤炱属子囊果具刚毛；发现了胶煤炱属无性型的分生孢子器不只是基部膨大，还有中上部膨大的形态出现，海绵胶煤炱两种形态的出现是多基因和环境共同作用的结果。
6、首次发现C. coffeae和Cyphellophora的有性型：通过分子生物学上的证据，清楚了C. coffeae的有性型和无性型的形态特征，将C. coffeae的有性型和无性型很好的联系起来；描述了Cyphellophora属下3个种的有性型特征，清楚了Cy. hylomeconis的有性型和无性型之间的联系。
7、通过对449份样本寄主植物种类的研究以及紫胶虫寄主植物上烟煤菌种类的分析，发现寄主植物的种类对烟煤菌的种类影响不大，初步推测烟煤菌的种类取决于蜜露的种类（即昆虫的种类）和环境因素共同作用。
本文结合分子系统学和形态学首次对云南烟煤菌做了系统的研究，描述了云南烟煤菌特征并构建了分类系统，对一些分类地位不清楚的种类和形态上难以区别的种类，提供了清晰的分子生物学证据，为烟煤菌鉴定和系统学研究提供了的重要科学依据。研究结果表明，烟煤菌在云南多样性极其丰富，还有很多的新种和新纪录种没有被发现，烟煤菌的分子数据还很欠缺，需要更多的分子资料结合形态特征进行研究。
Sooty mould is fungus with black hypha or hyphopodia and feeding on honeydew that isexcreted from sap-sucking insects like aphids, whiteflies and mealy bugs. Sooty mould iseatable and producing the industrial chloroperoxidase, pectinases and cellulases. The secondarymetabolite of sooty mould has better anti-cancer and antibacterial components that are thefoundation for researching the pharmaceuticals and biological pesticide. Researching theresource and systematic classification is the foundation of controlling and utilizing sooty mold.Yunnan is the kingdom of the animal and plant, with rich biological diversity, but investigationof sooty mould’s system resources and identification have not been reported. The purpose ofthis study is to investigate the resources of sooty mould, to research the morphology andmolecular phylogeny of sooty mould, to provide the scientific basis and material foundation forthe controlling and utilization of sooty mould.
Samples from27states of Yunnan Province were investigated and collected, identifiedand isolated. The natural classification system of sooty mould was achieved through thephylogenetic tree based on ITS,18S and28S sequences. The main results in this study arelisted below.
1.449samples were collected,96dry samples were deposited,62sooty mould strainswere isolated,24species belonging to8genera were identified.
2. the systemic morphology and molecular phylogeny of sooty mould, and themorphological characters of all genera belonging to Capnodiaceae, Chaetothyriaceae andTrichomeriaceae were listed. All species were descripted detailedly. The phylogenetic treebased on ITS,18S and28S sequences combined with morphological characteristics werediscussed taxonomic status of three families.
3.14new species and4new records of sooty mould were collected and described in thisstudy.4new species belonging to Capnodiaceae, i.e Leptoxyphium glochidion，Capnodium nonturgidum，C. lagerstroemiae，Phragmocapnias chinensis and3new records belonging tothe same family, i.e L. kurandae，P. siamensis，P. asiticus.6new species belonging toChaetothyriaceae, i.e Phaeosaccardinula multiseptata，Ph. dendrocalami，Ph. yunnanensis，Ph. dalbergiae，Cyphellophora schimae，Cy. Jingdongensis and Ceramothyrium thailandicumwas new records belonging to the same family.4new species belonging to Trichomeriaceae, i.eTrichomerium yunnanense， T. multisetariae， T. syzygii， T. Murrayae. The systematicsplacement and differences among similar species were discussed according to the results ofmolecular data and morphological character.
4. Revised the synonym of Capnodium. citri. Now C. salicinum was as a synonym of C.ciiri on the taxonomy and C. citri was the current name. Based on the phylogenetic tree of thisstudy, the relationship of two species was more distant and C. salicinum could not be asynonym of C. citri.
5. Morphological studys shows: the ascomata of Capndium with setae, and the pycnidiumof Scorias spongiosa was not only swells on the base, but also swells on the upper part.
6. This study firstly reported the telemorph of C. coffeae and Cyphellophora. Through thephylogenetic analysis, the morphological characteristics of C. coffeae teleomorph andanamorph were linked together. Sexual characteristics of3species belonging to Cyphellophorawere descripted, the sexual and asexual characteristics of Cy. hylomeconis were clear.
7. Through researching the kinds of host plants in449samples and sooty mould in lacinsect host plants, the kinds of host plants was few influenced by the kinds of sooty mould.Initially suppose that the species of sooty mould depended on the kinds of honeydew (i.e.insect species) and environmental factors.
It is an initial study on systematics of sooty mould combined with molecular andmorphologic studies. the main characteristics of sooty mould in Yunnan were described andmore natural classification system were constructed. We provide the clear molecular evidencefor the species which were difficult to identify only relying on morphology or taxonomic status.It is the important scientific basis for identifying and system studies. The results of this study show that sooty mould are rich in diversity of Yunnan, the molecule data is not enough andmore studies were needed on systematics and molecular phylogeny.
本研究从云南27个州县上调查采集烟煤菌资源，鉴定并分离培养烟煤菌的种类；通过ITS、18S和28S序列的分析，构建系统发育树，使烟煤菌的分类系统更趋于自然的分类系统，主要结果如下：
1、采集样本449份，保存干标本96份、分离获得纯培养物62份，鉴定出真菌属于8个属，24种。
2、对烟煤菌做了系统的形态学和分子系统学研究，列出了煤炱科、刺盾炱科和Trichomeriaceae烟煤菌所在属的属征，并详细描述了所采集到的种类。联合ITS、18S和28S序列建立系统发育树，结合形态特征，分别讨论了三个科系统发育及分类地位。
3、描述了烟煤菌14个新种和4个新纪录种，其中煤炱科四个新种：Leptoxyphiumglochidion，Capnodium nonturgidum，C. lagerstroemiae，Phragmocapnias chinensis；3个新纪录种：L. kurandae，P. siamensis，P. Asiticus。刺盾炱科6个新种：Phaeosaccardinulamultiseptata，Ph. dendrocalami，Ph. yunnanensis，Ph. dalbergiae，Cyphellophora schimae，Cy. jingdongensis；1个新纪录种：Ceramothyrium thailandicum。Trichomeriaceae的4个新种：Trichomerium yunnanense，T. multisetariae，T. syzygii，T. Murrayae，通过形态学和分子系统学研究确立了这些种的分类地位以及与其相似种的区别。
4、修订了Capnodium. citri的异名：现在分类学上把C. salicinum作为C. citri的异名，以C. citri作为现在的名字（Index Fungorum2014），但是根据本研究系统发育树来看，两者的亲缘关系较远，并不能作为异名使用。
5、增加了烟煤菌的形态特征：补充了煤炱科煤炱属子囊果具刚毛；发现了胶煤炱属无性型的分生孢子器不只是基部膨大，还有中上部膨大的形态出现，海绵胶煤炱两种形态的出现是多基因和环境共同作用的结果。
6、首次发现C. coffeae和Cyphellophora的有性型：通过分子生物学上的证据，清楚了C. coffeae的有性型和无性型的形态特征，将C. coffeae的有性型和无性型很好的联系起来；描述了Cyphellophora属下3个种的有性型特征，清楚了Cy. hylomeconis的有性型和无性型之间的联系。
7、通过对449份样本寄主植物种类的研究以及紫胶虫寄主植物上烟煤菌种类的分析，发现寄主植物的种类对烟煤菌的种类影响不大，初步推测烟煤菌的种类取决于蜜露的种类（即昆虫的种类）和环境因素共同作用。
本文结合分子系统学和形态学首次对云南烟煤菌做了系统的研究，描述了云南烟煤菌特征并构建了分类系统，对一些分类地位不清楚的种类和形态上难以区别的种类，提供了清晰的分子生物学证据，为烟煤菌鉴定和系统学研究提供了的重要科学依据。研究结果表明，烟煤菌在云南多样性极其丰富，还有很多的新种和新纪录种没有被发现，烟煤菌的分子数据还很欠缺，需要更多的分子资料结合形态特征进行研究。
Sooty mould is fungus with black hypha or hyphopodia and feeding on honeydew that isexcreted from sap-sucking insects like aphids, whiteflies and mealy bugs. Sooty mould iseatable and producing the industrial chloroperoxidase, pectinases and cellulases. The secondarymetabolite of sooty mould has better anti-cancer and antibacterial components that are thefoundation for researching the pharmaceuticals and biological pesticide. Researching theresource and systematic classification is the foundation of controlling and utilizing sooty mold.Yunnan is the kingdom of the animal and plant, with rich biological diversity, but investigationof sooty mould’s system resources and identification have not been reported. The purpose ofthis study is to investigate the resources of sooty mould, to research the morphology andmolecular phylogeny of sooty mould, to provide the scientific basis and material foundation forthe controlling and utilization of sooty mould.
Samples from27states of Yunnan Province were investigated and collected, identifiedand isolated. The natural classification system of sooty mould was achieved through thephylogenetic tree based on ITS,18S and28S sequences. The main results in this study arelisted below.
1.449samples were collected,96dry samples were deposited,62sooty mould strainswere isolated,24species belonging to8genera were identified.
2. the systemic morphology and molecular phylogeny of sooty mould, and themorphological characters of all genera belonging to Capnodiaceae, Chaetothyriaceae andTrichomeriaceae were listed. All species were descripted detailedly. The phylogenetic treebased on ITS,18S and28S sequences combined with morphological characteristics werediscussed taxonomic status of three families.
3.14new species and4new records of sooty mould were collected and described in thisstudy.4new species belonging to Capnodiaceae, i.e Leptoxyphium glochidion，Capnodium nonturgidum，C. lagerstroemiae，Phragmocapnias chinensis and3new records belonging tothe same family, i.e L. kurandae，P. siamensis，P. asiticus.6new species belonging toChaetothyriaceae, i.e Phaeosaccardinula multiseptata，Ph. dendrocalami，Ph. yunnanensis，Ph. dalbergiae，Cyphellophora schimae，Cy. Jingdongensis and Ceramothyrium thailandicumwas new records belonging to the same family.4new species belonging to Trichomeriaceae, i.eTrichomerium yunnanense， T. multisetariae， T. syzygii， T. Murrayae. The systematicsplacement and differences among similar species were discussed according to the results ofmolecular data and morphological character.
4. Revised the synonym of Capnodium. citri. Now C. salicinum was as a synonym of C.ciiri on the taxonomy and C. citri was the current name. Based on the phylogenetic tree of thisstudy, the relationship of two species was more distant and C. salicinum could not be asynonym of C. citri.
5. Morphological studys shows: the ascomata of Capndium with setae, and the pycnidiumof Scorias spongiosa was not only swells on the base, but also swells on the upper part.
6. This study firstly reported the telemorph of C. coffeae and Cyphellophora. Through thephylogenetic analysis, the morphological characteristics of C. coffeae teleomorph andanamorph were linked together. Sexual characteristics of3species belonging to Cyphellophorawere descripted, the sexual and asexual characteristics of Cy. hylomeconis were clear.
7. Through researching the kinds of host plants in449samples and sooty mould in lacinsect host plants, the kinds of host plants was few influenced by the kinds of sooty mould.Initially suppose that the species of sooty mould depended on the kinds of honeydew (i.e.insect species) and environmental factors.
It is an initial study on systematics of sooty mould combined with molecular andmorphologic studies. the main characteristics of sooty mould in Yunnan were described andmore natural classification system were constructed. We provide the clear molecular evidencefor the species which were difficult to identify only relying on morphology or taxonomic status.It is the important scientific basis for identifying and system studies. The results of this study show that sooty mould are rich in diversity of Yunnan, the molecule data is not enough andmore studies were needed on systematics and molecular phylogeny.
引文
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