中国华南及云南热带、亚热带地区枯枝暗色丝孢真菌分类研究
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摘要
在自然界中,枯枝真菌种类繁多,分布广泛。该类真菌在自然生态系统中对有机物的降解和在物质循环过程中具重要的生态学作用。深入系统地开展枯枝真菌的研究具有重要的经济意义和生态学意义。我国热带、亚热带地区植被繁茂,植物种类繁多,积累了丰富的枯枝资源,蕴含着众多的真菌种类。本文主要是对华南(包括海南、广东、湖南等省)和云南省热带、亚热带地区枯枝暗色丝孢真菌进行了分类研究,鉴定并描述了大量的属、种级分类单位,证明了我国枯枝暗色丝孢真菌存在多样性，丰富了我国的物种资源,为该类真菌资源的开发利用奠定了基础。
自2009年至2012年,从海南、广东、湖南和云南等省不同地区采集到枯枝标本1000余份,经过分离鉴定,从中发现了大量新的或值得重视的分类单位,取得了许多有价值的研究成果。建立了2个新属,即类枝孢属Cladosporiopsis Sh.C. Ren&X.G. Zhang和蒜孢属Sativumoides Sh.C. Ren, Jian Ma&X.G. Zhang；对24个属的91个分类单位进行了形态描述、绘图,并与相似的分类单位进行了详细的比较,阐明了各自的分布区域和有性型。在这些分类单位中，21个分类单位被鉴定为新种,20个分类单位为中国新记录种,1个新组合种,49个中国已报道种。所有自然基质研究标本及单孢分离培养物均保存于山东农业大学植物病理学标本室(HSAUP)。
21个新种为：卵圆类枝孢Cladosporiopsis ovata、麻楝拟棒孢Corynesporopsis chukrasiae、椭孢异氯霉Dischloridium elliptia、红楣爱氏霉Ellisembia annesleae、银叶树爱氏霉E. heritierae、莲叶桐爱氏霉E. hernandiae、具喙爱氏霉E. rostrata、含笑内隔孢Endophragmiella micheliae、南岭内隔孢E. nanlingensis、木莲棒梗孢Exserticlava manglietiae、红楣异参孢Heteroconium annesleae、梭孢异参孢H. fusiformis、新木姜子异参孢H. neulitseue、椭孢层出孢Repetophragma ellipsoidea、黑柃层出孢R. euryae、海南层出孢R. hainanensis、石榴蒜孢Sativumoides punicae、具喙异棒孢Solicorynespora rostrata、卵孢绒落孢Spiropes ovata、榄仁绒落菌S. terminaliae、云南葚孢Sporidesmium yunnunense、
1个中国新记录属：拟异参孢属Xenoheteroconium Bhat, W.B. Kendr.&Nag Raj。
20个中国新记录种：油棕密格孢Acrodictys elaeidicola M.B. Ellis、类隔孢密格孢A.septosporioides Matsush.、类粉衣棒孢Corynespora calicioidea (Berk.&Broome) M.B. Ellis、巨型棒孢C. gigaspora (Berk.&Broome) M.B. Ellis、天料木棒孢C. homaliicolaDeighton&M.B. Ellis.厚皮树棒孢C. lanneicola Deighton&M.B. Ellis.灰黄拟棒孢Corynesporopsis isabelicae Hol.-Jech..柱孢拟棒孢C. cylindrica B. Sutton.普瑞斯弯孢Curvularia prasadii R.L. Mathur&B.L. Mathur.蕉斑双球霉Diplococcium stoveri (M.B. Ellis) R.C. Sinclair, Eicker&Bhat.不等隔异氯霉Dischloridium inaequiseptata (Matsush.) Hol.-Jech.、拉恩斯异氯霉D. laeense (Matsush.) B. Sutton.三隔异氯霉D. triseptalum Hol.-Jcch..树皮内隔孢Endophragmiella corticola P.M. Kirk^美丽内隔孢E. pulchra (B. Sutton&Hodges) P.M. Kirk、树脂内隔孢E. resinae P.M. Kirk.具喙内隔孢E. rostrala P.M. Kirk.水松内隔孢E. taxi (M.B. Ellis) S. Hughes.姆兰杰异棒孢Solicorynespora mulanjeensis (B. Sutton) R.F. Castaneda, M. Stadler&Guarro.阔孢异棒孢S.pscudolmediae (R.F. Castaneda) R.F. Castaneda&W.B. Kendr..尖孢葚孢Sporidesmium aculisponim M.B. Ellis.双色拟异参孢Xenoheteroconium bicolor Bhat, W.B. Kendr.&Nag Raj。
1个新组合种：马努爱氏霉Ellisembia mamiensis (Matsush.) S.C. Ren&X.G. Zhang。
基于上述分类研究，总结出了与枝暗色丝孢真菌相关的行之有效的分类方法、分类标准和分类观点：
1．大多数枯枝丝孢真菌为腐生菌，它们的分生孢子形态、大小和颜色等特征易受培养基质和培养条件的影响，往往会与自然基质上的特征不同。枯枝暗音色丝孢真菌在自然界长期的进化过程中形成了较为稳定的形态特征，因此，对该类真菌的分类研究应以自然基质上的形态特征作为主要的鉴定依据，培养性状可作为必要的补允和辅助。
2．在属级分类标准水平上，分生孢子的形成方式和分生孢子梗（产孢细胞）的延伸方式是属级划分的重要标难。对于分生孢子的形成方式和分生孢子梗的延伸方式都相同的属，区分的标准主要根据分生孢子隔膜的类型、隔膜的发育特征以及产孢细胞的形态特征，分生孢子梗和分生孢子的萌发方式也是重要的标准。有时分生孢子的着生方式(单生或链生)往往成为区分一些属的唯一标准。此外，菌落形状、颜色以及生长特性在有些属于比较稳定的特征，对一些属的区分具有参考意义。
3．在种级分类标准水平上，分生孢子的形态特征，包括分生孢子的形状、大小、隔膜数、附属丝或喙的有无，颜色以及纹饰等是重要的鉴定标准。分生孢子梗（产孢细胞）的层出特征也可作为种的分类标准。另外，子座的有无、分生孢子梗是否分枝也可作为种级分类的标准。
真菌的分类鉴定是一个十分复杂的过程,一个分类单位的鉴定要综合考虑多种分类指标,借助不同的分类方法,既要观察其整体特征,又要关注其某局部特征,既要了解与相似分类单位的共同属性,又要注重其本质的差别。
The wood-inhabiting microfungi are very abundant and widely distributed in the nature. These fungi play an important ecological role in the decomposition and recycling of materials in natural ecosystems. There is economic and ecological significance to study these fungi. Abundant decaying wood and dead branches in the tropical and subtropical regions of China support a high level of dematiaceous hyphomycetes diversity. This study mainly aims at the taxonomic research of dematiaceous hyphomycetes on decaying wood collected in the tropical and subtropical regions of southern China and Yunnan Province.
During the investigations of conidial fungi from2009to2012, we collected more than1000specimens from different ecological regions in Hainan, Guangdong, Hunan and Yunnan Provinces. We have discovered quite a few new and noteworthy taxa and achieved many aluable results based on the careful identification. Two new genera, Cladosporiopsis Sh.C Ren&X.G. Zhang and Sativumoides Sh.C. Ren, Jian Ma&X.G. Zhang are established. One newly recorded genus from China, Xenoheteroconium Bhat, W.B. Kendr.&Nag Raj is introduced. Add up to91taxa included in24genera of dematiaceous hyphomycetes arc described, illustrated, and compared with closely related taxa.
Among them,21taxa are proposed to new species, they are Cladosporiopsis ovata Sh.C Ren&X.G. Zhang, Corynesporopsis chukrasiae S.C. Ren&X.G. Zhang, Dischloridium elliptia Sh.C. Ren&X.G. Zhang, Ellisembia annesleae S.C. Ren&X.G. Zhang, E. heritierae S.C. Ren&X.G. Zhang, E. hernandiae S.C. Ren&X.G. Zhang, E. rostrata S.C. Ren&X.G. Zhang, Endophragmiella micheliae S.C. Ren&X.G. Zhang, E. nanlingensis S.C. Ren&X.G. Zhang, Exserticlava manglietiae S.C. Ren&X.G. Zhang, Heleroconium annesleae S.C. Ren&X.G. Zhang,H. fusiformis S.C. Ren&X.G. Zhang, H. neolitseae S.C. Ren&X.G. Zhang, Repelophragma ellipsoidea S.C. Ren&X.G. Zhang, R. euryae S.C. Ren&X.G. Zhang, R. hainanensis S.C. Ren&X.G. Zhang, Sativumoides punicue Sh.C. Ren, Jian Ma&X.G. Zhang. Solicorynespora rostrala S.C. Ren&X.G. Zhang, Spiropes ovata S.C. Ren&X.G. Zhang. S. terminaliae S.C. Ren&X.G. Zhang, Sporidesmium yunnanense S. C. Ren&X.G. Zhang.
20species arc newly recorded to China:Acrodiclys elaeidicola M.B. Ellis, A. septosporioides Matsush., Corynespora calicioidea (Berk.&Broome) M.B. Ellis, C. gigaspora (Berk.&Broome) M.B. Ellis, C. homaliicola Deighton&M.B. Ellis, C. lanneicola Deighten&M.B. Ellis, Corynesporopsis isabelicae Hol.-Jech., C. cylindrica B.C. Sutton, Curvularia prasadii R.E. Mathur&B.L. Mathur, Diplococcium stoveri (M.B. Ellis) R.C Sinclair, Eickcr&Bhat, Dischloridium inaequiseplata (Matsush.) Hol.-Jech., D. laeense (Matsush.) B. Sutton, D. triseptatum Hol.-Jech., Ceska Mykol, Endophragmiella corticola P.M. Kirk, E. pulchra (B. Sutton&Hodges) P.M. Kirk, E. resinae P.M. Kirk, E. rostrata P.M. Kirk, E. laxi (M.B. Ellis) S. Hughes, Solicorynespora mulanjeensis (B. Sutton) R.F. Castaneda, M. Stadler&Guarro, S. pseudolmediae (R.F. Castaneda) R.F. Castaneda&W.B. Kendr., Sporidesmium acutisporum M.B. Ellis, Xenoheteroconium bieolor Bhat, W.B. Kendr.&Nag Raj.
1taxon, Ellisembia manuensis (Matsush.) S.C. Ren&X.G. Zhang is proposed to new combination.
49taxa are described as reported species from China. All specimens are deposited in the Herbarium of the Department of Plant Pathology, Shandong Agricultural University (HSAUP).
Based on our studied works, some useful taxonomic methods, criterias and viewpoints concerning this group of fungi were concluded:
1. Most of decaying wood dematiaceous hyphomycetes are saprophytic. Their conidial morphology and size are accessible to the culture condition and culture media, and different from those surviving on the natural substrates. During the long-term evolutionary process in nature, wood-inhabiting dematiaceous hyphomycetes have formed more constant morphological characteristics. Therefore, the morphology of these fungi on the natural substrates should be primary identifying bases, while culture features can be used essential supplements and assistances.
2. At genus level of taxonomic criteria, the patterns of conidial formation and conidiophores (conidiogenous cells) proliferation are the main criteria to separate different genera. To differentiate those genera having same patterns of conidial formation and conidiophores proliferation, the type of septum, the order of septum ontogenyand and the morphological characters of conidiogenous cells should be considered. Conidiophores and patterns of conidia germination also play very important roles. Sometimes, the conidial inserted characteristics (solitary or catenate) is the only criterion to separate few genera. In addition, the characters, color and growth traits of colonies are assistant to distinguish some genera.
3. At species level of taxonomic criteria, the morphological characteristics of conidia, especially the shape, size, septum, appendage, rostrate, color and ornamentation arc main diagnostic characteristics. The features of conidiophores (conidiogenous cells) proliferation is sometimes used to differentiate species in some genera. Moreover, stromata are presence or not, branched or simple and the size of conidiophores are different among some species, which can be one of taxonomic criteria at species level.
Fungal classification and identification is a quite complex process, and all kinds of taxonomic criterias should be considered generally, and depend on various taxonomic mathods. It is necessary to observe the overall characteristics, but also concern on local features, and not only concert on their common characteristics, but also focus on their essential differences.
自2009年至2012年,从海南、广东、湖南和云南等省不同地区采集到枯枝标本1000余份,经过分离鉴定,从中发现了大量新的或值得重视的分类单位,取得了许多有价值的研究成果。建立了2个新属,即类枝孢属Cladosporiopsis Sh.C. Ren&X.G. Zhang和蒜孢属Sativumoides Sh.C. Ren, Jian Ma&X.G. Zhang；对24个属的91个分类单位进行了形态描述、绘图,并与相似的分类单位进行了详细的比较,阐明了各自的分布区域和有性型。在这些分类单位中，21个分类单位被鉴定为新种,20个分类单位为中国新记录种,1个新组合种,49个中国已报道种。所有自然基质研究标本及单孢分离培养物均保存于山东农业大学植物病理学标本室(HSAUP)。
21个新种为：卵圆类枝孢Cladosporiopsis ovata、麻楝拟棒孢Corynesporopsis chukrasiae、椭孢异氯霉Dischloridium elliptia、红楣爱氏霉Ellisembia annesleae、银叶树爱氏霉E. heritierae、莲叶桐爱氏霉E. hernandiae、具喙爱氏霉E. rostrata、含笑内隔孢Endophragmiella micheliae、南岭内隔孢E. nanlingensis、木莲棒梗孢Exserticlava manglietiae、红楣异参孢Heteroconium annesleae、梭孢异参孢H. fusiformis、新木姜子异参孢H. neulitseue、椭孢层出孢Repetophragma ellipsoidea、黑柃层出孢R. euryae、海南层出孢R. hainanensis、石榴蒜孢Sativumoides punicae、具喙异棒孢Solicorynespora rostrata、卵孢绒落孢Spiropes ovata、榄仁绒落菌S. terminaliae、云南葚孢Sporidesmium yunnunense、
1个中国新记录属：拟异参孢属Xenoheteroconium Bhat, W.B. Kendr.&Nag Raj。
20个中国新记录种：油棕密格孢Acrodictys elaeidicola M.B. Ellis、类隔孢密格孢A.septosporioides Matsush.、类粉衣棒孢Corynespora calicioidea (Berk.&Broome) M.B. Ellis、巨型棒孢C. gigaspora (Berk.&Broome) M.B. Ellis、天料木棒孢C. homaliicolaDeighton&M.B. Ellis.厚皮树棒孢C. lanneicola Deighton&M.B. Ellis.灰黄拟棒孢Corynesporopsis isabelicae Hol.-Jech..柱孢拟棒孢C. cylindrica B. Sutton.普瑞斯弯孢Curvularia prasadii R.L. Mathur&B.L. Mathur.蕉斑双球霉Diplococcium stoveri (M.B. Ellis) R.C. Sinclair, Eicker&Bhat.不等隔异氯霉Dischloridium inaequiseptata (Matsush.) Hol.-Jech.、拉恩斯异氯霉D. laeense (Matsush.) B. Sutton.三隔异氯霉D. triseptalum Hol.-Jcch..树皮内隔孢Endophragmiella corticola P.M. Kirk^美丽内隔孢E. pulchra (B. Sutton&Hodges) P.M. Kirk、树脂内隔孢E. resinae P.M. Kirk.具喙内隔孢E. rostrala P.M. Kirk.水松内隔孢E. taxi (M.B. Ellis) S. Hughes.姆兰杰异棒孢Solicorynespora mulanjeensis (B. Sutton) R.F. Castaneda, M. Stadler&Guarro.阔孢异棒孢S.pscudolmediae (R.F. Castaneda) R.F. Castaneda&W.B. Kendr..尖孢葚孢Sporidesmium aculisponim M.B. Ellis.双色拟异参孢Xenoheteroconium bicolor Bhat, W.B. Kendr.&Nag Raj。
1个新组合种：马努爱氏霉Ellisembia mamiensis (Matsush.) S.C. Ren&X.G. Zhang。
基于上述分类研究，总结出了与枝暗色丝孢真菌相关的行之有效的分类方法、分类标准和分类观点：
1．大多数枯枝丝孢真菌为腐生菌，它们的分生孢子形态、大小和颜色等特征易受培养基质和培养条件的影响，往往会与自然基质上的特征不同。枯枝暗音色丝孢真菌在自然界长期的进化过程中形成了较为稳定的形态特征，因此，对该类真菌的分类研究应以自然基质上的形态特征作为主要的鉴定依据，培养性状可作为必要的补允和辅助。
2．在属级分类标准水平上，分生孢子的形成方式和分生孢子梗（产孢细胞）的延伸方式是属级划分的重要标难。对于分生孢子的形成方式和分生孢子梗的延伸方式都相同的属，区分的标准主要根据分生孢子隔膜的类型、隔膜的发育特征以及产孢细胞的形态特征，分生孢子梗和分生孢子的萌发方式也是重要的标准。有时分生孢子的着生方式(单生或链生)往往成为区分一些属的唯一标准。此外，菌落形状、颜色以及生长特性在有些属于比较稳定的特征，对一些属的区分具有参考意义。
3．在种级分类标准水平上，分生孢子的形态特征，包括分生孢子的形状、大小、隔膜数、附属丝或喙的有无，颜色以及纹饰等是重要的鉴定标准。分生孢子梗（产孢细胞）的层出特征也可作为种的分类标准。另外，子座的有无、分生孢子梗是否分枝也可作为种级分类的标准。
真菌的分类鉴定是一个十分复杂的过程,一个分类单位的鉴定要综合考虑多种分类指标,借助不同的分类方法,既要观察其整体特征,又要关注其某局部特征,既要了解与相似分类单位的共同属性,又要注重其本质的差别。
The wood-inhabiting microfungi are very abundant and widely distributed in the nature. These fungi play an important ecological role in the decomposition and recycling of materials in natural ecosystems. There is economic and ecological significance to study these fungi. Abundant decaying wood and dead branches in the tropical and subtropical regions of China support a high level of dematiaceous hyphomycetes diversity. This study mainly aims at the taxonomic research of dematiaceous hyphomycetes on decaying wood collected in the tropical and subtropical regions of southern China and Yunnan Province.
During the investigations of conidial fungi from2009to2012, we collected more than1000specimens from different ecological regions in Hainan, Guangdong, Hunan and Yunnan Provinces. We have discovered quite a few new and noteworthy taxa and achieved many aluable results based on the careful identification. Two new genera, Cladosporiopsis Sh.C Ren&X.G. Zhang and Sativumoides Sh.C. Ren, Jian Ma&X.G. Zhang are established. One newly recorded genus from China, Xenoheteroconium Bhat, W.B. Kendr.&Nag Raj is introduced. Add up to91taxa included in24genera of dematiaceous hyphomycetes arc described, illustrated, and compared with closely related taxa.
Among them,21taxa are proposed to new species, they are Cladosporiopsis ovata Sh.C Ren&X.G. Zhang, Corynesporopsis chukrasiae S.C. Ren&X.G. Zhang, Dischloridium elliptia Sh.C. Ren&X.G. Zhang, Ellisembia annesleae S.C. Ren&X.G. Zhang, E. heritierae S.C. Ren&X.G. Zhang, E. hernandiae S.C. Ren&X.G. Zhang, E. rostrata S.C. Ren&X.G. Zhang, Endophragmiella micheliae S.C. Ren&X.G. Zhang, E. nanlingensis S.C. Ren&X.G. Zhang, Exserticlava manglietiae S.C. Ren&X.G. Zhang, Heleroconium annesleae S.C. Ren&X.G. Zhang,H. fusiformis S.C. Ren&X.G. Zhang, H. neolitseae S.C. Ren&X.G. Zhang, Repelophragma ellipsoidea S.C. Ren&X.G. Zhang, R. euryae S.C. Ren&X.G. Zhang, R. hainanensis S.C. Ren&X.G. Zhang, Sativumoides punicue Sh.C. Ren, Jian Ma&X.G. Zhang. Solicorynespora rostrala S.C. Ren&X.G. Zhang, Spiropes ovata S.C. Ren&X.G. Zhang. S. terminaliae S.C. Ren&X.G. Zhang, Sporidesmium yunnanense S. C. Ren&X.G. Zhang.
20species arc newly recorded to China:Acrodiclys elaeidicola M.B. Ellis, A. septosporioides Matsush., Corynespora calicioidea (Berk.&Broome) M.B. Ellis, C. gigaspora (Berk.&Broome) M.B. Ellis, C. homaliicola Deighton&M.B. Ellis, C. lanneicola Deighten&M.B. Ellis, Corynesporopsis isabelicae Hol.-Jech., C. cylindrica B.C. Sutton, Curvularia prasadii R.E. Mathur&B.L. Mathur, Diplococcium stoveri (M.B. Ellis) R.C Sinclair, Eickcr&Bhat, Dischloridium inaequiseplata (Matsush.) Hol.-Jech., D. laeense (Matsush.) B. Sutton, D. triseptatum Hol.-Jech., Ceska Mykol, Endophragmiella corticola P.M. Kirk, E. pulchra (B. Sutton&Hodges) P.M. Kirk, E. resinae P.M. Kirk, E. rostrata P.M. Kirk, E. laxi (M.B. Ellis) S. Hughes, Solicorynespora mulanjeensis (B. Sutton) R.F. Castaneda, M. Stadler&Guarro, S. pseudolmediae (R.F. Castaneda) R.F. Castaneda&W.B. Kendr., Sporidesmium acutisporum M.B. Ellis, Xenoheteroconium bieolor Bhat, W.B. Kendr.&Nag Raj.
1taxon, Ellisembia manuensis (Matsush.) S.C. Ren&X.G. Zhang is proposed to new combination.
49taxa are described as reported species from China. All specimens are deposited in the Herbarium of the Department of Plant Pathology, Shandong Agricultural University (HSAUP).
Based on our studied works, some useful taxonomic methods, criterias and viewpoints concerning this group of fungi were concluded:
1. Most of decaying wood dematiaceous hyphomycetes are saprophytic. Their conidial morphology and size are accessible to the culture condition and culture media, and different from those surviving on the natural substrates. During the long-term evolutionary process in nature, wood-inhabiting dematiaceous hyphomycetes have formed more constant morphological characteristics. Therefore, the morphology of these fungi on the natural substrates should be primary identifying bases, while culture features can be used essential supplements and assistances.
2. At genus level of taxonomic criteria, the patterns of conidial formation and conidiophores (conidiogenous cells) proliferation are the main criteria to separate different genera. To differentiate those genera having same patterns of conidial formation and conidiophores proliferation, the type of septum, the order of septum ontogenyand and the morphological characters of conidiogenous cells should be considered. Conidiophores and patterns of conidia germination also play very important roles. Sometimes, the conidial inserted characteristics (solitary or catenate) is the only criterion to separate few genera. In addition, the characters, color and growth traits of colonies are assistant to distinguish some genera.
3. At species level of taxonomic criteria, the morphological characteristics of conidia, especially the shape, size, septum, appendage, rostrate, color and ornamentation arc main diagnostic characteristics. The features of conidiophores (conidiogenous cells) proliferation is sometimes used to differentiate species in some genera. Moreover, stromata are presence or not, branched or simple and the size of conidiophores are different among some species, which can be one of taxonomic criteria at species level.
Fungal classification and identification is a quite complex process, and all kinds of taxonomic criterias should be considered generally, and depend on various taxonomic mathods. It is necessary to observe the overall characteristics, but also concern on local features, and not only concert on their common characteristics, but also focus on their essential differences.
引文
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