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中国南方七省凋落枯枝暗色丝孢真菌分类研究
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摘要
我国陆地面积广阔,生态环境和植被结构类型复杂多样,积累了丰富的适于真菌生存繁衍的凋落枯枝资源,蕴涵着丰富多样的暗色丝孢真菌类群,但目前对该类基质上暗色丝孢真菌资源的挖掘十分贫乏,更缺乏对此类真菌属、种多样性的系统调查和分类研究。通过广泛征集我国热带及亚热带地区(主要是:海南、云南、广东、福建、广西、湖南及四川)不同生境及植被类型区域凋落枯枝,立足经典分类,借助显微描绘和照相技术,利用种菌形态学分类性状,系统开展了43个暗色丝孢真菌属的分类研究,修订完善了类棒孢属(Corynesporella)的属级特征,澄清了双曲孢属(Nakataea)、异参孢属(Heteroconium)、异棒孢属(Solicorynespora)及棕榈枝孢属(Spadicoides)种级分类单位的混乱,完善了具节束梗孢属(Gangliostilbe)等多个属相关物种的分类检索表,正确描述鉴定新种30个,中国新记录属2个,中国新记录种24个,中国已报道种27个,丰富了该类群真菌分类研究的内容,为深入开展其它生态地理区域和生存基质暗色丝孢真菌属、种资源多样性研究提供经验积累。
     合格描述鉴定的新种、中国新记录属(种)及已报道种分别是:
     新种30个:福建棒孢Corynespora fujianensis L.G. Ma&X.G. Zhang,福建阔领瓶梗孢Craspedodidymum fujianense L.G. Ma&X.G. Zhang,倒棍棒类棒孢Corynesporellaobclavata L.G. Ma&X.G. Zhang,三隔类棒孢Corynesporella trisepata L.G. Ma&X.G.Zhang,蒲葵双串孢Diplococcium livistonae L.G. Ma&X.G. Zhang,黄皮内隔孢Endophragmiella clausenae L.G. Ma&X.G. Zhang,五列木内隔孢Endophragmiellapentaphylacis L.G. Ma&X.G. Zhang,云南具节束梗孢Gangliostilbe yunnanensis L.G.Ma&X.G. Zhang,广东基膨孢Guedea guangdongense L.G. Ma&X.G. Zhang,榕生异参孢Heteroconium fici L.G. Ma&X.G. Zhang,观光木异参孢Heteroconiumtsoongiodendronis L.G. Ma&X.G. Zhang,云南异参孢Heteroconium yunnanensis L.G.Ma&X.G. Zhang,倒棍棒双曲孢Nakataea obclavata L.G. Ma&X.G. Zhang,广东双曲孢Nakataea guangdongense L.G. Ma&X.G. Zhang,四川双曲孢Nakataea sichuanenseL.G. Ma&X.G. Zhang,云南假绒落菌Pseudospiropes yunnanense L.G. Ma&X.G.Zhang,椭圆侧多隔孢Pleurophragmium ellipsoidea L.G. Ma&X.G. Zhang,云南侧多隔孢Pleurophragmium yunnanensis L.G. Ma&X.G. Zhang,棍形侧多隔孢Pleurophragmium clavata L.G. Ma&X.G. Zhang,云南暗色串孢Phaeomoniliayunnanense L.G. Ma&X.G. Zhang,四川拟侧耳霉Pleurotheciopsis sichuanense L.G. Ma&X.G. Zhang,广东层出孢Repetophragma guangdongense L.G. Ma&X.G. Zhang,云南类葚孢Sporidesmiella yunnanensis L.G. Ma&X.G. Zhang,四川类葚孢Sporidesmiellasichuanensis L.G. Ma&X.G. Zhang,云南棕榈枝孢Spadicoides yunnanensis L.G. Ma&X.G. Zhang,山茶棕榈枝孢Spadicoides camelliae L.G. Ma&X.G. Zhang,杆形棕榈枝孢Spadicoides bacilliformis L.G. Ma&X.G. Zhang,云南葚孢Sporidesmium yunnanensisL.G. Ma&X.G. Zhang,粗叶木异棒孢Solicorynespora lasianthi L.G. Ma&X.G. Zhang,双隔异棒孢Solicorynespora biseptata L.G. Ma&X.G. Zhang。
     中国新记录属2个:莫氏霉属Moorella P.Rag. Rao&D. Rao、布氏霉属Stephembruneria R.F. Casta eda。
     中国新记录种24个:球形瓶梗球孢Bahusutrabeeja globosa Bhat&W.B. Kendr.,不列颠毛内隔孢Chaetendophragmia britannica P.M. Kirk,层出阔领瓶梗孢Craspedodidymum proliferans V. Rao&de Hoog,新西兰小角孢Ceratosporella novae-zelandiae S. Hughes,圆柱形拟棒孢Corynesporopsis cylindrica B. Sutton,立陶宛暗双孢Cordana lithuanica Markovsk.,瓜达卡纳尔瓶梗附孢Cryptophiale guadalcanalensisMatsush.,西表瓶梗附孢Cryptophiale iriomoteanum Matsush.,宇田川瓶梗附孢Cryptophiale udagawae Piroz.&Ichinoe,非洲异氯霉Dischloridium ychaffrei (Bhat&B.Sutton) Hol.-Jech.,普尔尼双串孢Diplococcium pulneyense Subram.&Sekar,单隔棒梗孢Exserticlava uniseptata Bhat&B. Sutton,树皮生内隔孢Endophragmiella corticolaP.M. Kirk,非洲基膨孢Guedea sacra Rambelli&Bartoli,美丽异参孢Heteroconiumdecorosum R.F. Casta eda,Saikawa&Guarro,大洋洲异参孢Heteroconium ponapenseMatsush.,异生柱形孢Kylindria excentrica Bhat&B. Sutton,肥大柱形孢Kylindriaobesispora R.F. Casta eda,杯形黑头孢Melanocephala cupulifera S. Hughes,单生莫氏霉Moorella monocephala Matsush.,八丈岛假绒落菌Pseudospiropes hachijoensisMatsush.,假隔类葚孢Sporidesmiella pseudoseptata (M.B. Ellis) Subram.,澳大利亚棕榈枝孢Spadicoides australiensis Whitton, K.D. Hyde&McKenzie,雅致布氏霉Stephembruneria elegans R.F. Casta eda。
     中国已报道种27个:休斯蛛丝孢Arachnophora hughesii R.F. Casta eda&Guarro,单隔冠孢Coronospora uniseptata Matsushima,具柄小角孢Ceratosporella stipitata(Goidanich) S. Hughes,栎生拟棒孢Corynesporopsis quercicola (Borowska) P.M. Kirk,伊莎贝拉拟棒孢Corynesporopsis isabelicae Hol.-Jech.,黑色树状霉Dendryphiopsis atra(Corda) S. Hughes,印度指状孢Digitoramispora tambdisurlensis Pratibha,Raghuk.&Bhat,美丽内隔孢Endophragmiella pulchra (B. Sutton&Hodges) P.M. Kirk,新西兰基膨孢Guedea novae-zelandiae Hugues,卵形基膨孢Guedea ovata Morgan-Jones,R.C.Sinclair&Eicker,三隔棒梗孢Exserticlava triseptata (Matsush.) S. Hughes,土库曼异型格孢Hermatomyces tucumanensis Speg.,粗糙轮缘孢Ityorhoptrum verruculosum (M.B.Ellis) P.M. Kirk,印度束葚孢Morrisiella indica Saikia&A.K. Sarbhoy,山腊梅拟折孢Minimelanolocus chimonanthi Y.D. Zhang&X.G. Zhang,腐木拟虫囊孢Listeromycesinsignis Penz.&Sacc.,橄榄色拟折孢Minimelanolocus olivaceus R.F.Casta eda&Guarro,表生拟树状霉Paradendryphiopsis cambrensis M.B. Ellis,哥斯达黎加假绒落菌Pseudospiropes costaricensis (E.F. Morris) de Hoog&Arx,黑束柄霉Podosporiumnilgirense (Subram.) M.B. Ellis,棍棒类葚孢Sporidesmiella claviformis P.M. Kirk,竹生棕榈枝孢Spadicoides bambusicola D.Q. Zhou,Goh&K.D. Hyde,异隔棕榈枝孢Spadicoides versiseptatis M.K.M. Wong,Goh&K.D. Hyde,小孢葚孢Sporidesmiumparvum (S. Hughes) M.B. Ellis,尖孢葚孢Sporidesmium acutisporum M.B. Ellis,丹尼斯拟葚孢Sporidesmiopsis dennisii (J.L. Crane&Dumont) Bhat,W.B. Kendr.&Nag Raj,乌干达玉米孢Tretospeira ugandensis (Hansf) Pirozynski。
     上述研究标本均保存于山东农业大学植物病理学标本室(HSAUP)与中国科学院微生物研究所菌物标本馆(HMAS)。
     无性态真菌作为一独立类群,其分类研究主要是依据无性阶段的形态特征,分类界定标准因真菌学家所持观点不同而存在差异。在多年分类研究基础上,总结前人分类观点,系统地概括了凋落枯枝暗色丝孢真菌属、种级的划分标准:
     ①以腐生为主的凋落枯枝暗色丝孢真菌,应以自然基质上的形态特征为主要鉴别依据;对于单孢分离可培养的,其培养条件下的特征和产孢表型可作必要补充;
     ②以载孢体的类型,产孢细胞的延伸方式及孢子形成方式为属级分类的主要依据;产孢位点特征、分生孢子脱落方式(裂解式/破生式)及其突出形态特征如真/假隔膜,单生/链生等有稳定差异的特征,有时也作为属级鉴别标准;产孢梗的分枝类型、产孢细胞形状、刚毛及子座有无等特征有时对少数属的划分具一定参考价值;
     ③分生孢子的形状、大小、隔膜、色泽、孢壁纹饰及附属部件特征等是分种的主要依据。共同的、突出的和稳定的分生孢子形态特征对属的划分也有意义。分生孢子隔膜的有无,形成的顺序和位置,在不同种间,有时甚至在不同属间有稳定的差异,也可作为分种(属)的标准之一;
     ④在种级分类标准中,分生孢子的形状、大小、隔膜视为主要标准,而分生孢子附属物及色素沉积等则通常视为次要标准。
China has a large territory, complicated ecological environment and vegetation structure.There is a deep woody debris layer to be a rich habitat for fungi, containing a wide range ofdematiaceous hyphomycetes. However, the hyphomycetous fungi are still mostly unexploredin China, and the systematic investigation and classification at genera and species level arenecessary. By means of collecting dead branches from the forests of different habitats andvegetation in tropical and subtropical regions (mainly including Hainan, Yunnan, Guangdong,Guangxi, Fujian and Sichuan provinces), based on the traditional taxonomy, with the aid ofmicroscopy and photography, using the characteristics of morphology, we carried out thetaxonomic studies on43genera of anamorphic hyphomycete fungi. Meanwhile, wereviewedthe generic concept of Corynesporella Munjal&H.S. Gill, added new species and madeclarifications in the genera including Nakataea, Heteroconium, Solicorynespora, Spadicoides,improved the classification keys of related species of Gangliostilbe and some other genera,correctly described and identified30new species,2new Chinese record genera,24newChinese records,27Chinese known species, and enriched the numbers of dead wooddematiaceous hyphomycetes in China. These results could provide experience accumulationfor taxonomic studies of other dematiaceous hyphomycetes, and have significance for theexploration of their diversity on fungal resources.
     The described new species, new Chinese records and Chinese known species are asfollows:
     30new species are: Corynespora fujianensis L.G. Ma&X.G. Zhang, Craspedodidymumfujianense L.G. Ma&X.G. Zhang, Corynesporella obclavata L.G. Ma&X.G. Zhang,Corynesporella trisepata L.G. Ma&X.G. Zhang, Diplococcium livistonae L.G. Ma&X.G.Zhang, Endophragmiella clausenae L.G. Ma&X.G. Zhang, Endophragmiella pentaphylacisL.G. Ma&X.G. Zhang, Gangliostilbe yunnanensis L.G. Ma&X.G. Zhang, Guedeaguangdongense L.G. Ma&X.G. Zhang, Heteroconium fici L.G. Ma&X.G. Zhang,Heteroconium tsoongiodendronis L.G. Ma&X.G. Zhang, Heteroconium yunnanensis L.G.Ma&X.G. Zhang, Nakataea obclavata L.G. Ma&X.G. Zhang, Nakataea guangdongenseL.G. Ma&X.G. Zhang, Nakataea sichuanense L.G. Ma&X.G. Zhang, Pseudospiropesyunnanense L.G. Ma&X.G. Zhang, Pleurophragmium ellipsoidea L.G. Ma&X.G. Zhang,Pleurophragmium yunnanensis L.G. Ma&X.G. Zhang, Pleurophragmium clavata L.G. Ma&X.G. Zhang, Phaeomonilia yunnanense L.G. Ma&X.G. Zhang, Pleurotheciopsissichuanense L.G. Ma&X.G. Zhang, Repetophragma guangdongense L.G. Ma&X.G. Zhang, Sporidesmiella yunnanensis L.G. Ma&X.G. Zhang, Sporidesmiella sichuanensis L.G. Ma&X.G. Zhang, Spadicoides yunnanensis L.G. Ma&X.G. Zhang, Spadicoides camelliae L.G.Ma&X.G. Zhang, Spadicoides bacilliformis L.G. Ma&X.G. Zhang, Sporidesmiumyunnanensis L.G. Ma&X.G. Zhang, Solicorynespora lasianthi L.G. Ma&X.G. Zhang,Solicorynespora biseptata L.G. Ma&X.G. Zhang.
     2new Chinese record genera are: Moorella P.Rag. Rao&D. Rao, Stephembruneria R.F.Casta eda.
     24new Chinese record species are: Bahusutrabeeja globosa Bhat&W.B. Kendr.,Chaetendophragmia britannica P.M. Kirk, Craspedodidymum proliferans V. Rao&de Hoog,Ceratosporella novae-zelandiae S. Hughes, Corynesporopsis cylindrica B. Sutton, Cordanalithuanica Markovsk., Cryptophiale guadalcanalensis Matsush., Cryptophiale iriomoteanumMatsush., Cryptophiale udagawae Piroz.&Ichinoe, Dischloridium ychaffrei (Bhat&B.Sutton) Hol.-Jech., Diplococcium pulneyense Subram.&Sekar, Exserticlava uniseptata Bhat&B. Sutton, Endophragmiella corticola P.M. Kirk, Guedea sacra Rambelli&Bartoli,Heteroconium decorosum R.F. Casta eda,Saikawa&Guarro, Heteroconium ponapenseMatsush., Kylindria excentrica Bhat&B. Sutton, Kylindria obesispora R.F. Casta eda,Melanocephala cupulifera S. Hughes, Moorella monocephala Matsush., Pseudospiropeshachijoensis Matsush., Sporidesmiella pseudoseptata (M.B. Ellis) Subram., Spadicoidesaustraliensis Whitton,K.D. Hyde&McKenzie, Stephembruneria elegans R.F. Casta eda.
     27Chinese known species are: Arachnophora hughesii R.F. Casta eda&Guarro,Coronospora uniseptata Matsushima, Ceratosporella stipitata (Goidanich) S. Hughes,Corynesporopsis quercicola (Borowska) P.M. Kirk, Corynesporopsis isabelicae Hol.-Jech.,Dendryphiopsis atra (Corda) S. Hughes, Digitoramispora tambdisurlensis Pratibha,Raghuk.&Bhat, Endophragmiella pulchra (B. Sutton&Hodges) P.M. Kirk, Guedea novae-zelandiaeHugues, Guedea ovata Morgan-Jones,R.C. Sinclair&Eicker, Exserticlava triseptata(Matsush.) S. Hughes, Hermatomyces tucumanensis Speg., Ityorhoptrum verruculosum (M.B.Ellis) P.M. Kirk, Morrisiella indica Saikia&A.K. Sarbhoy, Minimelanolocus chimonanthiY.D. Zhang&X.G. Zhang, Listeromyces insignis Penz.&Sacc., Minimelanolocus olivaceusR.F.Casta eda&Guarro, Paradendryphiopsis cambrensis M.B. Ellis, Pseudospiropescostaricensis (E.F. Morris) de Hoog&Arx, Podosporium nilgirense (Subram.) M.B. Ellis,Sporidesmiella claviformis P.M. Kirk, Spadicoides bambusicola D.Q. Zhou,Goh&K.D.Hyde, Spadicoides versiseptatis M.K.M. Wong,Goh&K.D. Hyde, Sporidesmium parvum (S.Hughes) M.B. Ellis, Sporidesmium acutisporum M.B. Ellis, Sporidesmiopsis dennisii (J.L.Crane&Dumont) Bhat,W.B. Kendr.&Nag Raj, Tretospeira ugandensis (Hansf) Pirozynski
     All specimens are deposited in Herbarium of Shandong Agricultural University, Plant Pathology (HSAUP), and Mycological Herbarium, Institute of Microbiology, Chinese Academy of Sciences (HMAS).
     Anamorphic fungi is an independent taxonomic group, its taxonomic studies are mainly based on morphological characteristics of asexual stage. However, the criteria used for genera or species delimitation has a difference due to mycologist keeping different taxonomical views. Based on the continuing studies of dematiaceous hyphomycetes occurring on dead branches, and carefully reading documents, we summarize and point out the criteria of taxonomic studies of anamorphic hyphomycetes.
     ①The taxonomic classification of dematiaceous hyphomycetes on dead branches or rotten wood should be based on the morphological characteristics on natural substrates. Culture features can be used as a necessary complement for identification.
     ②The conidiomata type, the extension of conidiogenous cells/conidiophores, and the conidiogenesis pattern are key to distinguish hyphomycetes at genus level; the characteristics of conidiogenous loci, the conidial secession (schizolytic/rhexolytic) and features such as euseptation/distoseptation, solitary/catenate conidia are occasionally used as the criteria for identification at generic level; the branching pattern of conidiophores, conidiogenous cells shape, presence or absence of sterile setae and stroma, are also useful to distinguish a few genera.
     ③Conidial morphology, including shape, size, pigmentation, septation, ornamentation and appendages are the main criteria in distinguishing species. The common, outstanding and stable conidial morphological characteristics are also helpful with genera level. The presence or absence, the forming sequence, and position of septation are also considered as one of criteria in distinguishing genera or species for different genera or species that have stable differences.
     ④For the criteria of distinguishing species, conidial shape, conidial size and septation are regarded as primary criteria, and conidial appendage, pigmentation and ornamentation are usually the secondary criteria.
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