小盾壳科与微盾壳科真菌分类与分子系统学研究
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摘要
小盾壳科(Microthyriaceae)与微盾壳科(Micropeltidaceae)真菌是一类具有盾状囊壳和双囊壁子囊的真菌,这两科真菌国内外研究报道很少。依据目前的分类大纲小盾壳科包含了50个属,微盾壳科24个属,它们主要寄生于植物的叶片上或是腐生于枯枝落叶上。
本研究依据分类大纲,以各属的模式种标本和采集标本为研究材料,通过形态学上的显微结构特征和基于ITS、LSU、及SSU等3个基因片段的分子系统学研究,初步探讨了各属的分类地位以及系统发育关系,建立了小盾壳科的一个新属,发现了微盾壳科模式属的一个新种,从而建立了这两科真菌的更为清晰、更趋于自然的分类系统。主要研究结果和结论如下：
通过对小盾壳科的35个属的模式种以及1个新属的形态学特征和显微结构的研究,目前,我们暂时仅将7个属仍然放置于小盾壳科(Arnaudiella Petr.,Calothyriopsis Hohn., Caribaeomyces Cif.,Microthyrium Desm.,Palawania Syd.&P.Syd.,Paramicrothyrium H.X. Wu&K.D.Hyde.,Seynesiella G.Arnaud.)。我们将隶属于小盾壳科的10个属移置于星盾炱科(Asrerinaceae):Asterinella Theiss.,Asterinema Bat.&Gayao.,Asteritea Bat.&R. Garnier.,Govindua Bat.&H.Maia.,Maublancia G.Arnaud.,Polycyclinopsis Bat.A.F.Vital&I.H.Lima.,Polystomellina Bat.&A.F.Vital.,Resendea Bat.,Trichopeltosora Bat.&Cif., Xenostomella Svd.。将小盾壳科的2个属Cirsosina Bat.&J.L.Bezerra和Lembosiella Sacc.放置于Aulographaceae科。我们认为Trichothyrium Speg属,以及Actinopeltia Hohn属的模式种Actinopeltis peristomalis Hohn重新命名为Trichothyrium peristomalis(Hohn.)H.X. Wu&K.D.Hyde,comb.nov.归于Trichothyrium属,Trichothyriopsis Theiss属的模式种Trichothyriopsis densa(Racib.)Theiss.,我们使用它的同物异名Trichothyrium densum Racib.且将其并于Trichothyrium属,和Lichenopeltella Hohn属,Trichothyriomyces Bat.&H.Maia.属,Macrographa Etayo属,Pachythyrium G.Arnaud ex Spooner&P.M.Kirk属,全都放置于Trichothyriaceae科更为合适。通过,对于小盾壳科中的Asteronia(Sacc.)Henn属,Dictyoasterina Hansf.属,Tothia Bat属的研究我们认为这3个属应置于Venturiales目。通过对Muyocopron Speg属,Seynesiopeltis F. Stevens&R.W. Ryan属的研究,目前,我们暂时放置于座囊菌纲(Dothideomycetes),还不能进一步的确定其分类地位。Hidakaea I. Hino&Katum属暂放置于Hypocreales目。Helminthopeltis Sousa da Camara属放置于斑痣盘菌科(Rhytismataceae)。将Actinomyxa Syd.&P. Syd属归于肉星裂壳科(Stictidaceae)。 Stegothyrium Hohn属十分有趣,具有类似于双囊壁的子囊,但其又具有单囊壁子囊的结构；因此,我们将该属暂时放置于子囊菌门中(Ascomycota genera incertae cedis)。 Byssopeltis Bat., J.L. Bezerra&T.T. Barros属和Caudella Syd.&P. Syd属从小盾壳科中排除置于微盾壳科中。
目前,通过对微盾壳科的14个属的模式种以及该科模式属新种的形态学特征研究,暂时仍将8个属放置于微盾壳科：Chaetothyrina Theiss., Dictyopeltella Bat.&I.H. Lima., Dictyothyriella Speg., Haplopeltheca Bat., J.L. Bezerra&Cavalc., Micropeltis Mont. Stomiopeltis Theiss., StomiopeltopsisBat.&Cavalc., Stomiotheca Bat。我们将Bonaria Bat.属从微盾壳科中排除并放置于星盾炱科(Asterinaceae)。 Mitopeltis Speg属的形态特征与微盾壳科不同,被置于Microthyriales incertae cedis更为合适。Polypedia Bat.&Peres属从微盾壳科排除,放置于Parmulariaceae科。Dictyopeltis Theiss属, Hansfordiopsis Bat.属,Stigmatophragmia Tehon&G.L. Stout属依据于形态学特征,我们暂时只能将它们放置于Ascomycota genera incertae cedis这个类群,更为适合的分类地位需要进一步研究。
Scolecopeltidium F. Stevens&Manter.属隶属于小盾壳科,形态学特征与微盾壳科更为相似。通过对Scolecopeltidium F. Stevens&Manter.属的形态学特征进行了研究,发现该属建立式没有指定模式种。我们借阅到F.L. Stevens建立该属时,所包含的全部种。我们指定Scolecopeltidium mirabile F. Stevens&Manter作为该属的模式种,将Scolecopeltidium hormosporum F. Stevens&Manter仍放置于该属。Scolecopeltidium liciniae F. Stevens&Manter (寄主为Licenia sp)应该从该属排除,放置于Scolecopeltis属。建议Scolecopeltidium costi F. Stevens&Manter和Scolecopeltidium multiseptatum F. Stevens&Manter,以及Scolecopeltidium liciniae F. Stevens&Manter(寄主为Marantaceae科植物)从该属排除,放置于Micropeltis属更为合适。
分子系统学方面的研究表明：对采集标本的研究中,建立了2个新属Paramicrothyrium H.X. Wu&K.D. Hyde, gen. nov.属和Neomicrothyrium Boonmee, H.X. Wu&K.D. Hyde, gen. nov.属,并得到分子数据的支持。鉴定了Micropeltis属的1个新种Micropeltis zingiberacicola H.X. Wu&K.D. Hyde, sp. nov.。系统发育树上显示,Micropeltis zingiberacicola与Paramicrothyrium chinensis在具有盾状囊壳的真菌中,显示出更高的遗传多样性。Neomicrothyrium siamense与水生真菌中的Natupusilla属亲缘关系较近,但是没有明显相似的形态特征。Tothia fuscella (epitype)与Venturiaceae科聚为一枝,支持率为97%；同时,分子数据表明小盾壳(座)科和微盾壳(座)科真菌为非单起源类群。
The family Microthyriaceae and Micropeltidaceae sensu Lumbsch and Huhndorf2010are a poorly known but interesting family comprising50genera and24genera consisting of foliar epiphytes or saprobes on dead leaves and stems. We review two families based on examinations of generic types where possible. Members are distributed in Aulographaceae, Asterinaceae, Microthyriaceae, Micropeltidaceae and Palmulariaceae and notes are provided on each of these families. Ten genera are transferred from Microthyriaceae to Asterinaceae, and two to Aulographaceae based on the splitting or dissolving nature of the thyriothecia to release ascospores. New sequence data for a number of species and genera are provided. Microthyriaceous members growing on other fungi and lichens differ from Microthyriaceae sensu stricto and the family Trichothyriaceae is reinstated to accommodate these taxa. Other genera of Microthyriaceae belong in Rhytismataceae, Stictidaceae, Venturiales incertae cedis, Dothideomyetes genera incertae cedis, Hypocreales incertae cedis and Ascomycota genera incertae cedis.The family Microthyriaceae is reduced to seven genera characterised by superficial, flattened thyriothecia, with the cells of the upper wall radiating in parallel arrangement from the distinctcentral ostiolar opening, while the lower peridium is generally poorly developed. And the family Micropeltidaceae is reduced to eghit genera, one genus is transferred from Micropeltidaceae to Asterinaceae, one genus to Microthyriales incertae cedis, one genus to Parmulariaceae, three genera to Ascomycota genera incertae cedis. Sequence data is provided for five species with thyriothecia and Paramicrothyrium and Neomicrothyrium are described as new genera and Micropeltis zingiberacicola is introduced as a new species. Our phylogenetic analysis underscores the high genetic diversity for thyriotheciate species and there is no clear clade that can be well defined as Microthyriales. Nuclear ribosomal data support multiple polyphyletic lineages within Microthyriaceae and Micropeltidaceae. Some unexpected DNA based phylogenetic relationships such as those between Muyocopron and Saccardoella will require corroboration with more complete taxon sampling as well as additional non ribosomal markers. There are few differences between Aulographaceae, Asterinaceae and Palmulariaceae and these families may need synonymising.
本研究依据分类大纲,以各属的模式种标本和采集标本为研究材料,通过形态学上的显微结构特征和基于ITS、LSU、及SSU等3个基因片段的分子系统学研究,初步探讨了各属的分类地位以及系统发育关系,建立了小盾壳科的一个新属,发现了微盾壳科模式属的一个新种,从而建立了这两科真菌的更为清晰、更趋于自然的分类系统。主要研究结果和结论如下：
通过对小盾壳科的35个属的模式种以及1个新属的形态学特征和显微结构的研究,目前,我们暂时仅将7个属仍然放置于小盾壳科(Arnaudiella Petr.,Calothyriopsis Hohn., Caribaeomyces Cif.,Microthyrium Desm.,Palawania Syd.&P.Syd.,Paramicrothyrium H.X. Wu&K.D.Hyde.,Seynesiella G.Arnaud.)。我们将隶属于小盾壳科的10个属移置于星盾炱科(Asrerinaceae):Asterinella Theiss.,Asterinema Bat.&Gayao.,Asteritea Bat.&R. Garnier.,Govindua Bat.&H.Maia.,Maublancia G.Arnaud.,Polycyclinopsis Bat.A.F.Vital&I.H.Lima.,Polystomellina Bat.&A.F.Vital.,Resendea Bat.,Trichopeltosora Bat.&Cif., Xenostomella Svd.。将小盾壳科的2个属Cirsosina Bat.&J.L.Bezerra和Lembosiella Sacc.放置于Aulographaceae科。我们认为Trichothyrium Speg属,以及Actinopeltia Hohn属的模式种Actinopeltis peristomalis Hohn重新命名为Trichothyrium peristomalis(Hohn.)H.X. Wu&K.D.Hyde,comb.nov.归于Trichothyrium属,Trichothyriopsis Theiss属的模式种Trichothyriopsis densa(Racib.)Theiss.,我们使用它的同物异名Trichothyrium densum Racib.且将其并于Trichothyrium属,和Lichenopeltella Hohn属,Trichothyriomyces Bat.&H.Maia.属,Macrographa Etayo属,Pachythyrium G.Arnaud ex Spooner&P.M.Kirk属,全都放置于Trichothyriaceae科更为合适。通过,对于小盾壳科中的Asteronia(Sacc.)Henn属,Dictyoasterina Hansf.属,Tothia Bat属的研究我们认为这3个属应置于Venturiales目。通过对Muyocopron Speg属,Seynesiopeltis F. Stevens&R.W. Ryan属的研究,目前,我们暂时放置于座囊菌纲(Dothideomycetes),还不能进一步的确定其分类地位。Hidakaea I. Hino&Katum属暂放置于Hypocreales目。Helminthopeltis Sousa da Camara属放置于斑痣盘菌科(Rhytismataceae)。将Actinomyxa Syd.&P. Syd属归于肉星裂壳科(Stictidaceae)。 Stegothyrium Hohn属十分有趣,具有类似于双囊壁的子囊,但其又具有单囊壁子囊的结构；因此,我们将该属暂时放置于子囊菌门中(Ascomycota genera incertae cedis)。 Byssopeltis Bat., J.L. Bezerra&T.T. Barros属和Caudella Syd.&P. Syd属从小盾壳科中排除置于微盾壳科中。
目前,通过对微盾壳科的14个属的模式种以及该科模式属新种的形态学特征研究,暂时仍将8个属放置于微盾壳科：Chaetothyrina Theiss., Dictyopeltella Bat.&I.H. Lima., Dictyothyriella Speg., Haplopeltheca Bat., J.L. Bezerra&Cavalc., Micropeltis Mont. Stomiopeltis Theiss., StomiopeltopsisBat.&Cavalc., Stomiotheca Bat。我们将Bonaria Bat.属从微盾壳科中排除并放置于星盾炱科(Asterinaceae)。 Mitopeltis Speg属的形态特征与微盾壳科不同,被置于Microthyriales incertae cedis更为合适。Polypedia Bat.&Peres属从微盾壳科排除,放置于Parmulariaceae科。Dictyopeltis Theiss属, Hansfordiopsis Bat.属,Stigmatophragmia Tehon&G.L. Stout属依据于形态学特征,我们暂时只能将它们放置于Ascomycota genera incertae cedis这个类群,更为适合的分类地位需要进一步研究。
Scolecopeltidium F. Stevens&Manter.属隶属于小盾壳科,形态学特征与微盾壳科更为相似。通过对Scolecopeltidium F. Stevens&Manter.属的形态学特征进行了研究,发现该属建立式没有指定模式种。我们借阅到F.L. Stevens建立该属时,所包含的全部种。我们指定Scolecopeltidium mirabile F. Stevens&Manter作为该属的模式种,将Scolecopeltidium hormosporum F. Stevens&Manter仍放置于该属。Scolecopeltidium liciniae F. Stevens&Manter (寄主为Licenia sp)应该从该属排除,放置于Scolecopeltis属。建议Scolecopeltidium costi F. Stevens&Manter和Scolecopeltidium multiseptatum F. Stevens&Manter,以及Scolecopeltidium liciniae F. Stevens&Manter(寄主为Marantaceae科植物)从该属排除,放置于Micropeltis属更为合适。
分子系统学方面的研究表明：对采集标本的研究中,建立了2个新属Paramicrothyrium H.X. Wu&K.D. Hyde, gen. nov.属和Neomicrothyrium Boonmee, H.X. Wu&K.D. Hyde, gen. nov.属,并得到分子数据的支持。鉴定了Micropeltis属的1个新种Micropeltis zingiberacicola H.X. Wu&K.D. Hyde, sp. nov.。系统发育树上显示,Micropeltis zingiberacicola与Paramicrothyrium chinensis在具有盾状囊壳的真菌中,显示出更高的遗传多样性。Neomicrothyrium siamense与水生真菌中的Natupusilla属亲缘关系较近,但是没有明显相似的形态特征。Tothia fuscella (epitype)与Venturiaceae科聚为一枝,支持率为97%；同时,分子数据表明小盾壳(座)科和微盾壳(座)科真菌为非单起源类群。
The family Microthyriaceae and Micropeltidaceae sensu Lumbsch and Huhndorf2010are a poorly known but interesting family comprising50genera and24genera consisting of foliar epiphytes or saprobes on dead leaves and stems. We review two families based on examinations of generic types where possible. Members are distributed in Aulographaceae, Asterinaceae, Microthyriaceae, Micropeltidaceae and Palmulariaceae and notes are provided on each of these families. Ten genera are transferred from Microthyriaceae to Asterinaceae, and two to Aulographaceae based on the splitting or dissolving nature of the thyriothecia to release ascospores. New sequence data for a number of species and genera are provided. Microthyriaceous members growing on other fungi and lichens differ from Microthyriaceae sensu stricto and the family Trichothyriaceae is reinstated to accommodate these taxa. Other genera of Microthyriaceae belong in Rhytismataceae, Stictidaceae, Venturiales incertae cedis, Dothideomyetes genera incertae cedis, Hypocreales incertae cedis and Ascomycota genera incertae cedis.The family Microthyriaceae is reduced to seven genera characterised by superficial, flattened thyriothecia, with the cells of the upper wall radiating in parallel arrangement from the distinctcentral ostiolar opening, while the lower peridium is generally poorly developed. And the family Micropeltidaceae is reduced to eghit genera, one genus is transferred from Micropeltidaceae to Asterinaceae, one genus to Microthyriales incertae cedis, one genus to Parmulariaceae, three genera to Ascomycota genera incertae cedis. Sequence data is provided for five species with thyriothecia and Paramicrothyrium and Neomicrothyrium are described as new genera and Micropeltis zingiberacicola is introduced as a new species. Our phylogenetic analysis underscores the high genetic diversity for thyriotheciate species and there is no clear clade that can be well defined as Microthyriales. Nuclear ribosomal data support multiple polyphyletic lineages within Microthyriaceae and Micropeltidaceae. Some unexpected DNA based phylogenetic relationships such as those between Muyocopron and Saccardoella will require corroboration with more complete taxon sampling as well as additional non ribosomal markers. There are few differences between Aulographaceae, Asterinaceae and Palmulariaceae and these families may need synonymising.
引文
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