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作者简介:

蒋树浩(2000—),硕士研究生,研究方向为地衣生物学,(E-mail)jsh20000522@163.com。

通讯作者:

贾泽峰,博士,教授,研究方向为地衣生物学,(E-mail)zfjia2008@163.com。

中图分类号:Q949.34

文献标识码:A

文章编号:1000-3142(2024)04-0621-08

DOI:10.11931/guihaia.gxzw202311012

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参考文献
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NIAZI AR, AFSHAN NS, NASEER A, et al. , 2023. A new species and a new record of the genus Phaeophyscia Moberg (Lecanorales, Physciaceae) from Pakistan supported by phenotypic and molecular phylogenetic analyses [J]. Cryptog Mycol, 44(4): 51-59.
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目录contents

    摘要

    该文基于形态学、解剖学、化学及分子系统学的方法,对采自中国泰山的黑蜈蚣叶属地衣进行了分类学研究,发现了1新种,即泰山黑蜈蚣叶(Phaeophyscia taishanensis)。该新种的主要特征为地衣体上表面末端具稀疏的皮层毛;髓层白色;下表面黑色,裂片末端处呈灰白色或浅褐色;盘托上部偶有白色或浅色的皮层毛,常常稀疏可数;子囊孢子褐色,厚壁,Physcia型,孢子大小为(18.0~20.5) μm × (9.0~10.0) μm。该文还基于表型特征讨论了新种与相似种的异同,并基于分子数据以ITS序列构建最大似然系统发育树且进行了序列分析,同时提供了详细的形态学描述及特征图片。该新种的发现为蜈蚣衣科地衣生物多样性研究积累了基础资料。

    Abstract

    Based on morphological, anatomical, chemical and molecular systematic methods, a taxonomic study was carried out on the lichen genus Phaeophyscia collected from Mount Tai. One species, P. taishanensis is reported as new to science. It is characterized by the following characters: white or hyaline cortical hairs sparsely on the upper surface of the marginal peripheral zones of the lobes; white medulla; black lower surface with sometimes white or pale brown ends; sparse cortical hairs occasionally on upper portion of thalline margin; and ascospores brown, Physcia-type, sized (18.0-20.5) μm × (9.0-10.0) μm. Based on phenotypical characteristics, the similarities and differences between the new species and similar species were discussed. the maximum likelihood phylogenetic tree was constructed with ITS sequence, and the sequence analysis was carried out. A detailed morphological description and pictures of the characteristics of this new species are provided. The discovery of this new species has accumulated basic data for the study of Physciaceae biodiversity.

  • 黑蜈蚣叶属(Phaeophyscia Moberg)隶属于子囊菌门(Ascomycota)茶渍纲(Lecanoromycetes),粉衣目(Caliciales)蜈蚣衣科(Physciaceae),全世界有66种,中国已报道21种(魏江春,2020; 陈健斌和胡光荣,2022; Wijayawardene et al.,2022; 陈健斌,2023)。

  • 黑蜈蚣叶属(Phaeophyscia)是Moberg于1977年以Phaeophyscia orbicularis(Neck.)Moberg为模式建立的,除强调皮层结构、地衣化学之外,他还强调分生孢子在地衣分属中的重要性。于是,从蜈蚣衣属Physcia中分出该属,包括原蜈蚣衣属中那些皮层K-,不含atranorin,下皮层为假薄壁组织,个别为假厚壁长细胞组织,分生孢子为近椭圆形且小于4 μm长的类群(Moberg,1977)。后来,Esslinger承认且沿用了Moberg的黑蜈蚣叶属Phaeophyscia分属观点,在此基础上将蜈蚣衣属中地衣体上皮层K-,不含atranorin,下表面为淡白色,下皮层为假厚壁长细胞组织的几个种组建了新属Phyciella Essl.,包括Physciella denigrataP. melanchraP. nepalensis(Esslinger,1978)。但是,其观点并没有被地衣学家们广泛接受。

  • 黑蜈蚣叶属地衣的主要特征为叶状地衣体,上皮层K-,不含atranorin,髓层白色或橘红色,下皮层为假薄壁组织,个别为假厚壁长细胞组织,茶渍型子囊盘,Physcia型或Pachysporaria型子囊孢子,分生孢子近椭圆形且小于4 μm长。该属地衣的生长基物多样,包括树皮、枯木、苔藓、岩石、土壤(Moberg,1977; Aptroot &Sipman,1991; Liu &Hur,2019)。该属地衣的整体分布格局具有明显的北温带性质,分布或主要分布于北温带或北方植被带,东亚和北美洲是该属地衣两个重要的分布区域(陈健斌,2023)。

  • 本研究对保存于聊城大学生命科学学院地衣标本馆(LCUF)的地衣标本进行整理和鉴定时,发现了黑蜈蚣叶属地衣1新种,现予以报道。

  • 1 材料与方法

  • 1.1 研究标本

  • 采自山东泰山的地衣标本,现保存于聊城大学生命科学学院地衣标本馆(LCUF)。

  • 1.2 形态特征观察

  • 在OLYMPUS SZX16体式显微镜下观察地衣体的外表特征,并拍照。徒手纵切子囊盘,选其薄片置于配备有OLYMPUS DP74数字相机的OLYMPUS BX53光学显微镜下进行观察解剖特征,并拍照。

  • 1.3 化学物质检测

  • 使用试剂K(10%氢氧化钾水溶液)、C(次氯酸钠饱和水溶液)和P(对苯二胺在95%乙醇中的饱和溶液)滴加在地衣体皮层或髓层上进行颜色反应(CT),并综合薄层层析法(TLC)判断地衣所含次级代谢产物,所用溶剂为C系统(甲苯∶乙酸=20∶3,比值为体积比)(Culberson &Kristinsson,1970; Culberson,1972; Orange et al.,2001)。使用浓度为0.3%~1.0%的Lugol’s溶液对子囊盘纵切薄片进行染色,观察子囊孢子淀粉样蛋白反应。

  • 1.4 基因组提取、PCR扩增及测序

  • 从标本中选取适量地衣体,使用Sigma-Aldrich REDExtract-N-AmpTM试剂盒,参照产品说明书进行地衣型真菌总基因组的提取。使用引物ITS1F和ITS4(Gardes &Bruns,1993; White et al.,1990)扩增内源转录间隔区(ITS)片段,采用Zhao和Jia(2022)的PCR反应体系和反应条件进行扩增,使用1%琼脂糖凝胶电泳检测PCR扩增产物,送由铂尚生物技术有限公司(济南)进行测序。

  • 1.5 序列比对和系统发育分析

  • 测序返回序列使用软件Geneious v.9.0.2(Biomatters Ltd.,Auckland,NZ)进行拼接,并将序列上传至GenBank(OR769731)。自测序列与从GenBank中下载的37条序列,如表1所示,使用MAFFT v.7(Katoh et al.,2009)进行多重比对(使用“L-ING-i”运算法则,其他参数为默认值)。使用Gblocks V0.19b(Castresana,2000; Talavera &Castresana,2007)对矩阵中首尾及中间的模糊序列进行消除。使用PhyloSuite v.1.2.2(Zhang et al.,2020)进行系统发育分析,利用ModelFinder(Kalyaanamoorthy et al.,2017)的AIC准则分别为最大似然法(ML)分析和贝叶斯法(BI)分析选择最优模型,ML分析的最适模型为TIM2e+R2,BI分析的最适模型为SYM+I+G。在TIM2e+R2模型下,使用IQ-TREE(Nguyen et al.,2015)进行5 000次ultrafast(Minh et al.,2013),以及Shimodaira-Hasegawa-like检验(Guindon et al.,2010)构建ML树; 在SYM+I+G模型下,使用MrBayes V.3.2.6(Ronquist et al.,2012)运行两次并行的马尔科夫链(MCMC),每条链200万代,舍弃最初25%的样本数构建BI树。使用Figtree v.1.4.4进行系统发育树的查看和调整。

  • 表1 系统发育所用的物种及序列信息

  • Table1 Species and sequence information used for phylogenetic analysis

  • 注:本研究自测序列加粗表示。

  • Note: The new sequences generated in this study are indicated in boldface.

  • 2 结果与分析

  • 2.1 系统发育分析

  • 本研究系统发育分析所用序列来源于自测序列BLAST结果以及相关文献(Fayyaz et al.,2022; Niazi et al.,2023),以Physciella melanchra外类群,共包括ITS序列38条(表1)。基于ITS序列进行系统发育分析,结果显示ML树和BI树拓扑结构一致,文中仅展示ML系统发育树,ML系统发育树图见图1,同时将ML树和BI树的后验概率值及自检举值标注在每一个分支上。系统发育结果显示,泰山黑蜈蚣叶(Phaeophyscia taishanensis)与白腹黑蜈蚣叶(P. denigrata)聚在了一支,支持率为89%,形成姐妹群,说明这两个种有着较近的亲缘关系;P. microsporaP. kashmirensi聚在了一支,支持率为92%,BI后验概率为1,形成姐妹群,与Niazi等(2023)的结果一致;泰山黑蜈蚣叶和白腹黑蜈蚣叶所在分支与P. microsporaP. kashmirensis所在分支形成姐妹分支,支持率为89%。

  • 2.2 分类单元

  • Phaeophyscia taishanensis Sh. H. Jiang &Z. F. Jia sp. nov. Fig.2

  • Fungal names: FN 571697

  • Diagnosis: This new species is distinctly separate from the comparable species Phaeophyscia denigrata in that the upper portion of the thalline margin of the apothecia occasionally have sparse cortical hairs, and the lower cortex is paraplectenchymatous, ascospores are only Physcia-type.

  • Etymology: The specific epithet ‘taishanensis’ refers to the type locality of Mount Tai, Shandong Province, China.

  • Type: China, Shandong, Tai’an City, Mount Tai, Buyun Bridge, 36°14′51″ N, 117°06′23″ E,940 m alt., on the bark of Ulmus pumila L., 8 November 2020, M.L. Zhu TS18074 (holotype, LCUF, GenBank OR769731) .

  • Description: Thallus foliose, nearly orbicular or irregular, more than 7 cm in diameter, closely adnate to the substratum; lobes dichotomously to irregularly branched, separate, sometimes imbricate, 0.6-1.1 mm wide, terminally obtuse; thallus 187.8-273.0 μm thick; upper cortex grey-green to olive-green, flat or often slightly concave, occasionally have white or hyaline cortical hairs on the upper surface of marginal peripheral zones of the lobes, which sometimes can be sparse, lacking maculation, soredia, isidia, or lobules; upper cortex 26.1-39.7 μm thick; algal layer continuous, 53.3-85.8 μm thick; medulla loose, white, 70.3-109.5 μm thick; lower surface black, sometimes white or pale brown on the lobe ends, rhizinate; rhizines dense, black, the ends are sometimes white, about 0.5 mm long; lower cortex paraplectenchymatous, 28.6-38.4 μm thick.

  • Apothecia common, sessile, usually 0.5-1.5 mm in diameter; disc red-brown to black, epruinose; margin entire; the upper portion of the thalline margin of the apothecia occasionally have sparse and countable cortical hairs, usually found in the immature apothecia, most of the apothecium do not have cortical hairs; epithecium brown, 10.7-13.3 μm thick; hymenium hyaline or slightly brown, 80.5-93.6 μm thick; hypothecium light brown, 26.5-30.3 μm thick; asci clavate, eight-spored; ascospores brown, Physcia-type, (18-20.5) μm × (9.0-10.0) μm, I+ blue; pycnidia common, immersed into thallus, brown to black; conidia ellipsoid, (2.6-3.3) μm × (0.9-1.7) μm.

  • Chemistry: Thallus K-; medulla K-; no substances were detected by TLC.

  • 特征提要:该新种与白腹黑蜈蚣叶相似,区别在于该新种盘托上部偶有白色或浅色的皮层毛,下皮层组织为假薄壁组织,子囊孢子仅为Physcia型。

  • 词源:种加词 ‘taishanensis’表示该物种被发现的产地山东泰山。

  • 主模式:山东泰安市,泰山,步云桥,117°06′23″ E、36°14′51″ N,海拔940 m,2020-11-08,朱孟丽TS18074。

  • 图1 基于ITS序列构建的最大似然系统发育树

  • Fig.1 Maximum likelihood tree illustrating the phylogeny based ITS sequences

  • 地衣体叶状,近圆形或不规则状,直径大于7 cm,较紧密地贴生于基物;裂片二叉至不规则分裂,分离,或有时重叠,0.6~1.1 mm宽,末端常钝圆;地衣体厚187.8~273.0 μm;上表面灰绿色至橄榄褐色,平坦或常常稍凹,末端具稀疏的皮层毛,无白斑、粉芽、裂芽和小裂片;上皮层厚26.1~39.7 μm;藻层厚53.3~85.8 μm;髓层疏松,白色,厚70.3~109.5 μm;下表面黑色,裂片末端处呈灰白色或浅褐色,具假根;假根单一不分枝,黑色,末端有时呈白色,约0.5 mm长;下皮层厚28.6~38.4 μm,皮层组织为假薄壁组织。

  • 子囊盘常见,贴生,直径通常0.5~1.5 mm;盘面红褐色至黑色,无粉霜;盘缘完整;盘托上部偶有白色或浅色的皮层毛,常常稀疏可数,见于未成熟子囊盘,大部分子囊盘不具皮层毛;囊层被褐色,10.7~13.3 μm;子实层清晰状,稍带褐色,80.5~93.6 μm;囊层基清晰状,浅褐色,26.5~30.3 μm;子囊内含有8个孢子;子囊孢子褐色,厚壁,Physcia型,为(18~20.5) μm × (9.0~10.0) μm,I+,蓝色;分生孢子器常见,埋生于地衣体中,孔口凸起于地衣体上表面,棕色至黑色,圆点状;分生孢子椭圆形,(2.6~3.3) μm × (0.9~1.7) μm。

  • 图2 泰山黑蜈蚣叶(M. L. Zhu TS18074,LCUF)

  • Fig.2 Phaeophyscia taishanensis (M. L. Zhu TS18074, LCUF)

  • 化学反应:地衣体K-;髓层K-;TLC未检测出地衣化学物质。

  • 分布:该种产自中国东部山东省中部,该地区地带性森林类型为落叶阔叶林,气候为温带季风气候,为北温带成分,其生长基物为榆树(Ulmus pumila L.)树皮。截至目前,该种仅在标本模式产地发现。

  • 其他研究标本:山东泰安市,泰山,步云桥,117°06′23″ E、36°14′51″ N,海拔940 m,2020-11-08,朱孟丽TS18071、TS18072、TS18073。

  • 3 讨论

  • 本种的主要特征是地衣体上表面末端具稀疏的皮层毛,髓层白色,下表面黑色,裂片末端处呈灰白色或浅褐色,盘托上部偶有白色或浅色的皮层毛,常常稀疏可数,子囊孢子褐色,厚壁,Physcia型,(18~20.5) μm × (9.0~10.0) μm。在形态及化学上,本种与皮层毛黑蜈蚣叶(Phaeophyscia hirtella)、白刺毛黑蜈蚣叶(P. hirtuosa)和P. esslingeri相似。本种与P. hirtellaP. hirtuosa相比,后两者子囊盘基部都常具有黑色的假根,而本种子囊盘基部无假根。与P. esslingeri相比,后者地衣体上表面有白斑,盘托上部皮层毛较多,孢子较宽,为 [(17~)18~25] μm × [10~12(~14)] μm(Kondratyuk et al.,2016);但本种地衣体上表面无白斑,盘托上部偶有白色或浅色的皮层毛,常常稀疏可数,孢子较窄。

  • 在系统发育上,本种亲缘关系与白腹黑蜈蚣叶(Phaeophyscia denigrata)、P. microsporaP. kashmirensis相近。本种与P. denigrata相比,后者地衣体下表面灰白色,盘托上部不具皮层毛,子囊孢子为Pachysporaria型或Physcia型;而本种地衣体下表面黑色,裂片末端处呈灰白色或浅褐色,果托上部偶有白色或浅色的皮层毛,子囊孢子仅为Physcia型;P. denigrata曾置于Phyciella Essl.中,下表面灰白色,皮层组织是假厚壁长细胞,通过对本种裂片进行横切、纵切观察发现,本种下表面皮层组织为假薄壁组织,对于搭建PhysciellaPhaoephyscia 之间“桥梁”关系具有重要意义,仍需进一步研究。与P. microspora相比,后者子囊盘基部有黑色假根,孢子较小,为(14~17)μm ×(6.0~8.0)μm,未见分生孢子器(Schumm &Aptroot,2019);而本种子囊盘基部没有假根,孢子较大,分生孢子器可见。与P. kashmirensis相比,后者地衣体边缘具有大量的粉芽堆(Fayyaz et al.,2022),未见分生孢子器;而本种无粉芽堆,分生孢子器可见。

  • 致谢 感谢中国科学院微生物研究所陈健斌研究员在标本鉴定方面以及本文内容提供宝贵建议,感谢本课题组硕士生朱孟丽协助采集标本!

  • 参考文献

    • AP TROOT A, SIPMAN HJM, 1991. New lichens and lichen records from New Guinea [J]. Willdenowia, 20: 221-256.

    • CASTRESANA J, 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis [J]. Mol Biol Evol, 17(4): 540-552.

    • CHEN JB, HU GR, 2022. The lichen family Physciaceae (Ascomycota) in China Ⅶ. Five species new to China [J]. Mycosystema, 41(1): 155-159. [陈健斌, 胡光荣, 2022. 中国蜈蚣衣科地衣Ⅶ. 五个新记录种 [J]. 菌物学报, 41(1): 155-159. ]

    • CHEN JB, 2023. Flora lichenum Sinicorum. Physciaceae: Vol. 20 [M]. Beijing: Science Press: 1-262. [陈健斌, 2023. 中国地衣志. 蜈蚣衣科: 第20卷 [M]. 北京: 科学出版社: 1-262. ]

    • CULBERSON CF, 1972. Improved conditions and new data for identification of lichen products by standardized thin-layer chromatographic method [J]. J Chromatogr A, 72(1): 113-125.

    • ESSLINGER T, 1978. Studies in the lichen family Physciaceae. Ⅲ. A new species of Phaeophyscia from Hawaii [J]. Mycologia, 70: 1247-1249.

    • FAYYAZ I, AFSHAN NS, KHALID AN, 2022. Phaeophyscia kashmirensis sp. nov. (Lecanorales, Physciaceae) from Azad Jammu and Kashmir, Pakistan [J]. Lichenologist, 54(6): 355-361.

    • GARDES M, BRUNS TD, 1993. ITS primers with enhanced specificity for basidiomycetes - application to the identification of mycorrhizae and rusts [J]. Mol Ecol, 2(2): 113-118.

    • GUINDON S, DUFAYARD JF, LEFORT V, et al. , 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3. 0 [J]. Syst Biol, 59(3): 307-321.

    • KALYAANAMOORTHY S, MINH BQ, WONG TKF, et al. , 2017. ModelFinder: fast model selection for accurate phylogenetic estimates [J]. Nat Method, 14(6): 587-589.

    • KATOH K, ASIMENOS G, TOH H, 2009. Multiple alignment of DNA sequences with MAFFT [J]. Method Mol Biol (Clifton, N. J. ), 537: 39-64

    • KONDRATYUK SY, LÖKÖS L, FARKAS E, et al. , 2016. Phaeophyscia esslingeri sp. nov. (Physciaceae, lichen-forming Ascomycota) — a new lichen species from Eastern Asia, with a world-wide key to the hairy species of the genus [J]. Stud Bot Hung, 47(2): 251-262.

    • LIU D, HUR JS, 2019. Revision of the lichen genus Phaeophyscia and allied atranorin absent taxa (Physciaceae) in South Korea [J]. Microorganisms, 7(8): 242.

    • MINH BQ, NGUYEN MAT, VON HAESELER A, 2013. Ultrafast approximation for phylogenetic bootstrap [J]. Mol Biol Evol, 30(5): 1188-1195.

    • MOBERG R, 1977. The lichen genus Physcia and allied genera in Fennoscandia [J]. Symbolae Bot Upsalienses, 22: 1-108.

    • NGUYEN LT, SCHMIDT HA, VON HAESELER A, et al. , 2015. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies [J]. Mol Biol Evol, 32(1): 268-274.

    • NIAZI AR, AFSHAN NS, NASEER A, et al. , 2023. A new species and a new record of the genus Phaeophyscia Moberg (Lecanorales, Physciaceae) from Pakistan supported by phenotypic and molecular phylogenetic analyses [J]. Cryptog Mycol, 44(4): 51-59.

    • ORANGE A, JAMES PW, WHITE FJ, 2001. Microchemical methods for the identification of lichens [M]. London: British Lichen Society: 1-101.

    • RONQUIST F, TESLENKO M, VAN DER MARK P, et al. , 2012. MrBayes 3. 2: efficient bayesian phylogenetic inference and model choice across a large model space [J]. Syst Biol, 61(3): 539-542.

    • SCHUMM F, APTROOT A, 2019. Virtuelles Herbarium der Flechtengattungen Hyperphyscia, Phaeophyscia, Physcia und Physconia [M]. Norderstedt: Books on Demand: 1-632.

    • TALAVERA G, CASTRESANA J, 2007. Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments [J]. Syst Biol, 56(4): 564-577.

    • WEI JC, 2020. The enumeration of lichenized fungi in China [M]. Beijing: China Forestry Publishing House: 1-606. [魏江春, 2020. 中国地衣型真菌综览 [M]. 北京: 中国林业出版社: 1-606. ]

    • WHITE TJ, BRUNS T, LEE S, et al. , 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics [M]// INNIS MA, GELFAND DH, SNINSKY JJ, et al. PCR Protocols. San Diego: Academic Press: 315-322.

    • WIJAYAWARDENE NN, HYDE KD, DAI DQ, et al. , 2022. Outline of fungi and fungus-like taxa - 2021 [J]. Mycosphere, 13(1): 53-453.

    • ZHANG D, GAO F, JAKOVLIĆ I, et al. , 2020. PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies [J]. Mol Ecol Resourc, 20(1): 348-355.

    • ZHAO YF, JIA ZF, 2022. Astrothelium subsiamense sp. nov. from Fujian, China [J]. Mycotaxon, 137(3): 477-484.

  • 参考文献

    • AP TROOT A, SIPMAN HJM, 1991. New lichens and lichen records from New Guinea [J]. Willdenowia, 20: 221-256.

    • CASTRESANA J, 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis [J]. Mol Biol Evol, 17(4): 540-552.

    • CHEN JB, HU GR, 2022. The lichen family Physciaceae (Ascomycota) in China Ⅶ. Five species new to China [J]. Mycosystema, 41(1): 155-159. [陈健斌, 胡光荣, 2022. 中国蜈蚣衣科地衣Ⅶ. 五个新记录种 [J]. 菌物学报, 41(1): 155-159. ]

    • CHEN JB, 2023. Flora lichenum Sinicorum. Physciaceae: Vol. 20 [M]. Beijing: Science Press: 1-262. [陈健斌, 2023. 中国地衣志. 蜈蚣衣科: 第20卷 [M]. 北京: 科学出版社: 1-262. ]

    • CULBERSON CF, 1972. Improved conditions and new data for identification of lichen products by standardized thin-layer chromatographic method [J]. J Chromatogr A, 72(1): 113-125.

    • ESSLINGER T, 1978. Studies in the lichen family Physciaceae. Ⅲ. A new species of Phaeophyscia from Hawaii [J]. Mycologia, 70: 1247-1249.

    • FAYYAZ I, AFSHAN NS, KHALID AN, 2022. Phaeophyscia kashmirensis sp. nov. (Lecanorales, Physciaceae) from Azad Jammu and Kashmir, Pakistan [J]. Lichenologist, 54(6): 355-361.

    • GARDES M, BRUNS TD, 1993. ITS primers with enhanced specificity for basidiomycetes - application to the identification of mycorrhizae and rusts [J]. Mol Ecol, 2(2): 113-118.

    • GUINDON S, DUFAYARD JF, LEFORT V, et al. , 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3. 0 [J]. Syst Biol, 59(3): 307-321.

    • KALYAANAMOORTHY S, MINH BQ, WONG TKF, et al. , 2017. ModelFinder: fast model selection for accurate phylogenetic estimates [J]. Nat Method, 14(6): 587-589.

    • KATOH K, ASIMENOS G, TOH H, 2009. Multiple alignment of DNA sequences with MAFFT [J]. Method Mol Biol (Clifton, N. J. ), 537: 39-64

    • KONDRATYUK SY, LÖKÖS L, FARKAS E, et al. , 2016. Phaeophyscia esslingeri sp. nov. (Physciaceae, lichen-forming Ascomycota) — a new lichen species from Eastern Asia, with a world-wide key to the hairy species of the genus [J]. Stud Bot Hung, 47(2): 251-262.

    • LIU D, HUR JS, 2019. Revision of the lichen genus Phaeophyscia and allied atranorin absent taxa (Physciaceae) in South Korea [J]. Microorganisms, 7(8): 242.

    • MINH BQ, NGUYEN MAT, VON HAESELER A, 2013. Ultrafast approximation for phylogenetic bootstrap [J]. Mol Biol Evol, 30(5): 1188-1195.

    • MOBERG R, 1977. The lichen genus Physcia and allied genera in Fennoscandia [J]. Symbolae Bot Upsalienses, 22: 1-108.

    • NGUYEN LT, SCHMIDT HA, VON HAESELER A, et al. , 2015. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies [J]. Mol Biol Evol, 32(1): 268-274.

    • NIAZI AR, AFSHAN NS, NASEER A, et al. , 2023. A new species and a new record of the genus Phaeophyscia Moberg (Lecanorales, Physciaceae) from Pakistan supported by phenotypic and molecular phylogenetic analyses [J]. Cryptog Mycol, 44(4): 51-59.

    • ORANGE A, JAMES PW, WHITE FJ, 2001. Microchemical methods for the identification of lichens [M]. London: British Lichen Society: 1-101.

    • RONQUIST F, TESLENKO M, VAN DER MARK P, et al. , 2012. MrBayes 3. 2: efficient bayesian phylogenetic inference and model choice across a large model space [J]. Syst Biol, 61(3): 539-542.

    • SCHUMM F, APTROOT A, 2019. Virtuelles Herbarium der Flechtengattungen Hyperphyscia, Phaeophyscia, Physcia und Physconia [M]. Norderstedt: Books on Demand: 1-632.

    • TALAVERA G, CASTRESANA J, 2007. Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments [J]. Syst Biol, 56(4): 564-577.

    • WEI JC, 2020. The enumeration of lichenized fungi in China [M]. Beijing: China Forestry Publishing House: 1-606. [魏江春, 2020. 中国地衣型真菌综览 [M]. 北京: 中国林业出版社: 1-606. ]

    • WHITE TJ, BRUNS T, LEE S, et al. , 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics [M]// INNIS MA, GELFAND DH, SNINSKY JJ, et al. PCR Protocols. San Diego: Academic Press: 315-322.

    • WIJAYAWARDENE NN, HYDE KD, DAI DQ, et al. , 2022. Outline of fungi and fungus-like taxa - 2021 [J]. Mycosphere, 13(1): 53-453.

    • ZHANG D, GAO F, JAKOVLIĆ I, et al. , 2020. PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies [J]. Mol Ecol Resourc, 20(1): 348-355.

    • ZHAO YF, JIA ZF, 2022. Astrothelium subsiamense sp. nov. from Fujian, China [J]. Mycotaxon, 137(3): 477-484.