Academia.eduAcademia.edu
植 物 分 类 与 资 源 学 报 Plant Diversity and Resources 2014ꎬ 36 (6) : 781 ~ 787 DOI: 10.7677 / ynzwyj201413154 中国横断山的地衣研究 ( 6) : 灰梅属地衣修订 刘 ∗ ∗∗ 栋1ꎬ2 ꎬ 王欣宇2 ꎬ 李建文1ꎬ2 ꎬ 钱子刚1 ꎬ 王立松2 (1 云南中医学院ꎬ 云南 昆明 650500ꎻ 2 中国科学院昆明植物研究所生物多样性 与生物地理学重点实验室ꎬ 云南 昆明 650201) 摘要: 通过对灰梅属 ( Canoparmelia) 表型与基因型相结合的研究ꎬ 澄清了云南南部报道的珊瑚大叶梅 ( Parmotrema conformatum) 是裂芽灰梅 ( C� ecaperata) 的错误鉴定ꎮ 由于珊瑚大叶梅和裂芽灰梅在经典分 类鉴别上容易混淆ꎬ 通过 rDNA ITS 序列的系统发育分析ꎬ 证明了裂芽灰梅与粉芽灰梅 ( C� texana) 及同 心灰梅 ( C� concrescens) 有较近的亲缘关系ꎻ 首次记录了粉芽灰梅在云南的分布范围ꎻ 文中对两个新记录 种进行了详细描述和讨论ꎬ 并给出了中国已知灰梅属的检索表ꎮ 关键词: 灰梅属ꎻ 新记录ꎻ 地理分布ꎻ 分子系统学 中图分类号: Q 949� 34 文献标识码: A 文章编号: 2095-0845(2014)06-781-07 Contributions to the Lichen Flora of the Hengduan Mountainsꎬ China (6) : Revisional Study of the Genus Canoparmelia ( Lichenized Ascomycotaꎬ Parmeliacaea) ∗ LIU Dong 1ꎬ2 ꎬ WANG Xin ̄Yu 2 ꎬ LI Jian ̄Wen 1ꎬ2 ꎬ QIAN Zi ̄Gang 1 ꎬ WANG Li ̄Song 2∗∗ (1 Yunnan University of Traditional Chinese Medicineꎬ Kunming 650500ꎬ Chinaꎻ 2 Key Laboratory of Biodiversity and Biogeographyꎬ Kunming Institute of Botanyꎬ Chinese Academy of Sciencesꎬ Kunming 650201ꎬ China) Abstract: A revision of Canoparmelia from the Hengduan Mountains ( Yunnanꎬ China) revealed the occurrence of C� ecaperataꎬ based on specimens previously identified as Parmotrema conformatum. C� ecaperata is hereby reported as new to the flora of China. The study also led to the identification of C� texana as a component of the flora of Yun ̄ nan. In order to make sure the systematic position of these speciesꎬ a phylogenetic analysis of rDNA ITS sequence was also carried out in this paperꎬ indicating that C� ecaperata is closely related to C� texana and C� concrescens. De ̄ tailed information and discussion of these two new records are providedꎬ together with a key to all the known taxa of Canoparmelia recorded from China. Following our revisionꎬ Parmotrema conformatum should be excluded from the flora of China. Key words: Canoparmeliaꎻ New recordꎻ Geographic distributionꎻ Molecular phylogeny Canoparmelia Elix & Hale ( Parmeliaceaeꎬ Le ̄ segregated by Elix and Hale from Pseudoparmelia on ( Hawksworth et al.ꎬ 1995ꎻ Thell et al.ꎬ 2012) and and chemical characters ( Elix et al.ꎬ 1986) . Four canoralesꎬ Ascomycota ) comprises ca. 40 species is typified by C� texana ( Tuck.) Elix & Hale. It was the basis of morphologicalꎬ geographicꎬ ecologicalꎬ species of Canoparmelia were previously known from ∗ Funding: The National Natural Science Foundation of China ( No. 31170023ꎬ Y3216111W1) ꎬ Foundation of Key Laboratoryꎬ CAS ( KLBB ̄ 201210) and Flora Lichenum Sinicorum ( KSCX2 ̄EW ̄Z ̄9) ∗∗ Author for correspondenceꎻ E ̄mail: wanglisong@ mail� kib� ac� cn Received date: 2013-07-22ꎬ Accepted date: 2013-08-05 作者简介: 刘 栋 (1988-) 男ꎬ 硕士ꎬ 主要从事地衣分类学研究ꎮ E ̄mail: liudong@ mail� kib� ac� cn 782 植 物 分 类 与 资 源 学 报 第 36 卷 China: two from Taiwan ( Weiꎬ 1991ꎻ Laiꎬ 2000) ꎬ for 5 minꎬ followed by 30 cyclesꎬ each of three steps As part of an inventory of the lichen flora of the and a final extension at 72 ℃ for 7 min. and three from mainland China ( Jia et al.ꎬ 2008) . Hengduan Mountains ( Yunnanꎬ China) several spe ̄ (94 ℃ for 1 minꎬ 56 ℃ for 1 minꎬ 72 ℃ for 1� 5 min)ꎬ Twenty ̄seven sequences (18 retrieved from Gen ̄ cimens of Canoparmelia were collected. These be ̄ Bank and 9 obtained by authorsꎬ Table 1) were as ̄ cord to the flora of Chinaꎬ and to C� texanaꎬ which sequence database was subjected to automated align ̄ long to C� ecaperata Elix & Haleꎬ which is a new re ̄ is new to Yunnan Province. All specimens of Par ̄ motrema conformatumꎬ previously reported from Yun ̄ nan were misidentified and belong to C� ecaperata. 1 Materials and Methods The specimens were mainly collected from Yun ̄ nan Provinceꎬ and were deposited in KUN ( Herbar ̄ ium of Kunming Institute of Botany) . External mor ̄ phological descriptions were based on air ̄dried ma ̄ terialsꎬ and observation under stereomicroscopy (Don ̄ gwon OSM ̄1) . Chemical characteristics were exami ̄ ned by medullary color reactionꎬ with the solution for the color reaction following the methods described by Yoshimura (1974) : K ( 10% aqueous KOH so ̄ lution) ꎬ C ( NaClO solution) ꎬ KC (10% KOH fol ̄ lowed by C solution) ꎬ P ( 5% alcoholic P ̄pheny ̄ lenediamine) . Thin layer chromatography ( TLC) of acetone extracts was also carried out to further char ̄ acterize the chemical profile of the specimensꎬ using solvent systems C and G ( Culberson and Hör ̄Durꎬ 1970ꎻ Orange et al.ꎬ 2001 ) and Lethariella cla ̄ donioides ( Nyl.) Krog as control sample. DNA was extracted from freshly collected and frozen herbarium specimens grinded to a fine pow ̄ derꎬ using Axyprep Multisource Genomic DNA Mini ̄ prep Kit following the manufacturer’ s instructions. The nuclear ITS ( rDNA repeat) was amplified using primers ITS1 and ITS4 ( Whiteꎬ 1990ꎻ Gardes and Brunsꎬ 1993) in 25 μL reaction volume composed of 2 μL DNAꎬ 1 μL of each primerꎬ 12� 5 μL 2 × Taq PCR Mastermix (Aidlab) (contain Taq DNA Polymer ̄ ase: 0� 1 unit�μL-1 ꎻ MgCl2 : 4 mmol�L-1 ꎻ dNTPs: 0� 4 mmol�L -1 ) ꎬ 8� 5 μL dH 2 O. Amplifications were carried out in an automatic thermocycler (C 1000 )ꎬ TM with the following profile: initial denaturation at 94 ℃ sembled into a matrix for phylogenetic analyses. The ment and then manually inspected in BioEdit. Hypo ̄ trachynia adducta was set as the outgroup. Ambigu ̄ ous regions were deleted. Bayesian inference of phy ̄ logeny was performed with in MrBayes v3� 1� 2. Mod ̄ elTest 3� 7 was used to estimate separate best ̄fit models of evolution for ITS. Bayesian analyses were carried out using the Metropoliscoupled Markov chain Monte Carlo method ( MCMCMC ) . Analyses were run under the GTR model using a gamma ̄dis ̄ tributed rate parameter and a proportion of invariable sites. Two parallel MCMCMC runs were performedꎬ each using four chains and 1 million generationsꎬ sampling trees of every 100th generation. The initial burn ̄in was set to 50%. A 50% majority ̄rule con ̄ sensus cladogram was computed from the remaining treesꎻ the proportions of this tree correspond to Bayesian posterior probabilities ( BPP) ꎬ and clades with PP ≥0� 95 were considered to be significantly supported. The phylogenic trees are shown in Fig� 1. 2 2� 1 Result Key to the species of known Canoparmelia in China 1. Thallus without soredia or isidia ���� C� subtiliacea 1. Thallus sorediate or isidiate 2. Thallus sorediate 3. Medulla P+ yellowꎬ containing stictic and constictic acid acid �������������� C� crozalsiana 3. Medulla P-ꎬ containing divaricatic acid ����������������� C� texana 2. Thallus isidiate 4. Medulla P+ redꎬ containing protocetraric acid ��������������� C� amazonica 4. Medulla P-ꎬ containing divaricatic and usnic acid ����������������� C� ecaperata 6期 LIU Dong et al.: Contributions to the Lichen Flora of the Hengduan Mountainsꎬ China (6) : Revisional � Species name Canoparmelia concrescens C� ecaperata C� ecaperata Accession NO. Source GU994543 Kenya GenBank F� papillosa KC978844 China 12 ̄33440 F� springtonensis KC978843 KC978846 C� ecaperata GU994545 C� nairobiensis KC978847 C� texana KC978848 C� texana C� texana Flavoparmelia baltimorensis F� caperata China 12 ̄33611 China GenBank China 12 ̄37655 China 12 ̄37665 USA GenBank Spain AY586559 12 ̄34474 Kenya AY581059 JN943840 Fig� 1 10 ̄31934 Kenya HM010933 F� haywardiana F� marchantii China GU994547 GU994550 F� citrinescens Species and specimens of Parmeliaceae analyzed Location KC978845 C� ecaperata Table 1 GenBank Argentina New Zealand — GenBank GenBank GenBank GenBank Species name F� soredians F� subambigua F� subambigua Hypotrachynia adducta Parmotrema subtinctorium P� norsticticatum P� reticulatum P� subcaperatum P� subtinctorium P� tinctorum P� tinctorun Accession NO. Location Source HM010936 Uruguay GenBank EF042907 Australia GenBank AY586562 GU994551 Spain Argentina GenBank GenBank JN943839 Argentina KC978853 China 12 ̄35202 Portugal GenBank China GenBank China 12 ̄33444 KC978855 China GU994577 South Africa AY586557 Australia AY586570 Australia AY586579 GU593037 KC978854 Phylogenic construction of the systematic position of C� ecaperata based on rDNA sequence date. ▲ Sequences obtained by author 783 GenBank 12 ̄34288 GenBank GenBank GenBank 784 2� 2 植 物 分 类 与 资 源 学 报 Taxonomy Canoparmelia ecaperata Elix & Hale in Mycotaxon 27: 277ꎬ 1986. ———Parmelia ecaperata Müll. Arg. in Floraꎬ Jena 74: 378. 1891. ———Pseudoparmelia ecape ̄ rata (Müll. Arg.) Hale in Phytologia 29: 190ꎬ 1974. Diagnostic characters: Thallus folioseꎬ adnate on barkꎬ grayish yellow to yellow ̄greenꎬ 4 - 14 cm in diam.ꎻ lobes irregularly sublinearꎬ 1 - 3 mm wideꎬ without ciliaꎻ upper surface planeꎬ continuousꎬ some ̄ times cracked on the old partꎻ covered with dense isidiaꎬ simple and shortꎬ 0� 1 - 1 mm longꎬ cylindri ̄ cal or clavateꎬ mature ones with brown to black tipsꎻ medulla whiteꎬ 62� 5 - 75 μm thickꎻ lower surface Fig� 2 第 36 卷 blackꎬ turning brownish near the marginꎬ rhizines rareꎬ simple and blackꎬ 0� 5 - 1� 5 mm longꎻ lower cortex 25-75 μm thickꎬ Apothecia not seen (Fig� 2). Chemistry: Cortex K ̄ꎬ medulla K ̄ꎬ C ̄ (pink)ꎬ KC ̄ꎬ P ̄ꎻ atranorinꎬ divaricatic acidꎬ usnic acid and an unknown compound (Rf value between classes 4 and 5). Distribution: China: Yunnan Provinceꎬ Nanjian Co. ( Fig� 3)ꎻ Africaꎬ Nepalꎬ Indiaꎬ Thailand ( Hale and Masonꎬ 1976ꎻ Singh and Sinhaꎬ 2010ꎻ Swins ̄ cow et al.ꎬ 1988ꎻ Wolseley et al.ꎬ 2002) . Habitat and ecology: This species grows on the bark of Pinus yunnanensis from 1 300 m to 2 100 m in Nanjian Co.ꎬ Yunnan. Canoparmelia ecaperata. A. Habitatꎻ B. Cylindrical laminal isidia. C. Narrow and sublinear lobes. ( All based on Li S. Wang 12-37823) 6期 LIU Dong et al.: Contributions to the Lichen Flora of the Hengduan Mountainsꎬ China (6) : Revisional � Fig� 3 785 Distribution map of C� ecaperata ( left) and C� texana ( right) from Hengduan Mountains Remarks: Canoparmelia ecaperata is similar to Parmotrema conformatum and Flavoparmelia caperata. The former differs from C� ecaperata in the presence of fumarprotocetraric and protocetraric acids ( P + red) ꎬ ciliate lobes ( Chen et al.ꎬ 2005ꎻ Benatti and Marcelliꎬ 2009). Moreoverꎬ C� ecaperata occurs mainly on Pinusꎬ Wang Lisong et al. 12 - 33440ꎬ 30 March 2012ꎻ Langcang Co.ꎬ Mengla Villageꎬ on Pinusꎬ alt. 1 100 mꎬ Wang Lisong 10 - 31934ꎬ 11 Dec. 2010ꎻ Pu’er Cityꎬ Ning’er Co.ꎬ nearby road 213ꎬ N: 22°53′55� 44″ꎬ E: 101°02′28� 41″ꎬ alt. 1 334 mꎬ on barkꎬ Wang Lisong et al. 12 - 34474ꎬ 21 June in the tropical area of the Southern Hemisphere ( El ̄ 2012ꎻ Pu’er Cityꎬ N: 22� 48°ꎬ E: 100� 58°ꎬ Zhao whereas C� ecaperata has been reported from India Nov. 1960ꎻ Kunming Cityꎬ Heilongtanꎬ N: 25� 04°ꎬ ix and McCarthyꎬ 1998ꎻ Nöske and Sipmanꎬ 2004)ꎬ and Thailand ( Haleꎬ 1976) . Flavoparmelia caperata differs in the sorediate rather than isidiate lobesꎬ and the presence of protocetraric acid rather than atra ̄ Ji ̄ding & Chen Yu ̄ben 2942ꎬ HMAS 100308ꎬ 18 E: 102� 42°ꎬ Zhao Ji ̄ding & Chen Yu ̄ben 4748ꎬ HMAS 100307ꎬ 16 Dec. 1960. norin as the main compound. Canoparmelia concres ̄ Canoparmelia texana (Tuck.) Elix & Hale in Myco ̄ tinguished by the bright yellow thallus ( containing man in American Journal of Science and Arts series cens might be confused with C� ecaperata but is dis ̄ taxon 27: 278ꎬ 1986. ———Parmelia texana Tucker ̄ usnic acid in the cortex) ꎬ and its distribution only in 2 ( 253) : 424ꎬ 1858. ———Pseudoparmelia texana can be easily distinguished by the lack of usnic acid Diagnostic characters: Thallus grayꎬ upper sur ̄ Africa. Finally C� amazonicaꎬ which also has isidia ( Tuckerman) Hale in Phytologia 28: 191ꎬ 1974. and spare rhizines. face covered with dense soraliaꎬ punctiform or capi ̄ jian Co.: Leqiu Villageꎬ N: 25°00′25� 29″ꎬ E: 100° ines simpleꎻ containing atranorin and divaricatic acid. Specimens examined: Yunnan Province: Nan ̄ 21′43� 50″ꎬ alt. 2 060 mꎬ on Pinus yunnanensis barkꎬ Wang Lisong & Wang Xinyu 12 - 37814ꎬ 12 - 37820ꎬ 12 - 37822ꎬ 12 - 37823ꎬ 21 Dec. 2012ꎻ Gonglang Villageꎬ N: 24°43� 103′ꎬ E: 100°25� 514′ꎬ alt. 1 840 mꎬ on Pinus yunnanensis barkꎬ Wang Lisong et al. 12 - 33611ꎬ 25 March 2012. Longling Co.ꎬ Songshan Mt.ꎬ N: 24°44� 672′ꎬ E: 98°54� 281′ꎬ alt. 2 000 mꎬ tateꎻ lower surface blackꎻ moderately rhizinateꎬ rhiz ̄ Distribution: Pantemperate outside of Europe and pantropical at higher elevations. Shandong and Taiwan Province in China ( Jiaꎬ 2008ꎻ Laiꎬ 2000) ꎻ New to Yunnan Province ( Fig� 3) . Specimens examined: Yunnan Provinceꎬ Nan ̄ jian Co.: Leqiu Village. N: 25°00′25� 29″ꎬ E: 100° 21′43� 50″ꎬ alt. 2 060 mꎬ on Pinus yunnanensis barkꎬ 786 第 36 卷 植 物 分 类 与 资 源 学 报 Wang Lisong & Wang Xinyu 12 - 37805ꎬ 12 - 37810ꎬ 12- 37889ꎬ 21 Dec. 2012ꎻ Wuliang Medicine Valleyꎬ N: 24°52′2� 76″ꎻ E: 100°34′51� 39″ꎬ alt. 2 345 mꎬ on Pinus amandii barkꎬ Wang Lisong & Wang Xinyu Thallus greyꎬ upper surface without soredia or isidiaꎬ rhizines rare. Containing atranorin and caper ̄ atic acid ( Haleꎬ 1976) . 12 - 37655ꎬ 19 Dec. 2012ꎻ Hongxing Villageꎬ Wu ̄ 3 38′4� 52″ꎬ alt. 2 341 mꎬ on barkꎬ Wang Lisong & Wang Xinyu 12 - 36923ꎬ 16 Oct. 2012ꎻ Gonglang and Canoparmelia are similar in external morphology 24� 839′ꎬ alt. 2 270 mꎬ on barkꎬ Wang Lisong et al. 12 - 33742ꎬ 25 March 2012ꎻ Ziqiang Villageꎬ N: this study to verify the systematic position of these liang Medicine Valleyꎬ N: 24°50′40� 41″ꎬ E: 100° Countryꎬ Jinguo Villageꎬ N: 24°48� 557′ꎬ E: 100° 24°46′10� 55″ꎻ E: 100°25′08� 13″ꎬ alt. 1 640 mꎬ on barkꎬ Wang Lisong & Wang Xinyu 12 - 37674ꎬ 19 Dec. 2012ꎻ Ziqiang Villageꎬ N: 24°47′21� 60″ꎻ E: 100°25′07� 78″ꎬ alt. 1 740 mꎬ on barkꎬ Wang Li ̄ song & Wang Xinyu 12 - 37665ꎬ 19 Dec. 2012ꎻ Da ̄ dianshan Mt.ꎬ N: 24°51′48″ꎬ E: 100°34′39″ꎬ alt. 2 543 mꎬ on branchꎬ Wang Lisong et al. 12 - 32957ꎬ 21 March 2012. Canoparmelia amazonica ( Nyl.) Elix & Hale in Mycotaxon 27: 278ꎬ 1986 Thallus greyꎬ upper surface covered with rare branched isidiaꎬ rhizines dense. Containing atranorin and protocetraric acid ( Haleꎬ 1976) . Canoparmelia crozalsiana ( Bouly de Lesdain) Elix & Hale in Mycotaxon 27: 278ꎬ 1986 Thallus greenish or grayꎬ sorediateꎬ moderately rhizinate. Containing atranorinꎬ stictic acidꎬ constic ̄ tic acid ( Haleꎬ 1976) . Canoparmelia subtiliacea ( Nyl.) Elix & Hale in Mycotaxon 27: 279ꎬ 1986 Table 2 Thallus color Lobes Lower surface Rhizines Characteristic compounds Distribution Discussion Since the genera Flavoparmeliaꎬ Pamotremaꎬ ( Table 2 ) and therefore hard to distinguish from each otherꎬ the phylogenetic method was applied in taxa and validate the morphological identification. The genus Parmotrema is characterized by broad lobes ( usually more than 5 mm) and rareꎬ simple rhizines in the central part of the lower surfaceꎬ up ̄ per surface grayish greenꎬ without usnic acid ( Chen et al.ꎬ 2005 ) ꎻ Flavoparmelia is characterized by broad and roundish lobesꎬ containing usnic acid on the upper surface so that the color is yellow ̄green ( Haleꎬ 1986 ) ꎻ Canoparmelia has sublinear and rather narrow lobesꎬ closely attached to the sub ̄ strateꎬ has a subtropical to tropical distributionꎬ and always contains atranorin as the main compound ( Elix et al.ꎬ 1986) . Parmotrema conformatum was previously repor ̄ ted from Yunnan Province by Zhao et al. ( 1982) ꎬ basedꎬ howeverꎬ on two specimens ( HMAS 100307 100308) that lacked cilia and had P ̄ medullary re ̄ action. Careful examination of the two specimens re ̄ vealed that they belong to C� ecaperata according to morphologicalꎬ chemicalꎬ and distributional charac ̄ ters. This is further supported by inferences from mo ̄ lecular data whereby both specimens belong to Cano ̄ parmelia and to C� ecaperata in particular. Key differences among Canoparmeliaꎬ Flavoparmelia and Parmotrema Flavoparmelia Parmotrema Canoparmelia Yellowish ̄green Grey to greenish Grey to yellowish Broad (2-6 mm) Blackꎬ with a narrowꎬ brownꎬ naked marginal zone Sparse to moderateꎬ simple or branched Usnic acid Temperate Rather broad (5-10 mm) Medium (0� 5-5 mm) Sparse and simpleꎬ in the thallus center Moderate to denseꎬ Simple or tufted Brown to blackꎬ with broad naked zone. Atranorin Temperate to tropical Blackꎬ narrowꎬ brownꎬ marginal zone ( <1 mm) Atranorinꎬ usnic acid Subtropical to tropical 6期 LIU Dong et al.: Contributions to the Lichen Flora of the Hengduan Mountainsꎬ China (6) : Revisional � The phylogenetic tree showed a final alignment consisting of 480 charactersꎬ of which 281 characters are constantꎬ 31 variable parsimony ̄uninformativeꎬ and 168 parsimony ̄informative. All characters have equal weight. Gaps are treated as missing. The GC content of C� ecaperata is 54� 7%ꎬ lower than C� texana (56� 2%) . The molecular data showed that the phy ̄ logenetic relationship of C� ecaperata is closer to Canoparmeliaꎬ which is in accord with our morpho ̄ logical identification. The final result confirms the placement of this species in the Canoparmelia clade. In this analysisꎬ C� ecaperata is recovered as sister group of C� texana and C� concrescens with high support. In conclusionꎬ the presence of two taxa of Canop ̄ armelia has been confirmed from the Hengduan Mountains based on morphologicalꎬ chemical and phy ̄ logenetic study: C� ecaperata is reported as new to Chi ̄ na and C� texana as new to Yunnan. The previous re ̄ cord of Parmotrema conformatum from China is based on misidentification ( = C� ecaperata ) which implies that the species should be deleted from the flora of China. The study added to the lichen biodiversity of the Hengduan Mountainsꎬ and also showed that there is still a high potential for new discoveries in this region. Acknowledgment: We are grateful to Prof. Bernard Goffinet (University of Connecticutꎬ USA) and Prof. Schmidt ̄Vogt Di ̄ etrich (KIB) for reviewing the manuscriptꎬ Ms. Deng Hong for helping with the original specimens studyꎬ Dr. Li Guodongꎬ Dr. Yu Wenbinꎬ Tang Kemin and Liang Mengmeng for their help with the molecular analysisꎬ Dr. Ma Wenzhang for his as ̄ 787 tification of lichen products [ J] . Journal of Chromatography Aꎬ 46: 85—93 Elix JAꎬ Johnston Jꎬ Verdon Dꎬ 1986. Canoparmeliaꎬ Paraparmelia and Relicinopsisꎬ three new genera in the Parmeliaceae ( lichen ̄ ized Ascomycotina) [ J] . Mycotaxonꎬ 27: 271—282 Elix JAꎬ McCarthy PMꎬ 1998. Catalogue of the Lichens of the Smaller Pacific Islands [ M] . Bibliotheca Lichenologicaꎬ 70ꎬ J. Cramerꎬ Stuttgartꎬ 361 Gardes Mꎬ Bruns Tꎬ 1993. ITS primers with enhanced specificity for basidiomycetes ̄application to the identification of mycorrhizae and rusts [ J] . Molecular Ecologyꎬ 2 (2) : 113—118 Hale MEꎬ 1976. A monograph of the lichen genus Pseudoparmelia Lynge ( Parmeliaceae) [ J] . Smithsonian Contributions to Bota ̄ nyꎬ 31: 1—62 Hale MEꎬ 1986. Flavoparmeliaꎬ a new genus in the lichen family Parmeli ̄ aceae (Ascomycotina) [J]. Mycotaxonꎬ 25 (2) : 603—605 Hawksworth DLꎬ Kirk PMꎬ Cannon PF et al.ꎬ 1995. Ainsworth & Bis ̄ by’s Dictionary of the Fungi [ M] ꎬ 8th ed. Wallingfored: CAB Internationalꎬ 1—616 Jia ZFꎬ Li YLꎬ Hou J et al.ꎬ 2008. A study of lichen genus Canoparme ̄ lia from Taishan Mountain [J]. Mycosystemaꎬ 27 (3): 461—463 Lai MZꎬ 2000. Illustrated Macrolichens of Taiwan I [ M] . Taipei: Council of Agricultureꎬ 106—107 Nöske NMꎬ Sipman HJMꎬ 2004. Cryptogams of the reserva biológica san francisco ( Province Zamora ̄Chinchipeꎬ Southern Ecuador) II. Lichens [ J] . Cryptogamieꎬ Mycologieꎬ 25 (1) : 91—100 Orange Aꎬ James PWꎬ White FJ et al.ꎬ 2001. Microchemical Methods for the Identification of Lichens [ M] . London: British Lichen So ̄ cietyꎬ 1—101 Singh Kꎬ Sinha Gꎬ 2010. Indian Lichens: an Annotated Checklist [ M] . India: Botanical Survey of Indiaꎬ 1—572 Swinscow Tꎬ Douglas Vꎬ Krog Hꎬ 1988. Macrolichens of East Africa [ M] . London: British Museum ( Natural History) ꎬ 1—383 Thell Aꎬ Crespo Aꎬ Pradeep K et al.ꎬ 2012. A review of the lichen family Parmeliaceae ̄historyꎬ phylogeny and current taxonomy [ J] . Nordic Journal of Botanyꎬ 30: 641—664 sistance during our field workꎬ Prof. Du Zhizhi and Zhang Wei JCꎬ 1991. An Enumeration of Lichens in China [ M] . Beijing: also like to thank Yunnan Wei ̄he Pharmaceutical Co.ꎬ Ltd. White TJꎬ 1990. Amplification and direct sequencing of fungal riboso ̄ Hongxia for their support with the chemical analysis. We would (云南维和药业股份有限公司) for the financial support. References: Benatti MNꎬ Marcelli MPꎬ 2009. Species of Parmotrema ( Parmeli ̄ aceaeꎬ Ascomycota) of the central ̄southern coast of São Paulo Etateꎬ Brazil: I. Gyrophoric and lecanoric chemical group [ J] . Acta Botanica Brasilicaꎬ 23 (4) : 1013—1026 Chen JBꎬ Wang SLꎬ Elix JAꎬ 2005. Parmeliaceae ( Ascomycota) li ̄ chens in China′s mainland III. The genus Parmotrema [ J] . My ̄ cotaxonꎬ 91: 93—114 Culberson CFꎬ Hör ̄Dur Kꎬ 1970. A standardized method for the iden ̄ International Academic Publishers Ltd.ꎬ 1—278 mal RNA genes for phylogenetics [ A] . In: Innis MAꎬ Gelfand DHꎬ Sninsky JJ et al. ( eds.) ꎬ PCR Protocols [ M] . San Diego: Academic Pressꎬ 315—322 Wolseley Yꎬ Patricia Aꎬ Aguirre ̄Hudson B et al.ꎬ 2002. Catalogue of the lichens of Thailand [ J] . Bulletin of the Natural History Muse ̄ um: Botanyꎬ 32 (01) : 13—59 Yoshimura Iꎬ 1974. Lichen Flora of Japan in Color [ M] . Osaka: Hoikusha Publishing Co.ꎬ Ltd.ꎬ 124—159 Zhao JD ( 赵继鼎) ꎬ Xu LW ( 徐连旺) ꎬ Sun ZM ( 孙曾美) ꎬ 1982. The Preliminary of Lichen in China ( 中 国 地 衣 初 编 ) [ M] . Beijing: Science Pressꎬ 41—43