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A study of the pruinose species of Hypogymnia (, ) from China

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A study of the pruinose species of Hypogymnia (Parmeliaceae, Ascomycota) from China

Xin-Li WEI and Jiang-Chun WEI

Abstract: Six pruinose species of Hypogymnia are reported in this paper, including one new species Hypogymnia pruinoidea. The of Hypogymnia pseudopruinosa was found to be a mixture with H. laccata. Hypogymnia pseudopruinosa is therefore typified with a lectotype, and the description of H. pseudopruinosa is revised. Distributions of the six pruinose species are given and discussed. Com- ments on differences and similarities between pruinose species of Hypogymnia are made. Diagnostic characters of each species, and a key to the pruinose species of Hypogymnia in China, are also provided. Key words: H. pruinoidea, H. pseudopruinosa, lichen substances, pruina Accepted for publication 6 June 2012

Introduction Materials and Methods Although over 100 species of Hypogymnia Specimens treated here are preserved in the Lichen (Nyl.) Nyl. have been reported worldwide, Section of Herbarium Mycologicum Academiae Sinicae (HMAS-L). A dissecting microscope (ZEISS Stemi SV according to the latest statistics from Index 11), compound microscope (OPTON III), and scanning Fungorum, only five species have pruina on electron microscope (SEM, FEI, Quanta 200) were used the upper surface of the lobes (Wei & Jiang to study morphology. Colour test reagents (10% aqueous 1980; Chen 1994; Wei & Wei 2005). So far, KOH and concentrated alcoholic p-phenylenediamine) and thin-layer chromatography (TLC) were used for the all of these species are endemic to China. detection of lichen substances (Culberson & Kristinsson Thus, during our studies on the 1970; Culberson 1972). of Chinese species of Hypogymnia, the prui- nose species group attracted our attention. Taxonomy A new pruinose species collected from Mt. Taibaishan, Shaanxi Province, is described Lichen thalli often have a whitish, flour-like in this paper. The type of Hypogymnia pseudo- surface covering called pruina. It is some- pruinosa X. L. Wei & J. C. Wei (Wei & Wei times treated as a diagnostic character to 2005) was found to be a mixture with H. delimit lichen species (Giralt et al. 2001), laccata J. C. Wei & Y. M. Jiang, necessitating although in some genera it is considered to the lectotypification of H. pseudopruinosa. be highly variable and unreliable. To date, six pruinose species of Hypogymnia, H. prui- noidea X. L. Wei & J. C. Wei, sp. nov, H. lijiangensis J. B. Chen, H. pruinosa J. C. Wei & Y. M. Jiang, H. pseudopruinosa X. L. Wei & J. C. Wei, H. subfarinacea X. L. Wei & X-L. Wei and J-C. Wei (corresponding author): State J. C. Wei, and H. subpruinosa J. B. Chen, Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China. all endemic to China, have been found and Email: [email protected] reported. 784 THE LICHENOLOGIST Vol. 44

Key to the pruinose species of Hypogymnia in China 1 Soredia present ...... H. subfarinacea Soredia absent ...... 2 2(1) Medulla PD+ orange-red, containing physodalic acid in medulla ...... 3 Medulla PD-- ...... 4 3(2) Lobes divergent, with black borders; holes mainly on axils, rarely on lower surface ...... H. lijiangensis Lobes crowded, without black borders; holes on lobe tips...... H. subpruinosa 4(2) Pruina layer dense, with a sharp demarcation between pruinose and epruinose regions; containing alectoronic acid in thallus ...... H. pruinosa Pruina layer thin; lacking alectoronic acid in thallus...... 5 5(4) Thallus texture cartilaginous to slightly papery; lobes subdichotomously branched; holes mainly present on lobe tips ...... H. pseudopruinosa Thallus texture cartilaginous; lobes isodichotomously branched; holes present on lobe tips, axils, and lower surfaces ...... H. pruinoidea

The Species Known distribution. Sichuan, Tibet and Yunnan provinces of China (Fig. 4). Hypogymnia lijiangensis J. B. Chen Selected specimens examined. China: Sichuan: Xiang- Acta Mycologica Sinica 13 (2): 109 (1994); type: China, cheng, Mt. Wumingshan, 1983, J. J. Su, H. A. Wen, Yunnan, Lijiang county, 1981, X. Y. Wang, X. Xiao, B. Li 6036 (HMAS-L). Tibet: Riwoqeˆ county, 1976, J. J. Su 6991 (HMAS—holotype). Y. C. Zong, Y. Z. Liao 256-3 (HMAS-L). Yunnan: Lijiang county, Yulong Snow Mountain, alt. 3060 m, (Figs 1A, 2A, 3A) 1981, X. Y. Wang, X. Xiao, J. J. Su 5124 (HMAS-L); This species is characterized by discrete Shangri-La county, Mt. Tianbaoshan, alt. 3700 m, 1981, X. Y. Wang, X. Xiao, J. J. Su 5189 (HMAS-L). lobes with black borders, pruina limited to lobe tips, and presence of physodalic acid in Hypogymnia pruinoidea X. L. Wei & the thallus. It is similar to H. subfarinacea J. C. Wei sp. nov. X. L. Wei & J. C. Wei and H. subpruinosa J. B. Chen in the location of pruina and MycoBank No.: MB 564892 chemistry, but clearly differs in morphology. Species characterized by isodichotomously branched lobes, Hypogymnia lijiangensis has separated lobes, pruinose lobe tips and upper surface, and abundant per- holes mainly in the axils, lacks soredia on forations present on lobe tips, axils, and lower surface. Type: China, Shaanxi, Mt. Taibaishan, alt. 2800 m, the upper surface, and has a cartilaginous on Abies trunk, 3 August 2005, X. L. Wei 1727 texture; H. subfarinacea has granular soredia (HMAS—holotype). on the upper surface, being easily separated from the other pruinose species of Hypogym- (Figs 1B, 2B, 3B) nia, cartilaginous texture but slightly papery; Thallus foliose, up to 6 cm wide, tex- and H. subpruinosa has crowded lobes with ture cartilaginous, loosely adnate, with flat, holes only on the lobe tips. crowded, mostly isodichotomously branched, Chemistry. Thallus cortex K+ yellow; me- hollow lobes 05–10 mm wide and 05–20 dulla K+ pale reddish brown, PD+ orange- mm long, apices blunt; upper surface greyish red. TLC: atranorin, physodic acid, physo- green, rugose, dull, thinly pruinose or pruina dalic acid, 3-hydroxyphysodic acid, protoce- limited to the lobe tips, with a sharp demar- traric acid. cation between the pruinose and epruinose 2012 Pruinose Hypogymnia species from China—Wei & Wei 785

Fig.1. Hypogymnia species. A, H. lijiangensis (holotype); B, H. pruinoidea (holotype); C, of H. pruinosa (holotype); D, H. pseudopruinosa (lectotype); E, H. subfarinacea (holotype); F, H. subpruinosa (holotype). Scale ¼ 5mmincrements. region; soredia and isidia lacking; lobules Apothecia not seen. present; lower surface black, pale brown at Pycnidia mostly at lobe tips, immersed to lobe tips, rugose, dull, perforations present slightly protruding, black, mostly punctiform, on many lobe tips, axils, and lower surfaces, aggregated; conidia fusiform, simple, 60– the holes not rimmed; medulla hollow, both 75 10 mm. the ceiling and floor of the cavity white to Chemistry. Reagent tests: cortex K+ yellow, pale dirty brown. P-- ; medulla K+ pale reddish brown, KC+ 786 THE LICHENOLOGIST Vol. 44

Fig. 2. Pruina of Hypogymnia species. A, H. lijiangensis (holotype); B, H. pruinoidea (holotype); C, H. pruinosa (holotype); D, H. pseudopruinosa (lectotype); E, H. subfarinacea (holotype); F, H. subpruinosa (holotype). Scale ¼ 5mm increments. pale pink, P-- . All specimens contain atra- province (Fig. 4). The high-elevation moun- norin, physodic and 3-hydroxyphysodic acids, tain habitats are cool and moist, supporting most specimens also contain vittatolic acid. lichen-rich forests, woodlands and subalpine areas. These areas are rich in Hypogymnia Etymology. The epithet ‘pruinoidea’ refers species such as H. arcuata Tchaban. & to the pruinose lobe tips and upper surface McCune, H. austerodes (Nyl.) Ra¨sa¨nen, H. in this species. flavida McCune & Obermayer, H. hypotrypa Habitat, distribution and substratum. Hypo- (Nyl.) Rass., and others. gymnia pruinoidea occurs on bark and wood, Remarks. Morphologically the new species usually of Abies in Mt. Taibaishan, Shaanxi resembles H. pseudopruinosa, but differs in 2012 Pruinose Hypogymnia species from China—Wei & Wei 787

Fig.3.Hypogymnia species, location of holes indicated by arrows. A, H. lijiangensis (holotype); B, H. pruinoidea (holotype); C, H. pruinosa (holotype); D, H. pseudopruinosa (lectotype); E, H. subfarinacea (holotype); F, H. subpruinosa (holotype). Scale ¼ 5 mm.

having a thallus with a more cartilaginous differs by perforations sometimes grouped texture, isodichotomously branched lobes, on the lower surface and containing physodic and abundant perforations present on lobe and 3-hydroxyphysodic acids. Hypogymnia tips, axils, and lower surface, whereas H. pruinosa is characterized by perforations main- pseudopruinosa is characterized by a thallus ly at lobe tips, and contains alectoronic acid. with a cartilaginous to slightly papery texture, Specimens examined. China: Shaanxi Province:Mt. subdichotomously branched lobes, and per- Taibaishan, 2800 m, on trunk of Abies tree, 2005, X. L. forations mainly present on lobe tips. The Wei 1728, 1729; 3136 m, on trunk of Abies tree, 2005, new species also resembles H. pruinosa, but X. L. Wei 1662, 1663, 1664 (HMAS-L). 788 THE LICHENOLOGIST Vol. 44

Hypogymnia pruinosa J. C. Wei & Apothecia rare, 1–3 mm diam., stipitate; Y. M. Jiang disc yellow-brown or red-brown, glossy, con- Acta Phytotax. Sin. 18 (3): 386 (1980); type: China, cave at first and then slightly plane with en- Tibet, Changdu, 1976, Y. C. Zong, Y. Z. Liao 215 tire and thin margin; epithecium brown, 7–9 (HMAS—holotype; HMAS—isotype). mm thick; hymenium colourless, 32–36 mm thick; asci clavate, 235–250 8–14 mm, 8- (Figs 1C, 2C, 3C) spored; spores simple, colourless, ellipsoid This species is characterized by almost cir- to nearly spherical, 55–70 35–40 mm; cular thalli, wide and short lobes, a layer of paraphyses linear, septate, 2 mm wide, slightly dense pruina on the upper surface, presence swollen at the tips; hypothecium colourless, of a sharp demarcation between pruinose 27–36 mm thick. and epruinose regions, and in containing Pycnidia common, brown to black; conidia alectoronic acid in the thallus. weakly bifusiform, c.25 (rarely to 5 mm) 10 mm. Chemistry.TLC:atranorin,alectoronicacid. Chemistry. Cortex K+ yellow, P-- ; medulla Distribution. Shaanxi, Sichuan, Tibet, and K+ yellow, P-- ; containing atranorin and Yunnan provinces of China (Fig. 4). physodic acid. Selected specimens examined. China: Shaanxi:Mei county, Mt. Taibaishan, 2005, X. L. Wei 1624 (HMAS- Etymology. The epithet ‘pseudopruinosa’ L). Sichuan: Kangding county, Minya Konka, alt. 3600 refers to the presence of pruina only on the m, 1982, X. Y. Wang, X. Xiao, B. Li 9438 (HMAS-L). Tibet: Linzhi county, alt. 3011m, 2004, X. L. Wei 1041 lobe tips. (HMAS-L). Yunnan: Lijiang county, Yulong Snow Distribution. Shaanxi and Yunnan provinces Mountain, alt. 3100 m, 1981, X. Y. Wang, X. Xiao, J. J. Su 4829 (HMAS-L). of China (Fig. 4).

Hypogymnia pseudopruinosa X. L. Wei Remarks. This species resembles H. macro- & J. C. Wei spora (Zhao 1964; Wei 1991) at first sight, but differs by having a dense layer of pruina Mycotaxon 94: 155 (2005); type: China, Yunnan, Deˆqeˆn county, 1981, X. L. Wei 7606 (HMAS—lectotype desig- limited to the tips of the lobes, holes not nated here). rimmed, and smaller ascospores. The holo- type of H. pseudopruinosa was found to be (Figs 1D, 2D, 3D) mixed with H. laccata. Some lobes of the Thallus foliose, tightly appressed, texture two species overlapped each other, which cartilaginous to slightly papery; lobes sub- led to some phenotypic characters of H. dichotomously branched, 1–2 mm wide, 5 laccata being wrongly assigned to H. pseudo- mm long; dense layer of pruina limited to pruinosa, whereas some diagnostic characters the lobe tips; upper surface greyish green to were actually based on both species. The dark brownish yellow, partly black, rugose; diagnostic characters mistakenly attributed isidia, soredia and lobules lacking; lower sur- to H. pseudopruinosa include the glossy upper face black, brown near the apices, rugose, surface and PD+ medulla (physodalic acid), and perforations mainly present on the lobe which were actually based on H. laccata. tips; medulla hollow, the ceiling of the cavity A lectotype is chosen here (ICBN Art. 9.9) white to pale brown, and floor of the cavity from the separated true H. pseudopruinosa white to dark brown. portion of the original holotype and isotype. Upper cortex prosoplectenchymatous, pale The H. laccata in the mixture has been sepa- yellow, 120–145 mm thick; algal layer 100– rated out as specimen no. 7606-1. When this 145 mm thick, consisting of green, subspher- species was published as new, it was based ical cells of 75–95 mm diam.; hyphae in on a single specimen from Yunnan Province medulla colourless, c.2mm diam.; lower cor- but, during our recent research, at least four tex prosoplectenchymatous, pale yellow, 10 additional specimens were collected from mm thick. Shaanxi province. 2012 Pruinose Hypogymnia species from China—Wei & Wei 789

Fig. 4. Distribution of the six pruinose species of Hypogymnia. H. lijiangensis (b), H. pruinoidea (n), H. pruinosa (e), H. pseudopruinosa (D), H. subfarinacea (C), H. subpruinosa (j).

Selected specimens examined. China: Shaanxi:Mei Distribution. Sichuan and Yunnan provinces county, Mt. Taibaishan, 1988, C. H. Ma 074 (HMAS-L); of China (Fig. 4). 2011, X. L. Wei w11139, w11143, w11150 (HMAS-L). Specimens examined. China: Yunnan: Lijiang county, Hypogymnia subfarinacea X. L. Wei & Yulong Snow Mountain, alt. 2900 m, 1981, X. Y. J. C. Wei Wang, X. Xiao, J. J. Su 4892 (HMAS-L); ibid., alt. 3100 m, 1981, X. Y. Wang, X. Xiao, J. J. Su 6591 Mycotaxon 94: 156 (2005); type: China, Sichuan, Nan- (HMAS-L). ping county, Jiuzhai Gou, 1983, X. Y. Wang & X. Xiao 10582 (HMAS—holotype). Hypogymnia subpruinosa J. B. Chen (Figs 1E, 2E, 3E) Acta Mycologica Sinica 13 (2): 107 (1994); type: China, This species superficially resembles H. Yunnan, Zhongdian county, 1981, X. Y. Wang, X. Xiao, J. J. Su 7094 (HMAS—holotype). farinacea Zopf, but differs in having more separated lobes, the pruina limited to the (Figs 1F, 2F, 3F) lobe tips, and containing physodalic acid; H. farinacea contains atranorin, chloroatranorin, This species is characterized by wide and physodic acid, 3-hydroxyphysodic acid, and short lobes with round tips, strongly rugose 20 -O-methylphysodic acid (McCune 2002). upper surface, pruina limited near to the lobe tips, and presence of physodalic acid in Chemistry. Cortex K+ yellow, P-- ; medulla the thallus. It is similar to H. pruinosa in ex- K+ yellow, P+ orange-yellow orange-red. ! ternal appearance, but the latter has pruina TLC: atranorin, physodic acid, physodalic all over the upper surface and contains alec- acid, 3-hydroxyphysodic acid, and protoce- toronic acid. traric acids. 790 THE LICHENOLOGIST Vol. 44

Table 1. Comparison of phenotypic characters of the six pruinose species of Hypogymnia from China

H. H. H. H. H. H. lijiangensis pruinoidea pruinosa pseudopruinosa subfarinacea subpruinosa

Texture Cartilaginous + + + + + + Papery ------+ + -- Lobes* Crowded -- -- + -- -- + Isodichotomous -- + ------Subdichotomous + -- + + + + Soredia Present ------+ -- Holes (on) Lobe tip -- + + + -- + Axil + + -- -- + -- Lower surface -- + -- -- + -- Medulla Ceiling colour† + + + + + + Floor colour 1 -- + ------Floor colour 2 + -- + + + + PD+ + ------++

* Isodichotomous: isodichotomously branched; subdichotomous: subdichotomously branched † Ceiling colour: white to pale brown in all species; floor colour 1: white to pale brown; floor colour 2: pale to dark brown

Chemistry. Cortex K+ yellow; medulla K+ sodalic acid, with the result of the medulla pale reddish brown, PD+ orange-red. TLC: being PD+ orange-red or PD-- , respectively, atranorin, physodic acid, physodalic acid, is sometimes a significant diagnostic character 3-hydroxyphysodic acid, and protocetraric to delimit species of Hypogymnia.Twoprui- acids. nose species H. lijiangensis and H. subpruinosa, have a PD+ medulla. Hypogymnia lijiangensis Distribution. Heilongjiang, Shaanxi, Si- has separate rather than contiguous lobes chuan, Tibet, and Yunnan provinces of China with black borders, and holes mainly in the (Fig. 4). axils, rarely on the lower surface; H. sub- Selected specimens examined. China: Heilongjiang: Dai- pruinosa has crowded lobes, without black ling, s. n. (HMAS-L). Shaanxi: Mt. Taibaishan, alt. borders, and holes only on the lobe tips. The 3280 m, 2005, X. L. Wei 1986 (HMAS-L). Sichuan: -- Maerkang county, Mt. Mengbishan, alt. 3700 m, X. Y. three species with a PD medulla are chemi- Wang & X. Xiao 11503 (HMAS-L). Tibet: Linzhi cally distinct: alectoronic acid is present in county, alt. 3010 m, X. L. Wei 1036 (HMAS-L). H. pruinosa, but absent in the other two Yunnan: Shangri-La county, Mt. Tianbaoshan, alt. species, H. pseudopruinosa and H.pruinoidea. 3800 m, 1981, J. J. Su, X. Xiao, X. Y. Wang 6808 Hypogymnia pseudopruinosa superficially re- (HMAS-L). sembles the epruinose species, H. macro- spora, but differs by having a dense layer of Discussion pruina limited to the tips of the lobes and smaller ascospores. Compared with H. prui- Of the six pruinose species of Hypogymnia, noidea, the thallus of H. pseudopruinosa has a only H. subfarinacea has granular soredia on cartilaginous to papery texture, subdichoto- the upper surface, easily separating it from mously branched lobes, and perforations the other five. Presence or absence of phy- mainly present on lobe tips, whereas H. prui- 2012 Pruinose Hypogymnia species from China—Wei & Wei 791

Fig. 5. SEM photographs of pruina on Hypogymnia species. A, whewellite in H. lijiangensis (holotype); B, weddellite of crystal type 3 (white arrow) and whewellite (black arrow) in H. pruinoidea (holotype); C, weddellite of crystal type 2 in H. pruinosa (holotype); D, whewellite in H. pseudopruinosa (lectotype); E, weddellite of crystal type 2 (black arrow) and whewellite (white arrow) in H. subfarinacea (holotype); F, weddellite of crystal type 3 (arrow on right) and whewellite (upper arrow) in H. subpruinosa (holotype). Scales: A–D & F ¼ 10 mm; E ¼ 5 mm. 792 THE LICHENOLOGIST Vol. 44 noidea has a thallus with a more cartilaginous based on SEM study. Types 1–3 belong to texture, isodichotomously branched lobes, the weddellite, and type 4 belongs to the and abundant perforations present on lobe whewellite (Wadsten & Moberg 1985). In tips, axils and lower surface. Table 1 com- our SEM study, three types of crystal struc- pares six pruinose species of Hypogymnia ture have been confirmed on the pruina of more clearly, including the main different the six pruinose Hypogymnia species: weddel- phenotypic characters, such as thallus tex- lite type 2, type 3, and whewellite. Hypogymnia ture, lobes, holes, soredia, and medulla PD pruinoidea has two crystal types, weddellite reactions. type 3 and whewellite (Fig. 5A); H. lijiangensis Among the six pruinose species of Hypo- has only one crystal type, whewellite (Fig. gymnia, all except H. pruinoidea occur in 5B); H. pruinosa has only one crystal type, south-western China, and two species, H. weddellite type 2 (Fig. 5C); H. pseudopruinosa pruinosa and H. subpruinosa,extendtonorth- has only one crystal type, whewellite (Fig. western and (or) north-eastern China. Hypo- 5D); H. subfarinacea has two crystal types, gymnia pruinoidea is found only in Shaanxi weddellite type 2 and whewellite (Fig. 5E); province, north-western China. Located on and H. subpruinosa has two crystal types, a low latitude plateau, south-western China weddellite type 3 and whewellite (Fig. 5F). It arises from south-east Tibet, crosses over is postulated that the formation of calcium west Sichuan, and expands to central and oxalate on the surface of is caused by north Yunnan. A combination of complex a need for the lichen to dispose of an excess of landforms with advantageous moisture con- calcium, and there seems to be some relation- ditions in south-western China results in an ship between different types of crystal struc- extremely abundant , with many ture and habitat, especially with regard to endemic species of animals and plants. Hy- different conditions of acidity and water con- pogymnia species are also abundant in south- tent (Wadsten & Moberg 1985). However, western China. Over 80% of known Hypo- the taxonomic significance of different crystal gymnia species in China can be found in this structures of calcium oxalate is not yet clear. area, and about 18% of the species are In conclusion, based on the above men- endemic. South-western China can therefore tioned phenotypic characters, presence or be regarded as a main centre of speciation for absence of soredia and black borders, loca- Hypogymnia. Mount Taibaishan is the high- tion of pruina and thallus perforations, and est peak of Qinling, Shaanxi province, which chemistry, all the six pruinose species can be is the watershed of Yangtze River and Yellow easily separated. All the phenotypic charac- River. As the highest mountain in the eastern ters used to define the six pruinose species half of China, it has a very complex geogra- of Hypogymnia are commonly used in the phy and climate. The marked difference in taxonomy of Hypogymnia, while their objec- elevation leads to a vertical distribution of tivity and the phylogenetic placement of climate, animals, and plants, and many new each species based on them will require species of plants endemic to Taibaishan molecular methods. have been reported. Over 30% of known Hy- This project was supported by the Main Direction Pro- pogymnia species in China can be found in gram of Knowledge Innovation of the Chinese Academy this area, including the new endemic species, of Sciences (KSCX2-EW-Z-9). We thank Dr Irwin M. H. pruinoidea. Brodo for his careful modification of English grammar The common character of the six species in the manuscript. Professor Bruce McCune of Oregon State University is thanked for the significant discussion of Hypogymnia treated in this paper is having about Hypogymnia with the first author, and he and one pruina. Pruina is a whitish, flour-like surface anonymous reviewer are also thanked for their valuable covering, consisting primarily of superficial suggestions on the manuscript. Professor Jianbin Chen deposits, most commonly calcium oxalate gave constructive comments on an earlier version of our paper. Mr Chunli Li helped take SEM photograghs of (Bu¨del & Scheidegger 1996). There are two pruina. The authors are also grateful to Ms. Hong Deng kinds of calcium oxalate found in lichen for giving considerable assistance during the studies in pruina, weddellite and whewellite, and four HMAS-L. types of crystal structure have been reported 2012 Pruinose Hypogymnia species from China—Wei & Wei 793

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