Psora altotibetica (Psoraceae, Lecanorales), a new lichen species from the Tibetan part of the Himalayas

Einar Timdal; Walter Obermayer; Mika Bendiksby

doi:10.3897/mycokeys.13.8824

Abstract

In the present study, we describe the new species, Psora altotibetica, from nine localities in China (Tibetan area) and Nepal. The study includes analyses of anatomy, secondary chemistry, and DNA sequence data of P. altotibetica and presumed close relatives. Psora altotibetica resembles P. indigirkae morphologically, but is phylogenetically closer to P. tenuifolia and P. vallesiaca. It differs from P. indigirkae in the colour of the apothecia, the size of the ascospores, and in the secondary chemistry. The species is terricolous and was collected in the alpine zone of the Great Himalayas between 4230 and 5000 m altitude. Psora tenuifolia and P. vallesiaca are here reported as new to China and the Himalayas.

Key words

Asia, Lecanorales , lichenized ascomycetes, taxonomy

Introduction

The genus Psora consists of c. 30 species growing on soil and rock, mainly in arid areas, from the arctic to the subtropical regions of the world (Timdal 2002). The current concept of the genus was proposed by Schneider (1980) and emended by Timdal (1984). No world monograph exists, but there are two revisions of the genus in North America (Timdal 1986, 2002) and keys to the species in Europe (Poelt and Vězda 1981) and Asia (Timdal and Zhurbenko 2004). Ekman and Blaalid (2011) published a molecular phylogeny of the Psoraceae, including 18 species of Psora.

In the current Chinese checklist (Wei 1991), five species of Psora are listed: (1) P. asahinae (Zahlbr.) J.C.Wei, (2) P. asiae-centralis (H.Magn.) N.S.Golubk., (3) P. crenata (Taylor) Reinke, (4) P. decipiens (Hedw.) Hoffm., and (5) P. lurida (Ach.) DC. However, three species on this list (1, 2, and 5) are currently excluded from Psora, and for one (3), the single Chinese record is based on incorrectly identified material [Psora asahinae was synonymised with Psorula rufonigra (Tuck.) Gotth.Schneider (Schneider 1980), P. asiae-centralis was placed in Toninia as T. tristis ssp. asiae-centralis (H.Magn.) Timdal by Timdal (1992), P. lurida was placed in Romjularia as R. lurida (Ach.) Timdal by Timdal (2008), and the report of P. crenata is based on the holotype of Lecidea undulata H.Magn. (Bohlin 82, S!) which was placed in synonymy with Anamylopsora pulcherrima (Vain.) Timdal by Timdal (1991)]. Hence, P. decipiens (4) is the only currently known species of Psora in China. It is listed from Xizang by Wei (1991) and from Yunnan by Wang (2012, including a beautiful photograph), and is, together with Psora himalayana (C.Bab.) Timdal (described from Uttar Pradesh, India, at 4700 ft), the only known Psora species in the Himalayas.

On a visit to the herbarium of Institut für Pflanzenwissenschaften, Karl-Franzens-Universität Graz (GZU) in 1992, one of us (ET) discovered an apparently undescribed species of Psora collected near the Khumbu Glacier south of Mt Everest (Nepal) by Josef Poelt in 1962. Two years later, ET was asked by Torstein Engelskjøn to identify some specimens collected by him in 1993 in the Rongbuk Valley north of Mt Everest (Tibet) during a joint Chinese-Norwegian scientific expedition. The material comprised the same apparently undescribed species of Psora. The material from both expeditions was rather poor and put aside pending richer collections. In 2013, during a visit to GZU, MB was made aware of the Psora collection that WO had made in the Himalayas in 1994 and 2000. This collection contained the putative new species, and a renewed study, including DNA analysis, has led us to describe it here as Psora altotibetica.

Material and methods

The specimens

This study is based on 15 collections of Psora made by WO in the Himalayas in 1994 and 2000, the collection of Psora altotibetica by J. Poelt (GZU), the two by T. Engelskjøn (TROM), two by G. & S. Miehe made in the Himalayas in 1993 (GZU), four collections from the Karakoram Range (Pakistan) in GZU which are here identified as P. himalayana and P. vallesiaca (Schaer.) Timdal, and two collections of P. indigirkae Timdal & Zhurb. from Yakutia in O. Voucher data for these 26 specimens are given in the list of examined specimens and in Table 1. The table also contains voucher data for the specimens of which DNA was downloaded from GenBank or obtained through the Norwegian Barcode of Life project (NorBOL). With the exception of one collection (P. himalayana, Zhurbenko 98161), we have examined all collections listed in Table 1 by morphology during this project or previously.

Table 1.

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Psora specimens used in this study with voucher information, major lichen substances, and GenBank accession numbers. New sequences are indicated by accession numbers in bold.

Taxon, Specimen	Voucher Information	Major Lichen Substances	GenBank Accession Number
Taxon, Specimen	Voucher Information	Major Lichen Substances	ITS	mtSSU
P. altotibetica 1	China, Xizang, Obermayer 5282 (GZU), holotype	gyrophoric acid	KU863638	KU863651
P. altotibetica 2	China, Xizang, Miehe & Miehe 9573/23/02 (GZU)	gyrophoric acid	KU863639	KU863652
P. altotibetica 3	China, Xizang, Obermayer 5223 (GZU)	gyrophoric acid	KU863640	KU863653
P. altotibetica 4	China, Xizang, Obermayer 4365 (GZU)	gyrophoric acid	KU863642	KU863655
P. altotibetica 5	China, Xizang, Obermayer 3967 (GZU)	gyrophoric acid	KU863641	KU863654
P. altotibetica 6	China, Xizang, Obermayer 4485 (GZU)	gyrophoric acid	KU863643	KU863656
P. altotibetica	China, Xizang, Engelskjøn T-030b (TROM L-42812)	gyrophoric acid	-	-
P. altotibetica	China, Xizang, Engelskjøn T-036 (TROM L-42813)	gyrophoric acid	-	-
P. altotibetica	China, Xizang, Miehe & Miehe 9573/23/03 (GZU)	-	-	-
P. altotibetica	China, Xizang, Obermayer 4350 (GZU)	gyrophoric acid	-	-
P. altotibetica	China, Xizang, Obermayer 4502 (GZU)	-	-	-
P. altotibetica	China, Xizang, Obermayer 4981 (GZU)	gyrophoric acid	-	-
P. altotibetica	Nepal, Poelt 1138 (GZU)	gyrophoric acid	-	-
P. californica	USA, California, Timdal SON139/04 (O-L-60112)	bourgeanic acid, gyrophoric acid	EF524322	EF524292
P. globifera 1	Greenland, Timdal 10149 (O-L-139171)	no substances	EF524323	EF524294
P. globifera 2	Norway, Klepsland JK11-L619 (O-L-183774)	no substances	KU873928	-
P. globifera 3	Norway, Bendiksby et al. 12914 (O-L-184327)	no substances	KU873930	-
P. globifera 4	Norway, Klepsland JK11-L213 (O-L-177145)	no substances	KU873929	-
P. globifera 5	Norway, Hjelmstad s.n. (O-L-184143)	no substances	KU873932	-
P. himalayana	Russia, Yakutia, Zhurbenko 98161 (M-0066792)	-	AY425635	-
P. himalayana	Pakistan, Miehe & Miehe 3529 (GZU)	no substances	-	-
P. himalayana	Pakistan, Poelt K91-416	no substances	-	-
P. hyporubescens	USA, California, Bratt & Timdal 7052 (O-L-22483), holotype	anthraquinones, gyrophoric acid	EF524311	EF524295
P. indigirkae 1	Russia, Yakutia, Haugan & Timdal YAK19/03 (O-L-19148), holotype	bourgeanic acid, gyrophoric acid	EF524302	-
P. indigirkae 2	Russia, Yakutia, Haugan & Timdal YAK17/24 (O-L-19086), paratype	bourgeanic acid, gyrophoric acid	KU863631	KU863644
P. indigirkae 3	Russia, Yakutia, Zhurbenko 92185 (O-L-118686), paratype	bourgeanic acid, gyrophoric acid	KU863632	KU863645
P. nitida	Mexico, Baja California, Timdal SON33/06 (O-L-15546)	gyrophoric acid	EF524313	EF524296
P. pacifica	USA, California, Rosentreter 14580 (O-L-126265)	gyrophoric acid, unknown accessory	EF524314	EF524297
P. peninsularis	Mexico, Baja California, Timdal SON32/07 (O-L-15539), holotype	norstictic acid	EF524320	EF524298
P. russellii	Mexico, Baja California, Timdal SON31/03 (O-L-15531)	norstictic acid	EF524321	EF524300
P. tenuifolia 1	Russia, Yakutia, Haugan & Timdal YAK17/26 (O-L-19088)	norstictic acid, zeorin	EF524309	EF524303
P. tenuifolia 2	China, Xizang, Obermayer 4487 (GZU)	norstictic acid, zeorin	KU863636	KU863649
P. tenuifolia 3	China, Xizang, Obermayer 5236 (GZU)	zeorin	KU863637	KU863650
P. tenuifolia	China, Sichuan, Obermayer 9791 (GZU)	norstictic acid, zeorin	-	-
P. tenuifolia	China, Xizang, Obermayer 4525 (GZU)	zeorin	-	-
P. testacea	Greece, Rui & Timdal TH06/04 (O-L-59263)	atranorin	EF524315	EF524301
P. tuckermanii	USA, Arizona, Rui & Timdal US240/05 (O-L-59926)	no substances	EF524317	EF524304
P. vallesiaca 1	Greece, Rui & Timdal 7993 (O-L-15186)	norstictic acid	EF524324	EF524291
P. vallesiaca 2	China, Xizang, Obermayer 3227 (GZU)	norstictic acid	KU863633	KU863646
P. vallesiaca 3	China, Xizang, Obermayer 5279 (GZU)	no substances	KU863635	KU863648
P. vallesiaca 4	Pakistan, Poelt K91-705 (GZU)	norstictic acid	KU863634	KU863647
P. vallesiaca 5	Norway, Bendiksby et al. 12979 (O-L-184392)	norstictic acid	KU873926	-
P. vallesiaca 6	Norway, Klepsland JK11-L624 (O-L-183778)	norstictic acid	KU873927	-
P. vallesiaca 7	Norway, Klepsland JK11-L601 (O-L-183760)	norstictic acid	KU873931	-
P. vallesiaca	China, Xizang, Obermayer 4482 (GZU)	norstictic acid	-	-
P. vallesiaca	Pakistan, Poelt s.n. (GZU)	norstictic acid	-	-

Anatomy

Microscope sections were cut on a freezing microtome and mounted in water, 10% KOH (K), lactophenol cotton blue, a modified Lugol’s solution in which water was replaced by 50% lactic acid, and chlor-zinc-iodine. Amyloid reactions were observed in the modified Lugol’s solution after pretreatment in K. Chlor-zinc-iodine was used to locate remnants of algae in the cortex, and polarized light was used to locate crystals of secondary metabolites and calcium oxalate. Calcium oxalate was identified by adding 25% sulphuric acid to the section; the oxalate crystals dissolve and needle shaped crystals of calcium sulphate precipitate. Ascospore measurements are given as X ± 1.5×SD, where X is the arithmetic mean and SD – the standard deviation.

Secondary chemistry

Thin-layer chromatography (TLC) was performed in accordance with the methods of Culberson (1972), modified by Menlove (1974) & Culberson and Johnson (1982). All specimens were examined by TLC, with the exception of Miehe & Miehe 9573/23/03 and Obermayer 4502, which were omitted due to being represented by richer material from the same localities.

DNA extraction, PCR and sequencing

We performed DNA extraction, PCR amplification, PCR purification, and cycle sequencing as described by Bendiksby and Timdal (2013). DNA was extracted from apothecia of 13 specimens (Table 1; GenBank Accession Numbers KU863631–KU863656). All DNA isolates produced for the present study are deposited in the DNA collection at Natural History Museum, University of Oslo. We amplified and sequenced the nuclear ribosomal internal transcribed spacer (ITS) and the mitochondrial ribosomal small subunit (mtSSU) using the primer pairs ITS5/ITS4 (White et al. 1990) and mtSSU1/mtSSU3R (Zoller et al. 1999), respectively.

Data analyses

Sequences were assembled and edited using SEQUENCHER v.4.1.4 (Gene Codes Corporation, Ann Arbor, Michigan, U.S.A.). Alignments were established in BIOEDIT 7.2.3 (Hall 1999) using the “ClustalW/Multiple alignment” option with subsequent manual adjustments. We analysed and summarized the data with parsimony and Bayesian phylogenetic methods, including model testing, as described in Bendiksby et al. (2015). The nuclear and mitochondrial datasets were analysed separately and in combination (concatenated) with indels treated as missing data.

Results

Species identifications

The 24 Central Asian specimens were identified by morphology and secondary chemistry as Psora altotibetica (13), P. himalayana (2), P. vallesiaca (5), and P. tenuifolia Timdal (4).

Anatomy

The following key characters for including P. altotibetica in Psora were observed in the new species: the upper cortex contained remnants of algae throughout both the lower stainable layer and the upper epinecral layer (‘Scheinrindentyp’ of Poelt 1958); the hypothecium contained calcium oxalate crystals; the epihymenium contained orange crystals which dissolved in K with a purple diffusion (assumed to be anthraquinones); and the ascus contained a well-developed, amyloid tholus with a central, deeper amyloid tube structure (Porpidia-type).

The following species level characters were observed in P. altotibetica: Upper cortex composed of thin-walled hyphae with rounded lumina; lower cortex composed of mainly periclinally oriented hyphae; crystals of calcium oxalate and assumedly gyrophoric acid (dissolving in K) present both in upper cortex and medulla; no crystals in lower cortex; ascospores 9–14 × 5–7 µm.

Secondary Chemistry

The results of the TLC examinations are given in Table 1. All examined specimens of P. altotibetica contained gyrophoric acid; no traces of fatty acids were detected.

Molecular data

Altogether 26 DNA sequences were generated from 13 specimens for the present study (13 ITS and 13 mtSSU), including two specimens of P. indigirkae from Yakutia in O. In addition, seven unpublished ITS sequences of P. globifera (Ach.) A.Massal. and P. vallesiaca from Norway were generated by the lichen DNA barcode project, OLICH, at the Norwegian Barcode of Life (NorBOL). Moreover, 24 ITS and mtSSU sequences from 13 Psora specimens were downloaded from GenBank. GenBank accession numbers of all 57 sequences are given in Table 1.

Alignments and phylogenetic analyses

The ITS matrix of 32 accessions was 676 basepairs long and contained 181 parsimony-informative characters. The basepairs and parsimony-informative characters for the mtSSU matrix of 25 accessions were 881 and 28, respectively. The estimated best fit model of evolution for ITS was SYM+G and for mtSSU it was HKY+I+G. Both parsimony jackknife and Bayesian trees of ITS vs mtSSU were congruent but resolved to various extents (not shown). Therefore, for the final analyses, a concatenated dataset of 1557 bp was used. In the Bayesian analysis of the concatenated dataset, the average standard deviation of split frequencies had fallen to 0.0045 at termination (four million generations) and the first 1000 saved trees (i.e. 25%) were discarded as burn-in. The Bayesian 50% majority rule consensus tree, rooted with P. testacea, is presented with both Bayesian and parsimony branch support superimposed (Fig. 1). The molecular data group, with high support, multiple accessions of each species according to species determination based on morphology. The single exception is one accession of P. himalayana, which falls out nested within a P. vallesiaca clade. The latter consists of strongly supported subclades. Psora tenuifolia is strongly supported as phylogenetic sister species to P. altotibetica. The P. tenuifolia – P. altotibetica clade is in turn sister to the P. vallesiaca clade. A clade consisting of P. hyporubescens Timdal and P. pacifica Timdal is also strongly supported. Apart from this, the molecular data does not support any further inter-species relationships.

Figure 1.

The Bayesian 50% majority rule consensus tree based on a concatenated alignment of ITS and mtSSU sequences of 33 accessions of 14 Psora species (see Table 1). Parsimony jackknife support values above 50% are shown below branches and Bayesian posterior probabilities above. The curly branch leading to P. testacea has been shortened to reduce the size of a broad figure.

Discussion

Psora tenuifolia is the sister species of P. altotibetica in our phylogeny (Fig. 1). It differs in having thinner, ascending, less pruinose, more white-edged squamules containing zeorin and usually norstictic acid, and in having a well-developed lower cortex composed of mainly anticlinally oriented hyphae which are densely covered by calcium oxalate crystals (Timdal 1986). Psora tenuifolia was previously known from Alaska and arctic Canada (Timdal 1986) and from Yakutia (Zhurbenko 2003). In two specimens (Obermayer 4525 and 5236) norstictic acid was not detected by TLC; these specimens represent a previously unknown chemotype of P. tenuifolia. The species is here reported as new to China (Sichuan and Xizang) and the Himalayas. One collection (Obermayer 4487) is a mixture of P. altotibetica and P. tenuifolia.

Psora vallesiaca is the phylogenetic sister species of the P. altotibetica - P. tenuifolia clade (Fig. 1). It differs from P. altotibetica in having less pruinose squamules with a more up-turned and white-edged margin, and in containing norstictic acid. It is morphologically more similar to P. tenuifolia than to P. altotibetica; see Timdal (1986) for discussion on the differences between P. tenuifolia and P. vallesiaca. Two specimens here identified as P. vallesiaca (Obermayer 4482 and 5279) do not contain lichen substances and were first thought to represent P. himalayana. We obtained sequences from the latter, and it clusters with P. vallesiaca in our phylogeny (Fig. 1, specimen P. vallesiaca 3). The sequence of P. himalayana downloaded from GenBank (specimen not examined by us, chemistry unknown) also clusters with those of P. vallesiaca in our phylogeny. Timdal (1986) remarked that P. himalayana and P. vallesiaca are sometimes difficult to distinguish morphologically, but that the chemistry is diagnostic. It now seems that there is a norstictic acid deficient chemotype of P. vallesiaca, making the taxonomic status of P. himalayana in need of revision. Psora vallesiaca is here reported as new to China (Sichuan and Xizang) and the Himalayas.

The other species of Psora known from the Himalayas, P. decipiens, differs in having orange to red or rose, more regularly rounded squamules with a usually more upturned and crenulate margin, in having strictly marginal apothecia, and in lacking lichen substances or more rarely containing norstictic acid or very rarely hyposalazinic and hypostictic acids (see, e.g., Timdal 2002).

Nine other Psora species contain gyrophoric acid (Timdal 1986, 2002, Timdal and Zhurbenko 2004). Five of those are in our phylogeny (Fig. 1; i.e. P. californica Timdal, P. hyporubescens, P. indigirkae, P. nitida Timdal, and P. pacifica), but none are closely related to P. altotibetica. Psora indigirkae is the morphologically most similar Psora species, but differs from P. altotibetica in having brown apothecia, often with a reddish hue, larger ascospores (14–17 × 7–8 µm; n=50), and in its secondary chemistry: gyrophoric acid is accompanied by bourgeanic acid (Timdal and Zhurbenko 2004). None of the four remaining gyrophoric acid containing Psora species are morphologically similar to P. altotibetica. Psora nipponica (Zahlbr.) Gotth.Schneider and P. rubiformis (Ach.) Hook. have, e.g., a lower cortex similar to that of P. tenuifolia (Timdal 1986), P. montana Timdal has medium to castaneous brown squamules and brown, more plane apothecia, and P. russellii (Tuck.) A.Schneider has larger, more rounded squamules, often with a central depression, and almost always contains norstictic acid as the major compound (gyrophoric acid being minor to trace or lacking).

Taxonomy

Psora altotibetica Timdal, Obermayer & Bendiksby, sp. nov.

MycoBank No: Mycobank: MB 816840

Fig. 2

Diagnosis

Similar to Psora indigirkae, but apothecia black, ascospores shorter, and bourgeanic acid absent from the thallus.

Figure 2.

Psora altotibetica, part of holotype. Scale bar = 1 mm.

Type

CHINA. Xizang: Himalaya Range, 165 km SSE of Lhasa, 40 km W of Lhünze, little village on way to Nera Tso (=Ni La Hu), 28°23'N, 92°05'E, 4300–4400 m alt., dry-valley, N-exposed dry slopes, on the ground, 1 Aug 1994, W.Obermayer 5282 (holotype: GZU!).

Description

Thallus squamulose; squamules up to 3 mm wide, rounded, adnate, dispersed to adjacent, weakly concave to plane; upper surface medium brown, dull, becoming moderately to densely pruinose, smooth when young, later with fissures in the cortex; margin concolorous with upper side or partly white, straight to slightly down-turned or slightly up-turned, entire; upper cortex 40–60 μm thick, composed of pale brown, thin-walled hyphae with rounded lumina, containing remnants of algae throughout (Chlor-zinc-iodine!), containing crystals of gyrophoric acid (assumedly, dissolving in K) and (in pruinose squamules) crystals of calcium oxalate; epinecral layer hardly developed. Medulla not amyloid, containing both lichen substances (dissolving in K) and calcium oxalate; lower cortex poorly developed, composed of mainly periclinally oriented, pale brown hyphae, not containing crystals; lower surface brown. Apothecia up to 1.2 mm diam., marginal or submarginal on the squamules, plane and indistinctly marginate when young, soon becoming convex and immarginate, black, epruinose or faintly white pruinose at the margin. Hypothecium colourless, containing crystals of calcium oxalate; epihymenium yellowish brown, containing orange crystals dissolving in K, K+ purple. Ascus clavate, with a well-developed, amyloid tholus containing a deeper amyloid tube, lacking an ocular chamber (Porpidia-type); ascospores ellipsoid, simple, hyaline, 9–14 × 5–7 µm (n = 30). Conidiomata not seen.

Chemistry

Gyrophoric acid (by TLC); upper cortex and medulla K–, C+ faintly red, KC+ faintly red, P–.

Habitat and distribution

The species is terricolous and known from nine localities in China (Tibet) and Nepal at altitudes between 4230 and 5000 m (Fig. 3).

Figure 3.

Psora altotibetica, known distribution. Open circle = holotype locality.

Etymology

The name refers to its occurrence at high altitude in the Tibetan part of the Himalayas.

Other specimens examined

Psora altotibetica. CHINA. Xizang: Shegar, Rongbuk Valley, S of Rongpu-si, 5000 m alt., moraine hill, 7 Jul 1993, T.Engelskjøn T-030b (TROM L-42812); 5000 m alt., rock base, patchwise on silt, 7 Jul 1993, T.Engelskjøn T-036 (TROM L-42813); Upper Tsangpo basin, N of Saga, 29°34'N, 85°15'E, alt. 4760 m, 28°W, grazed stony relicts of Juniperus dwarf-scrub, on ground, 26 Aug 1993, G.Miehe. & S.Miehe 9573/23/02 (GZU) & 9573/23/03 (GZU); 120 km SSW of Quamdo (=Changtu), 10 km S of Bamda, 30°09'N, 97°17'E, 4500–4700 m alt., on mosses, overhang, NNW-exp., 6 Aug 1994, W.Obermayer 3967 (GZU); Himalaya Range, 160 km S of Lhasa, dry valley of Kuru river, 10 km NW Lhozag, 28°24'N, 90°39'E, 4230 m alt., N-exposed steep rocks in a glen, on soil, 17 Jul 1994, W.Obermayer 4350 (GZU) & 4365 (GZU); Himalaya Range, 170 km S of Lhasa, between Lhozhag and Lhakhang Dzong, W-facing slopes of Dhalari mountain, 28°20'N, 90°58'E, 4300 m alt., NNW-exposed, ±underhang, 20 Jul 1994, W.Obermayer 4485 (GZU) & 4502 (GZU); Himalaya Range, 170 km SE of Lhasa, 110 km SSE of Tsetang (Nedong), 28°35'N, 92°23'E, 4700 m alt., alpine meadows with Kobresia pygmaea, ground with Ochotona-burrows, 26 Jul 1994, W.Obermayer 4981 (GZU); Himalaya Range, 130 km SE of Lhasa, 50 km SSE of Tsetang (Nedong), on way to the pass Putrang La, 28°52'N, 92°06'E, 4400 m alt., dry slope, on soil, 2 Aug 1994, W.Obermayer 5223 (GZU). NEPAL. Mahalangur Himal, Khumbu, Moränen des Khumbu-Gletschers bei Lobuche, 4950–5000 m alt., Sep 1962, J.Poelt 1138 (GZU).

Psora himalayana. PAKISTAN. Karakorum, Naz Bar (Yasin), 36°17–25'N, 73°0–17'E, 3400–3470 m alt., subalpine Juniperus communis dwarf-scrub with Juniperus macropoda trees in Seriphidium maritimum steppe; on silt in rock crevices, 11 Sep 1990, G.Miehe & S.Miehe 3529 (GZU); Karakorum, Baltistan, Haramosh Range, “Alm” Pakora SE Ganto La, 35°41'N, 75°21'E, 3600–3800 m alt., pasture and rocks around the alm, rocky slopes, 3 Jul 1991, J.Poelt K91-416 (GZU).

Psora indigirkae. RUSSIA. Sakha Republic: Momskii region, along the river Indigirka, c. 48 km NNW of Tyubelyakh, 65°48'N, 142°53'E, 200–300 m alt., on calcareous soil in limestone cliffs, 20 Jul 1992, R.Haugan & E.Timdal YAK17/24 (O L-19086); c. 54 km N-NNW of Tyubelyakh, 65°51'N, 143°01'E, 200–300 m alt., 20 Jul 1992, M.P.Zhurbenko 92185 (O L-118686).

Psora tenuifolia. CHINA. Sichuan: Tibetan fringe mountains (=Hengduan Shan), Shaluli Shan, on the outskirts of Yajiang, 200 m E of the river Yalong Jiang, 30°02’22”N, 101°00’16”E, 2610 m alt., NE-exposed dry slopes with schist outcrops, on thin soil crust (over schist), 12 Aug 2000, W.Obermayer 9791 (GZU); Xizang: Himalaya Range, 170 km S of Lhasa, between Lhozhag and Lhakhang Dzong, W-facing slopes of Dhalari mountain, 28°20'N, 90°58'E, 4300 m alt., NNW-exposed, ±underhang, 20 Jul 1994, W.Obermayer 4487 (GZU); Himalaya Range, 175-180 km S of Lhasa, between Lhozhag and Lhakhang Dzong, Kuru river valley, pass, 28°12'N, 91°00'E, 3600 m alt., on soil, 21 Jul 1994, W.Obermayer 4525 (GZU); Himalaya Range, 210 km SE of Lhasa, 15 km ESE of Lhünze, way to Qayü, dry-valley of Subansiri, 28°24'N, 92°37'E, 4100–4200 m alt., on soil (+ mosses), 31 Jul 1994, W.Obermayer 5236 (GZU).

Psora vallesiaca. CHINA. Sichuan: Shalui Shan Mts, 30 km NE Batang, S Yidun, 30°16'N, 99°25'E, 3750–3800 m alt., on marble outcrops, soil, 25 Jun 1994, W.Obermayer 3227 (GZU); Xizang: Himalaya Range, 165 km SSE of Lhasa, 40 km W of Lhünze, little village on way to Nera Tso (=Ni La Hu), 28°23'N, 92°95'E, 4300–4400 m alt., dry-valley, N-exposed dry slopes, on the ground, 1 Aug 1994, W.Obermayer 5279 (GZU); Himalaya Range, 170 km S of Lhasa, between Lhozhag and Lhakhang Dzong, W-facing slopes of Dhalari mountain, 28°20'N, 90°58'E, 4300 m alt., NNW-exposed, ±underhang, 20 Jul 1994, W.Obermayer 4482 (GZU); PAKISTAN. Karakorum, Baltistan, Haramosh Range, between “Alm” Matumdus (3620 m, 35°42'N, 75°23'E) and Hemasil in the Basna Valley, 3100–3200 m alt., 7 Jul 1991, J.Poelt K91-705 (GZU); Karakorum, Baltistan, Basna valley, Basnald, 2500 m alt., 10 Jul 1991, J.Poelt s.n. (GZU).

Acknowledgments

We are grateful to Torstein Engelskjøn, Tromsø, and the late Josef Poelt for the loan of material. The two expeditions of Walter Obermayer to the Tibetan area were financially supported by the Austrian Science Fund in 1994 (project number P09663-BIO) and in 2000 (project number P13676-BIO). The seven Norwegian ITS sequences were provided by the Norwegian Barcode of Life project, funded by the Norwegian Taxonomy Initiative (Norske Artsprosjektet) administered by the Norwegian Biodiversity Information Centre (ArtsDatabanken) (project number 70184216).

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