Rare and noteworthy boreal lichens from the Altai Mountains - BioOne

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Dec 1, 1997 - groups that are relatively poorly investigated in. Siberia: ''biatoroid'' lichens, species from the. Lecanora varia group and species that commonly.
Rare and noteworthy boreal lichens from the Altai Mountains (South Siberia, Russia) Evgeny A. Davydov1,3 and Christian Printzen2 1

Altai State University, Lenin Ave. 61, and Tigirek State Reserve, Nikitina Str. 111, Barnaul, 656049, Russia; 2 Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany

ABSTRACT. Based on field trips between 1995–2007, rare and noteworthy lichens from the Altai Mountains in southern Siberia are described. Bacidia zerovii Oxner is recognized as a synonym of Scoliciosporum chlorococcum (Graewe ex Stenh.) Veˇzda. Lecidea brenneri H. Magn. is lectotypified and synonymized with Lecania cyrtellina (Nyl.) Sandst. The new combination Puttea exsequens (Nyl.) Printzen & Davydov is introduced for Lecidea exsequens Nyl. and is the oldest validly published name for Lecidea gibberosa sensu Th. Fr. non Ach. Of the 33 species reported here, five species – Lecania croatica, Lecanora pseudosarcopidoides, Lecidea, L. sanguineoatra, Puttea exsequens and Rimularia fuscosora are new for Asia and Russia, Buellia arborea is new to Asia, Lecanora farinaria is new to Russia, Caloplaca herbidella, C. sorocarpa, Fellhanera subtilis, Lecidea pullata, and Pertusaria coccodes are new to Siberia. Several species are reported as new to West or South Siberia. Most corticolous lichen species display a more or less circumboreal distribution and also occur in Europe and North America. KEYWORDS. Bacidia zerovii, Lecidea brenneri, Lecania, Rimularia, Puttea exsequens, Asia.

¤ Most forest lichen species of the Northern Hemisphere display a more or less circumboreal distribution with frequent intercontinental and sometimes bipolar disjunctions. Major problems for the interpretation of biogeographical patterns of such species, especially those with intercontinental disjunctions, are a lack of fossil records and, above all, incomplete knowledge of distributional ranges. South Siberia includes extensive mountain systems, which acted as glacial refugia for many boreal and nemoral species in North Asia. The boreal zone can be divided into several major geographical units: 3

Corresponding author’s e-mail: [email protected] DOI: 10.1639/0007-2745.115.1.61

¤

¤ Europe, Caucasus, Siberia, Far East, Western and Eastern North America. As a very crude approximation, the distribution area of boreal lichen species in the Northern Hemispere may be described as presence or absence in these regions. But for many species, reliable data does not even exist on this scale. In this study we present new information on the distribution of remarkable boreal lichens in the Altai Mountains (South Siberia). Our main focus was on groups that are relatively poorly investigated in Siberia: ‘‘biatoroid’’ lichens, species from the Lecanora varia group and species that commonly appear without fruit bodies and therefore require secondary metabolite chemistry for identification. The Altai is an extensive mountain region situated in central Eurasia and constitutes the highest

The Bryologist 115(1), pp. 61–73 Copyright E2012 by The American Bryological and Lichenological Society, Inc.

0007-2745/12/$1.45/0

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mountain range in Siberia. The highest summits rise above the snow line (2700–2950 m), with the tallest mountain being Mt. Belukha, at 4506 m. These mountains give rise to several creeks and rivers, most of which eventually join the Ob’ River. The regional climate is severely continental, and the continentality increases from the West, where the mean annual temperature is more than +2uC to the East where it is less than 24 uC. Because of the influence of the Asiatic anticyclone, the winter is long (5–9 months) and very cold. January temperatures range from 210 to 215uC in the foothills to 225 to 230uC in the sheltered hollows of the east. Summer at the elevation of 1000 m a. s. l. is short and cool. July temperatures range from 18–20uC in North-West to 14–16uC in the South-East. The transitions between summer and winter are sharp, particularly in the alpine belt. The western and north-eastern portions of the Altai experience a humid climate, with more than 1500 mm of precipitation more or less equally distributed throughout the year. By contrast, the south-eastern Altai has a climate characterized by low annual precipitation (100–200 mm) and snowless winters. The Altai is situated largely within the steppe zone. The altitudinal vegetation belts are represented by: shrub-steppe, forest-steppe, different types of taiga forest, Betula forests, subalpine meadows with shrubs and herbs, alpine meadows, and mountain tundra. Taiga forests are mainly composed of Abies sibirica Ledeb., Pinus sibirica Du Tour, and Larix sibirica Ledeb. The timberline in the Altai mountains ranges from 1700–1800 m in the north to 2200–2300 m in the south (Kuminova 1960; Kamelin 1998). The lichen flora of the Altai Mts. has been rather thoroughly studied previously (Davydov 2001, 2004; Davydov & al. 2007; Makryi 1986; Sedelnikova 1990, 2003). At present the number of lichens known from the Altai Mts. exceeds one thousand species. Nevertheless, our data presented below shows that a considerable portion of the flora is still unrecognized and that considerable range extensions for many boreal European and North American species can be expected.

MATERIALS AND METHODS Herbarium specimens collected by the authors during a joint field trip in 2003 from the Altai Mountains (Siberia) composed the core material for

Figure 1. Study area in South Siberia.

this study (Fig. 1). Additional material was collected by the first author predominantly in the West Altai during 1995–2007. Voucher specimens are deposited in the herbaria ALTB and FR. In addition, collections deposited in B, H-NYL, HBG, KW, S, TRH & UPS were investigated. Morphological and anatomical characters were analysed by applying standard light microscopical methods. Chemical analyses of secondary products were made with standard thin-layer chromatography techniques (Culberson & Kristinsson 1970). Abbreviated collection data of examined specimens are provided for every species. Altitudes are indicated in meters above sea level. Names of collectors were reduced to ED and CP respectively.

RESULTS AND DISCUSSION Bacidia beckhausii Ko¨rb. This inconspicuous species is widespread in the boreal and temperate zone (see e. g. Esslinger 2011; Kurokawa 2003; Santesson et al. 2004; Urbanavichus 2010) but apparently rare in the area, as witnessed by the single collection we found. In Siberia it is reported from the South Baikal area (Urbanavichene 1998) and the West Siberian Plain around Surgut (Makarova et al. 2002). New for the Altai Mts. where it was collected in humid wood pasture with Larix sibirica, Pinus sibirica, Picea obovata and abundant dead wood, on Salix shrub in swamp. Specimen examined: ALTAI REPUBLIC, Ust’-Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8768 (FR). Bacidia hemipolia (Nyl.) Malme The species is rare in Europe and has a scattered distribution in Russia, where reports are based on old collections from Karelia in the European part

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(Norrlin 1876; Ra¨sa¨nen 1939) and from the Konda River basin on the West Siberian Plain (Vainio 1928). New for South Siberia, found on bark of Abies sibirica in subalpine Larix sibirica – forest with krummholz Abies sibirica. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Gladkaya griva Mt. 51u03910.00 N, 82u51950.90 E, alt. 1220 m, CP 8405 (FR).

Biatora chrysantha (Zahlbr.) Printzen [TLC: gyrophoric a.] One of the most widespread species of the genus, known from Europe, North America and Asia. In Russia the species occurs in the European part and Siberia (Urbanavichus 2010). Recently reported from dark taiga forests in Mongolia (Hauck & Javkhlan 2006; Palka & S´liwa 2006). One of the few species of the genus that tolerates moderately dry conditions in relatively open forests. Within the area it is relatively common on Abies sibirica and Betula spp. in Abies mountain taiga forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Kolyvansky range, Malaya Belaya River, 51u059 N, 82u299 E, alt. 800–1000 m, ED 6728 (ALTB); Tigireksky range, Bolshaya Uskuchevka River 51u11.29 N, 82u48.39 E, alt. 560 m, ED 5664, 5665 (ALTB); same range, 51u04.209 N, 82u50.549 E, alt. 764 m, ED 5666 (ALTB); same range, 50u59.79 N, 82u47.89 E, alt. 550 m, ED 5739 (ALTB); same range, Berlozhia River, 50u579340 N, 83u029590 E, alt. 717 m, ED 6729 (ALTB); same range, Bolshaya Uskuchevka River, 51u11919.30 N, 82u48930.60 E, alt. 450 m, CP 8308, 8310, 8326, 8328 (FR); same range Malaya Amelikha River, 51u04940.40 N, 82u49957.90 E, alt. 850 m, CP 8356, 8369 (FR); same locality, 51u04934.10 N, 82u50928.00 E, alt. 750 m, CP 8375 (FR); same range, Batalikha River, 51u03.09 N, 82u53.19 E, alt. 1525 m, ED 6727 (ALTB) [TLC: gyrophoric a., atranorin (tr)]. Biatora fallax Hepp This species that typically occurs in undisturbed humid coniferous forests is rare in central and northern Europe (Printzen 1995) and in Western North America (Printzen & Tønsberg 1999). In Russia, the species has a scattered distribution and occurs in the European North, North Ural, Arctic and South Siberia (in the South

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Baikal area) and the Northern part of the Far East (Urbanavichus 2010). New for the Altai Mts. where it grows in Abies sibirica dominanted mountain taiga relict forest with nemoral forbs, on bark of Abies sibirica. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Belaya River, 50u559590 N, 82u579250 E, alt. 720–800 m, ED 6798 (ALTB).

Biatora meiocarpa (Nyl.) Arnold This relatively rare species of the genus may have been overlooked in large parts of its range. In 1994 the only known and confirmed localities were located in Sweden, Finland and Russian Karelia (Ekman 1994). In the meantime, the species has also been reported from Central Europe (van den Boom 2002), Japan (Kurokawa 2003), the Caucasus and southwestern North America (Printzen 2004), and other parts of Russia (European part, North Ural, West and South Siberia, Far East; Urbanavichus 2010). The chemically distinct B. meiocarpa var. tacomensis (Printzen & Tønsberg) Printzen & Tønsberg grows in the Pacific West of North America and Central Norway (Printzen & Tønsberg 1999). Material from the Altai Mts. has relatively narrow spores. More material is needed to see whether it represents a distinct species. The reports of B. helvola and B. sylvana by Biazrov (1974), Cogt (1994) and Svirko (2003) may belong here. In the Altai it grows on the bark of Abies sibirica, Sorbus sibirica, Picea obovata and Salix sp. in coniferous forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Belaya River, 50u579340 N, 83u029590 E, alt. 717 m, ED 6796, 6797 (FR); same range, the velley of the Irkutka River 50u579380 N, 82u589410 E, alt. 650 m, ED 6799 (FR); same range, 50u579200 N, 82u589270 E, alt. 700–800 m, ED 6720 (ALTB), ED 6800 (FR); same range, Irkutka River, 51u019140 N, 82u589060 E, alt. 885 m, ED 6801 (FR); same range, Zagornaya Amelikha River, 51u03937.90 N, 82u51912.20 E, alt. 1010 m, CP 8392 (FR); ALTAI REPUBLIC, Ust’-Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8719, 8767 (FR). Biatora vernalis (L.) Fr. [TLC: no substances or trace of gyrophoric a. (contaminations)]

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Another circumboreal species of the genus reported from North America, Europe and Asia (e.g. Andreeva, 1983; Cogt 1994; Hauck & Javkhlan 2006; Spribille et al. 2010; Urbanavichus 2010). Literature reports must be treated with care though, because many unrelated species have in the past been subsumed under B. vernalis. Found only once by us, associated with B. chrysantha on old Betula pendula in Abies sibirica forest. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, valley of Bolshaya Uskuchevka River 51u11919.30 N, 82u48930.60 E, alt. 450 m, CP 8306 (FR).

Buellia arborea Coppins & Tønsberg [TLC: atranorin, placodiolic a.] This species was described based on Norwegian and Scottish collections (Tønsberg 1992), and was subsequently reported for several European countries (Bogdan 2002; Brackel & Kocourkova´ 2006; Foucard & Nordin 1999), European Russia (Urbanavichus 2010) and for North America (Spribille & Bjo¨rk 2008; Tønsberg & McCune 2001). New for Asia. In the Altai Mts. it is lignicolous on hard and smooth wood in open situations in forests. Buellia arborea shows a Holarctic disjunctive distribution and seems to be not rare in Asia, but without TLC it is possible to confuse it with Buellia griseovirens and Xylographa vitiligo, which both occur in the Altai Mts. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Irkutka River. 51u019140 N, 82u589060 E, alt. 885 m, ED 6747 (ALTB); ALTAI REPUBLIC, Ust’-Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8733, 8735 (FR). Buellia griseovirens (Turner & Borrer ex Sm.) Almb. [TLC: norstictic & connorstictic a., atranorin] In Russia the specied has a scattered distribution from the European part to East Siberia (Urbanavichus 2010). New for the Altai Mts. where it was collected on the bark of Picea obovata, Abies sibirica, Sorbus sibirica, and Betula pendula. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Zagornaya Amelikha River, 51u03.379 N, 82u51.129 E, alt. 986 m, ED 6753 (ALTB); Soloneshensky district, Bashchelaksky range, Shinok

River, 51u21942.50N, 83u34953.10E, alt. 870 m, CP 8557 (ALTB); ALTAI REPUBLIC, Ust’-Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8706 (ALTB); Ust’-Koksinsky district, Katunsky range, at the junction of the Khazinikha and Ioldo Rivers, 49u539 N, 86u049 E, alt. 1800 m, ED 6754 (ALTB); Ongudaysky district, 51u05939.70 N, 85u38917.10 E, alt. 1270 m, CP 8855 (FR).

Caloplaca herbidella (Hue) H. Magn. [TLC: no substances] The concept of the species was recently clarified by Arup and A˚kelius (2009). The species occurs in Europe (ibid.), the southern part of European Russia, the South Ural, Caucasus and the southern part of the Russian Far East (Urbanavichus 2010). New to Siberia. Both fertile (more common) on bark of Betula tortuosa in Pinus sibirica – Abies sibirica forest and sterile specimens on bark of Sorbus sibirica in Abies sibirica – tall forb forest, lacking orange pigment were collected in the Altai Mts. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Bol’shoi Tigirek River, 51u039050 N, 82u589370 E, alt. 1490 m, ED 6795 (ALTB); same range, Belaya River, 50u579340 N, 83u029590 E, alt. 717 m, ED 6751 (ALTB). Caloplaca sorocarpa (Vain.) Zahlbr. [TLC: rocellic a.] This relatively rare species is reported from few regions in Europe, North America (Wetmore 2004; Spribille et al. 2010) and Russia, where it was collected in the Komi Republic (Pystina 2003) and on Kamtshatka peninsula (Khodosovtsev et al. 2004). Caloplaca sorocarpa has an endophloeodic thallus and usually occurs on deciduous trees and shrubs. New to Siberia. In the region it was collected on Sorbus sibirica and Abies sibirica. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Irkutka River, 51u019140 N, 82u589060 E, alt. 885 m, ED 6717, 6791 (ALTB); same range, Belaya River, 50u579340 N, 83u029590 E, alt. 717 m, ED 6718 (ALTB). Fellhanera subtilis (Veˇzda) Diederich & Se´rus. The very inconspicuous species has a scattered distribution in North Eurasia and is also known from British Columbia in North America (Goward et al. 1996), although it is likely overlooked over much of

Davydov & Printzen: Lichens of Altai Mountains

its range. In Russia it is known from the European part, the North Ural, Caucasus and the Far East (Urbanavichus 2010). New for Siberia, where it was collected on bark of Betula pendula in Abies sibirica forest on a NEfacing slope. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, valley of Bolshaya Uskuchevka River, 51u11919.30 N, 82u48930.60 E, alt. 450 m, CP 8321 (FR).

Lecania croatica (Zahlbr.) Kotlov [TLC: no substances] A mostly sterile, sorediate species that was often overlooked or misidentified. Harris & Lendemer (2010) provided a thorough description and illustrations. Sterile specimens are characterized by small punctiform soralia and the lack of secondary metabolites in TLC. The species is widespread in Europe and Eastern North America. Reported here for the first time from Asia and Russia. In the Altai Mts. it grows on Salix sp. and Populus tremula in coniferous forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Bolshaya Uskuchevka River, 51u11919.30 N, 82u48930.60 E, alt. 450 m, CP 8312 (FR); Soloneshensky district, Bashchelaksky range, Shinok River, 51u21942.50 N, 83u34953.10 E, alt. 870 m, CP 8576 (FR); ALTAI REPUBLIC, Ongudaysky district, 51u05939.70 N, 85u38917.10 E, alt. 1270 m, CP 8856, 8858 (FR). Lecania cyrtellina (Nyl.) Sandst. Lecidea brenneri H. Magn., Svensk Bot. Tidskr. 30: 255 (1936). TYPE: RUSSIA: GUV. JENISEJSK, Gorinskij volok, 59u 209 N, auf glatter Rinde [bark of Sorbus aucuparia acc. to protologue], 03.X.1876, M. Brenner 465s (S – lectotype!, designated here, UPS – isolectotype!). Lecidea brenneri has not been reported since its original description by Magnusson (1936) based on collections made in 1876 by M. Brenner. In the protologue, Magnusson (1936) compared it to Lecidea sphaerella Hedl., but that species has ascospores 3.5–5.0 wide as opposed to 2.5–3.0 in L. brenneri. Printzen (1995) suggested a relationship with Lecania cyrtella (Ach.) Th. Fr., which is also

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characterized by larger ascospores of 4–5 mm in width. Comparison with material of Lecania cyrtellina (Nyl.) Sandst. showed that both are conspecific. The material collected in the Altai Mts. has rather dark apothecia and may be confused with Biatora globulosa Flo¨rke but is distinguished by scattered wide hyphal lumina in the hypothecium, rather wide excipular hyphae and a diffuse apothecial pigmentation that is not restricted to the epihymenium as in B. globulosa but extends throughout the hymenium. In the lectotype as well as in our collection some apothecia had a peculiar reddish brown pigmentation. Our collection was too small for chemical analysis. Due to its small apothecia and the endosubstratal thallus, the species may be overlooked. It was collected in forest of Picea obovata, Betula and Salix along creek, where it grew on bark of Salix. Additional specimen examined: ALTAISKY KRAI, Soloneshensky district, Bashchelaksky range, Shinok River, 51u21942.50 N, 83u34953.10 E, alt. 870 m, CP 8578 (FR). Specimens examined for comparison: Czech Republic, W Bohemia, Sˇumava Mts., SE slope of Mt. Stozˇec, 48u529 N, 13u509 E, alt. ca. 900 m, 5.VIII.1996, Z. Palice & C. Printzen (FR); S Bohemia, Sˇumava Mts., Volary, Cˇerny´ Krˇı´zˇ, Mt. Jeleni vrch, E slope, alt. 865 m, 15.V.2011, Z. Palice 14404 & V. Pouska (FR-dupl.). Germany, Hessen, Haidtra¨nktal im Taunus, alt. 400 m, X.1860, J. A. Metzler (FR). Russia, Guv. Tobolsk, Kalimski, 61uN, auf Espe [Populus tremula, Salix according to protologue], 31.V.1876, M. Brenner (UPS – paratype!).

Lecanora albellula var. albellula Nyl. [TLC: isousnic a., usnic a.] A widespread taxon in the northern hemisphere. Van den Boom and Brand (2008) provide a thorough description. The distinction from L. albellula var. macropycnidiata and L. pseudosarcopidoides requires chemical data and well developed pycnidia, which are not always easy to find. Literature reports of this species therefore have to be treated with care, and the distribution of the species has to be re-investigated. The record confirms the presence of this species in South Siberia. It was collected in subalpine Larix sibirica-forest with krummholz Abies sibirica on a trunk of Larix sibirica. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Gladkaya

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griva Mt. 51u03910.00 N, 82u51950.90 E, alt. 1220 m, CP 8406 (FR).

Lecanora boligera (Norman ex Th. Fr.) Hedl. [TLC: fumarprotocetraric a.] This species has a circumboreal distribution (Esslinger 2011; Urbanavichus 2010) New for West Siberia. It was found growing on bark of Picea obovata and Pinus sibirica in humid wood pasture with Larix sibirica, Pinus sibirica, Picea obovata and abundant dead wood. Specimen examined: ALTAI REPUBLIC, Ust’Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8704, 8711, 8798 (FR). Lecanora farinaria Borrer [TLC: atranorin, roccellic a.] This species belongs to the Lecanora subfusca group and occurs in Europe (see Kukwa & Kubiak 2007), North America (Esslinger 2011) and South Asia (Aptroot & Seaward 1999). Reported here as new for North Asia and Russia. It grows in Abies sibirica forest, on bark of Abies sibirica. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Belaya River, 50u569 N, 82u439 E, alt. 800–900 m, ED 6757 (ALTB); same range, Batalikha River, 51u02.39 N, 82u52.19 E, alt. 950 m, ED 6749 (ALTB). Lecanora pseudosarcopidoides M. Brand & van den Boom [TLC: isousnic a.] This apparently common but misunderstood species was only known from Europe (van den Boom & Brand 2008). New to Asia and Russia. The specimen was collected on wood of Larix sibirica in subalpine Larix sibirica-forest with krummholz Abies sibirica. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, ca. Gladkaya griva Mt. 51u03910.00 N, 82u51950.90 E, alt. 1220 m, CP 8411 (FR). Lecanora subintricata (Nyl.) Th.Fr. [TLC: usnic a.] A common circumboreal species of the ‘‘L. albellula’’-group characterized by its small ascospores, shallow hymenium and amphithecial cortex of regular thickness (Printzen 2001; van den Boom & Brand 2008). North American and European material of this species seem to differ in

secondary chemistry (usnic and 6 isosunic acids in North America, usnic acid and 6 brialmontin 1 in European collections). Only usnic acid was found in the material cited below. New to Altai Mts. The species grows on Pinus sibirica in humid wood pasture with Larix sibirica, Pinus sibirica, Picea obovata and abundant dead wood, and on dry stump in meadow in open forest of Picea obovata, Pinus sibirica, Larix sibirica and Betula sp. Specimens examined: ALTAI REPUBLIC, Ust’Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8778, 8814 (FR); Shebalinsky district, 51u05939.70 N, 85u38917.10 E, 1270 m, CP 8849 (FR).

Lecanora varia (Hoffm.) Ach. [TLC: usnic a., psoromic a.] A common, lignicolous western Eurasian species that seems to be largely absent from North America (Printzen 2001) and has not been reported from South Asia. It is easily identified by its chemistry and the peculiar anatomy of its amphithecium (Printzen 2001), however, literature reports of this species lacking these data have to be treated with care. Therefore, the record confirms the presence of this species in South Siberia. Common within the area. Specimen examined: ALTAI REPUBLIC, Ust’-Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8774 (FR); Ongudaysky district, 51u05939.70 N, 85u38917.10 E, alt. 1270 m, CP 8850 (FR). Lecidea betulicola (Kullh.) H. Magn. A relatively rare, apparently circumboreal species in Eurasia found mostly in old-growth forests. Reported for Russia from its European part, the North Ural and the northern part of the Far East (Urbanavichus 2010). In Siberia it was reported only once from its plain part (Kataeva et al. 2005). New to South Siberia. It appears to be also rare within the Altai region where it was collected in Abies sibirica dominanted mountain taiga relict forest (Chern’) on bark of Abies sibirica. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Belaya River, 50u579200 N, 82u589270 E, alt. 700–800 m, ED 6719 (ALTB, FR). Lecidea erythrophaea Flo¨rke ex Sommerf. This circumboreal species also is known from Europe, European and Asian Russia and North

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America, where it mostly grows on deciduous trees, but occasionally also on conifer bark. We collected it on Salix, Picea obovata and Abies sibirica. New to Altai Mts. Specimens examined: ALTAISKY KRAI, Soloneshensky district, Bashchelaksky range, Shinok River, 51u21943.60 N, 83u35920.40 E, alt. 910 m, CP 8537, 8539 (FR); same locality, 51u21942.50N, 83u34953.10E, alt. 870 m, CP 8560 (FR); ALTAI REPUBLIC, Ust’-Kansky district, 51u21924.00 N, 83u34906.00 E, alt. 670 m, CP 8639 (FR); same locality, 51u21921.40 N, 83u33950.20 E, alt. 1020 m, CP 8684 (FR).

Lecidea pullata (Norman) Th. Fr. This common, probably circumboreal species, is mostly found sterile and probably overlooked in the area by previous collectors. The species is widespread in North Europe and also known from British Columbia, Colorado, Que´bec and Alaska in North America (Goward & Thor 1992; Spribille & Bjo¨rk 2008; Spribille et al. 2010). Reported for Russia from its European part, the North Ural, Caucasus and the northern part of the Far East (Urbanavichus 2010). New to Siberia. Sterile specimens are characterized by the occurrence of sphaerophorin and isosphaeric acid. It was collecred on wood in humid wood pasture with Larix sibirica, Pinus sibirica, Picea obovata and abundant dead wood, and on bark of Betula sp. in open forest of Picea obovata, Pinus sibirica, Larix sibirica and Betula. Specimens examined: ALTAI REPUBLIC, Ust’Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8806 (FR); [TLC: sphaerophorin, isosphaeric a. (tr.)]; Ongudaysky district, 51u05939.70 N, 85u38917.10 E, alt. 1270 m, CP 8853, 8854 (FR); [TLC: sphaerophorin]. Lecidea sanguineoatra auct. non (Wulfen) Ach. This species, which is widespread in the Northern Hemisphere, belongs to a poorly understood species complex around Lecidea hypnorum. Some authors synonymize the name with L. hypnorum. We follow the concept of Aptroot et al. (2009). New to Asia and Russia. The species was collected in a stand of Abies sibirica, Betula pendula and Salix caprea, on bark of Salix caprea. Specimen examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Malaya

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Amelikha River, 51u04922.10 N, 82u50950.40 E, alt. 780 m, CP 8382 (FR).

Megaspora verrucosa (Ach.) Hafellner et V.Wirth var. mutabilis (Ach.) Nimis & Cl. Roux [TLC: no substances] Megaspora verrucosa is a cosmopolitan species, usually occurring on soil, bryophytes or plant debris in polar and alpine zones. It is widespread in the European and Asian parts of Russia (Urbanavichus 2010). Megaspora verrucosa var. mutabilis is distinguished by a shallow hymenium (ca. 100 mm) and slightly larger ascospores and grows on bark of trees in forests. The variety is new for West Siberia and the Altai Mts. where it was collected on bark of Salix sp. and Populus tremula in coniferous forests. Sedelnikova (1990) reported Megaspora verrucosa var. verrucosa for the high mountain belt of the Altai Mts. where it grows on soil or plant debris. Thus the varieties seem to display the same differences in altitudinal and substrate preferences in the Altai as in western North America (Nash & Ryan 2007). Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, 51u069490 N, 83u019010 E, alt. 1000 m, ED 6772 (ALTB); ALTAI REPUBLIC, Ust’-Koksinsky district, Katunsky range, 49u539 N, 86u049 E, alt. 1800 m, ED 6773 (ALTB). Pertusaria coccodes (Ach.) Nyl. [TLC: norstictic & connorstictic a.] This subcosmopolitan species is known in Russia from the European part, the Ural Mts and the Far East (Urbanavichus 2010). New for Siberia. It was collected on bark of Abies sibirica at timberline, and on Picea obovata in the forest of Picea obovata, Betula and Salix. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Batalikha River, 51u03.09 N, 82u53.19 E, alt. 1525 m, ED 6739 (ALTB); Soloneshensky district, Bashchelaksky range, Shinok River, 51u21942.50 N, 83u34953.10 E, alt. 870 m, CP 8555 (FR). Pertusaria pupillaris (Nyl.) Th. Fr. [TLC: fumarprotocetraric or fumarprotocetraric & protocetraric a.] The species is reported from Western Europe, North America (Spribille et al. 2010; Tønsberg 1993)

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and from a few localities in Russia from the European North, North Ural and West Siberian Plain (Urbanavichus 2010). New for South Siberia. It was collected on bark of Abies sibirica, Picea obovata and Betula pendula in coniferous mountain forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Belaya River, 50u579200 N, 82u589270 E, alt. 700–800 m, ED 6740 (ALTB); same range, 50u579340 N, 83u029590 E, alt. 717 m, ED 6743, 6744 (ALTB); ALTAI REPUBLIC, Ust’Kansky district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8755 (FR).

Pycnora leucococca (R. Sant.) R. Sant. [TLC: alectorialic a.] One of the most common sterile sorediate crusts in the North American boreal forests (Spribille & Bjo¨rk 2008) but under-reported in Eurasia where it is known so far from parts of Europe and Russia (northern and central parts of European Russia, the northern and southern Ural Mts, Arctic Russia, West and South Siberia, Chukchi Peninsula; Urbanavichus 2010). New for the Altai Mts. where it was collected on bark of Abies sibirica and on wood in coniferous mountain forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range Belaya River, 50u579200 N, 82u589270 E, alt. 700–800 m, ED 6723 (ALTB); ALTAI REPUBLIC, Ust’-Kansky district, 51u21921.40 N, 84u33950.20 E, alt. 1020 m, CP 8661 (FR); same district, 50u43949.70 N, 85u24915.80 E, alt. 1680 m, CP 8730, 8743 (FR); Ust’-Koksinsky district, Katunsky range, Ozyornaia River, 49u489 N, 85u499 E, alt. 1516–1700 m, ED 6724 (ALTB). Puttea exsequens (Nyl.) Printzen & Davydov comb. nov. [5 L. gibberosa sensu Th. Fr. non Ach.] BASIONYM: Lecidea exsequens Nyl., Flora 64: 179 (1881). TYPE: ROMANIA, infra montem Domugled in Banatu, ad lignum putrescens, 1874, H. Lojka 2713 (H-NYL 21346 – holotype!). This species is rarely collected and hitherto only reported from Europe but is probably much more widespread as witnessed by the two collections cited below. Apart from the Latin protologue there are

only a few brief descriptions in Latin (e.g. Fries 1874, under L. gibberosa, Arnold 1887), German (e.g. Lettau 1912, Lindau 1923) or Swedish (Foucard 2001). We therefore give a brief description here. Description: Thallus crustose, very thin and whitish or endosubstratal, esorediate. Apothecia rounded to slightly irregular in outline (Fig. 2), sessile with a weakly constricted base, 0.3–0.7 mm diam., flat to moderately convex, disc ochre or orange- to red-brown, matte, with a pruinose appearance when wet, margin lacking or only slightly prominent, mostly concolorous or slightly lighter than disc. Hypothecium colourless, 25–175 mm, hyphae with narrow lumina of c. 1 mm. Exciple colourless, covered by a granular epipsamma, composed of strongly gelatinized anticlinal hyphae with narrow lumina of 0.5–1.0, apically to 1.2 mm. Hymenium 45–55 mm high, colourless with a yellowish-ochre granular epihymenium of 5–10 mm, granules often streaking into hymenium. Paraphyses weakly to moderately branched and anastomosing, with thin lumina of 0.8–1.0, apically to 1.2 mm. Asci 8-spored, of the Pilocarpaceae-type, with a fine amyloid tube structure in the tholus. Ascospores colourless, fusiform-ellipsoid, 8–11 3 3.0–4.5 mm. Pycnidia not seen by us (cylindrical and 4–6 mm long according to Fries 1874). Chemistry: usnic acid detected by TLC in one specimen. Distribution and habitat: Europe, new to Asia and Russia. The species was growing on wood, the typical substratum, in forest of Picea obovata along a creek. Discussion: Originally described from Romania, Lecidea exsequens has been informally treated as the correct name for the Scandinavian L. gibberosa sensu Th. Fr. non Ach. by Printzen (1995) and Spribille et al. (2008). Vainio (1934) first discovered that L. gibberosa as treated by Fries (1874) was not conspecific with L. gibberosa Ach. The type material of L. gibberosa Ach. [‘‘Germania, Schrader’’ H-ACH 250] consists of a few damaged apothecia on wood. Vainio (1934), who was unable to safely identify the material, suggested that it belongs to a Lecanora. We confirm this observation. In Santesson (1984) and following checklists and identification keys of Scandinavian lichens (e.g. Foucard 2001) L. exsequens was treated as ‘‘L. gibberosa sensu Th. Fr. non Ach.’’.

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Figure 2. Puttea exsequens. A. Habit. B. Apothecial section in water. C. Apothecial section in lactophenol cotton blue. Scale – A: 1 mm; B, C: 50 mm.

Because of its anatomical similarity with Puttea margaritella (Hulting) S. Stenroos & Huhtinen (see below) it is here transferred to Puttea. Differentiation: Puttea exsequens resembles species of Biatora, few of which occur on wood. From these it can easily be distinguished by the narrow excipular hyphae. Biatora species with an epipsamma include B. flavopunctata (Tønsberg) Hinteregger &

Printzen and B. vacciniicola (Tønsberg) Printzen, which are sorediate. Biatora aegrefaciens Printzen and B. vacciniicola are best distinguished by their C+ red reaction of apothecial sections due to the content of gyrophoric acid. Biatora subgilva (Arnold) Hinteregger has smaller apothecia of 0.2–0.5 mm diam. and considerably longer ascospores of 10–27 3 3–4 mm. Puttea exsequens could also be confused

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with Lecanora symmicta (Ach.) Ach., but can be distinguished on account of its fusiform ascospores and the Pilocarpaceae-type ascus. Lecanora symmicta is also uncommon on wood. Puttea margaritella, Lecidea symmictella Nyl. and Catillaria retigena (Hedl.) H. Magn. are the most similar species. All three have narrower ascospores of 1.5–3.0 mm width. Catillaria erysiboides is also externally similar but characterized by distinctly two-celled spores. Specimens examined: RUSSIA: ALTAI REPUBLIC, Ust’-Kansky district, 51u21921.40 N, 84u33950.20 E, alt. 1020 m, CP 8658 (FR); same locality, 51u21924.00 N, 83u34906.00 E, alt. 670 m, CP 8629 (FR). Additional specimens examined: AUSTRIA: Steiermark, Sulzbach, VIII.1876, J. Glowacki (HBG). NORWAY: Sør-Trøndelag, Selbu, by river Ra˚na, c. 345 m, 22.VI.2007, J. T. Klepsland L-099 (TRH). ROMANIA: Ad rivam rivuli ‘‘Riu sor’’ infra alpem Retyeza´t, com. Hunyad in Transsylvania, 1873, H. Lojka 2354 (FR). SWEDEN, So¨dermanland, St. Malm, Sandviksskogen, 6.VII.1895, G.O.A. Malme [Malme, Lich. suec. exs. 44] (B); Uppland, Uppsala par., Fredrikslund, i barrskog, [on wood], 27.VII.1862, T. M. Fries (UPS, under Lecidea gibberosa) [TLC: usnic a.].

Pycnora sorophora (Vain.) Hafellner [TLC: alectorialic a.] This often overlooked species is so far known from Europe, Russia (North and Center of the European part, North and South Ural, West Siberia; Urbanavichus 2010), and North America (Spribille & Bjo¨rk 2008). New for South Siberia. The species grows on wood or on bark of Abies sibirica in coniferous mountain forests. Specimens examined: ALTAISKY KRAI, Krasnoshchokovsky district, Tigireksky range, 51u11922.60 N, 82u59929.60 E, alt. 925 m, ED 6793 (ALTB); ALTAI REPUBLIC, Ust’-Koksinsky district, Katunsky range, Ozyornaia River, 49u489 N, 85u499 E, alt. 1516–1700 m, ED 6725 (ALTB). Rimularia fuscosora Muhr & Tønsberg [TLC: norstictic a.] The species was known only from Norway, Sweden and Scotland (Tønsberg 1992). New to Asia and Russia. It was collected in flooded Betula pendula forest on bark of Betula pendula.

Specimen examined: ALTAISKY KRAI, Krasnoshchokovsky district, Tigireksky range, Tigirek, 51u089 N, 83u039 E, alt. 480 m, ED 6752 (ALTB).

Rinodina excrescens Vain. [TLC: pannarin] The species was described from West Siberia by Vainio (1928) and is known from very few localities in Europe (Giralt et al. 1994; Spribille et al. 2006) and eastern North America (Sheard & Tønsberg 1995; see Galanina et al. 2011 for more details). The species was recently found in Sokhondinskiy Zapovednik (Dahuria, South Siberia) and in many localities in the Russian Far East (Galanina et al. 2011) where Rinodina excrescens is a widespread and primarily corticolous species on the bark of deciduous and coniferous trees. New for the Altai Mts. where it was collected in forest of Picea obovata, Betula and Salix along a creek, on a trunk of Betula. Specimens examined: ALTAISKY KRAI, Soloneshensky district, Bashchelaksky range, Shinok River, 51u21942.50N, 83u34953.10E, alt. 870 m, CP 8550 (FR). Scoliciosporum chlorococcum (Graewe ex Stenh.) Veˇzda Syn. nov.: Bacidia zerovii Oxner Botanicheskiy Zhurnal Akad. Nauk Ukrain. SSSR 5: 95 (1948) TYPE: [RUSSIA], North Altai, Oyrotia, Shebalinsky district, Olygash, Pinus sibirica forest with Larix, below pass to Sarychet Mt., on Berberis sibirica, 17.VIII.1940, A. Oxner (KW – holotype!). Oxner described Bacidia zerovii from the Altai and, due to the size and colour of its apothecia and ascospores, suggested that it is isolated in the genus and probably related to the Bacidia compacta group (Oxner 1948). Reports of further authors (Golubkova 2003; Sedelnikova 1990; Urbanavichus 2010) were based on the single collection made by Oxner. The holotype of Bacidia zerovii Oxner agrees in all studied characters with material of Scoliciosporum chlorococcum (Graewe ex Stenh.) Veˇzda and we conclude that the name is a synonym of this widely distributed Holarctic species.

Scoliciosporum perpusillum J. Lahm ex. Ko¨rb. This inconspicuous species was reported only from few localities in Russia in the South Baikal area

Davydov & Printzen: Lichens of Altai Mountains

(Urbanavichene & Urbanavichus 1999) and on the Kola Peninsula (Kuusamo; Urbanavichus et al. 2008). New for West Siberia. The species grows on bark of Abies sibirica in coniferous mountain forests. Specimens examined: ALTAISKY KRAI, Zmeinogorsky district, Tigireksky range, Bolshaya Uskuchevka River, 51u11.29 N, 82u48.39 E, alt. 560 m, ED 6780 (ALTB); same range Belaya River, 50u59.79 N, 82u47.89 E, alt. 550 m, ED 6779 (ALTB); same locality, 51u00.39 N, 82u45.89 E, alt. 540 m, ED 6777 (ALTB); same locality, 50u579200 N, 82u589270 E, alt. 700– 800 m, ED 6778 (ALTB); ALTAI REPUBLIC, Ust’-Koksinsky district, Katunsky range, Khariuzovka River, 49u499 N, 85u489 E, alt. 1516–1800 m, ED 6781 (ALTB).

ACKNOWLEDGMENTS We gratefully acknowledge help from the curators and staff of the herbaria B, H-NYL, HBG, KW, S, TRH, & UPS. The authors thank G. P. Urbanavichus and D. E. Himelbrant for their comments on some species’ distributions. We are very grateful to T. Spribille and C. Bjo¨rk who reviewed the manuscript for their valuable suggestions. This study was financially supported by the Deutsche Forschungsgemeinschaft (DFG grant No.436 RUS 17/36/05).

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