Stereocaulon from the Holarctic, with a Key to the Known Species
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Opuscula Philolichenum, 8: 9-39. 2010. Lichenicolous fungi and lichens growing on Stereocaulon from the Holarctic, with a key to the known species MIKHAIL P. ZHURBENKO1 ABSTRACT. – A total of 22 species of lichenicolous fungi and three lichens are reported on Stereocaulon species and discussed. Lichenopeltella stereocaulorum sp. nov., Lichenosticta dombrovskae sp. nov. and Odontotrema stereocaulicola sp. nov. are described from Stereocaulon. Arthonia stereocaulina, Opegrapha stereocaulicola, Rhymbocarpus stereocaulorum and Sphaerellothecium stereocaulorum are reported new to North America (except Greenland); Endococcus nanellus is new to Kazakhstan, Alaska and Greenland; Arthonia stereocaulina and Opegrapha stereocaulicola are new to Svalbard; Catillaria stereocaulorum, Cercidospora stereocaulorum, Lasiosphaeriopsis stereocaulicola, Polycoccum trypethelioides and Taeniolella christiansenii are new to the U.S.A.; Catillaria stereocaulorum and Niesslia peltigericola are new to the Canadian Arctic; Lasiosphaeriopsis stereocaulicola and Taeniolella christiansenii are new to British Columbia; and Catillaria stereocaulorum is new to Mongolia. Anzina carneonivea, Endococcus rugulosus s. l., Niesslia peltigericola, Phaeosporobolus alpinus and Protothelenella sphinctrinoidella are newly documented on Stereocaulon. Seventeen species of Stereocaulon are new hosts for various lichenicolous fungi. A key to the 39 species of fungi and lichens known to occur on Stereocaulon is provided. Study of the host relationships of lichenicolous fungi on Stereocaulon does not support a close relationship between the crustose and fruticose species of the genus. Terricolous arctic species of Stereocaulon with more solid thallus structure are more frequently colonized by fungi when compared to species with lax thalli. INTRODUCTION Stereocaulon Hoffm. is a widespread lichen genus comprising 137 species, most of which are fruticose macrolichens (Kirk et al. 2008). A decade ago just nine species of lichenicolous fungi were known to occur on members of the genus (Zhurbenko 2000). However, recent studies have revealed much a higher diversity of stereocaulicolous fungi (Etayo 2002, Ihlen & Wedin 2007, Zhurbenko & Triebel 2008, Zhurbenko et al. 2009). The aims of this paper are: 1) describe three new species of stereocaulicolous fungi and provide information about another two undescribed species of unclear taxonomic position; 2) present new information on ecology, geography and species composition of mycobiota growing on Stereocaulon; 3) provide a worldwide key for these fungi; and 4) test the relationship of crustose and fruticose species of Stereocaulon on the basis of their associated fungi. MATERIALS AND METHODS The study is based on 392 samples of lichenicolous fungi and lichens from 27 regions of the Holarctic, mostly from the Arctic (Plate 1). These finds occurred on 20 fruticose host species (Stereocaulon alpestre, S. alpinum, S. arcticum, S. arenarium, S. botryosum, S. capitellatum, S. depressum, S. glareosum, S. grande, S. groenlandicum, S. incrustatum, S. intermedium, S. myriocarpum, S. paschale, S. rivulorum, S. saxatile, S. subcoralloides, S. symphycheilum, S. tomentosum and S. vesuvianum) and one crustose host species (S. plicatile). Two hundred fifty-three specimens were collected by the author; the others were found in undetermined lichen gatherings from various collectors (see acknowledgements) donated to the 1 MIKHAIL P. ZHURBENKO — Lab. of the Systematics and Geography of Fungi, Komarov Botanical Institute, Russian Academy of Sciences, Professor Popov 2, St.-Petersburg, 197376, Russia. – e-mail: mzhurb@g- mail.com 9 Plate 1. Geographical location of the main collecting regions: 1 – Alaska (USA), 2 – British Columbia (CANADA), 3 – Canadian Arctic Archipelago (CANADA), 4 – Greenland (DENMARK), 5 – Svalbard (NORWAY), 6 – Troms County (NORWAY), 7 – Murmansk Region (RUSSIA), 8 – Karelia (RUSSIA), 9 – Nenets Region (RUSSIA), 10 – Franz Josef Land (RUSSIA), 11 – Polar Ural (RUSSIA), 12 – KAZAKHSTAN, 13 – Gydan Pen. (RUSSIA), 14 – Severnaya Zemlya (RUSSIA), 15 – Taimyr Pen. (RUSSIA), 16 – Putorana Plateau (RUSSIA), 17 – Evenkiya (RUSSIA), 18 – Tyva, Sayan and Altai Mts. (RUSSIA), 19 – Baikal Siberia (RUSSIA), 20 – MONGOLIA, 21 – mainland Yakutiya (RUSSIA), 22 – New Siberian Islands (RUSSIA), 23 – Khabarovsk Territory (RUSSIA), 24 – Magadan Region (RUSSIA), 25 – Kamchatka Pen. (RUSSIA), 26 – Wrangel’ Island (RUSSIA), 27 – Chukotka (RUSSIA). author. Crustose Stereocaulon species were not specifically searched for the associated fungi. Endophytic fungi of Stereocaulon species (see Petrini et al. 1990) were omitted from this study. Examined materials are mostly housed in the LE-Fungi herbarium; some specimens are deposited in H and M. The material was studied and photographed using Zeiss Stemi 2000-CS microscopes and an Axio Imager A1 equipped with Nomarski differential interference contrast optics. Microscopical examination was done in water, 10 % KOH (K), Lugol’s iodine, directly (I) or after a KOH pre-treatment (K/I), HNO 3, Brilliant Cresyl blue (BCr) or phloxin (1% in water) solutions. The length, breadth and length/breadth ratio (l/b) of asci, ascospores and conidia are mostly given as: (min–){ X -SD}–{ X +SD}(–max), where min and max are the extreme values, X the arithmetic mean and SD the corresponding standard deviation. Microscopical measurements were made in water, unless otherwise indicated. Sizes of asci were rounded to the nearest 1 μm, those of ascospores and conidia to the nearest 0.5 μm. Terms for simple plane shapes and colors mostly follow Stearn (1992: 539) and Petersen (1996), respectively. Classification and nomenclature of taxa above the generic level follow Lumbsch & Huhndorf (2007). Authors of host taxa are omitted throughout this paper. THE SPECIES Abbreviations: Is. – Island, Mt(s). – Mountain(s), Pen. – Peninsula. Lichens are designated by “*”. 10 1. *Anzina carneonivea (Anzi) Scheid. NOTES. – This lichen usually grows on mosses, bark or lignum (Santesson et al., 2004). It has also been reported on lichens such as Cladonia and Peltigera (e.g. Hafellner 2000), but not previously on Stereocaulon. o o SPECIMEN EXAMINED. – RUSSIA. KARELIA: Kandalakshskii Gulf of White Sea, Keret’ Is., 66 19’N, 33 42’E, alt. 40 m, rocks, on Stereocaulon saxatile (healthy-looking phyllocladia), 9.vii.1998, D. Himelbrant (LE 210402:b). 2. Arthonia stereocaulina (Ohlert) R. Sant. NOTES. – Ascomata apothecial, usually superficial, convex, subglobose or irregular in shape, sometimes applanate from above or mostly immersed and only slightly elevating above the host thallus surface, (0.05–)0.1–0.25(–0.5) mm diam., smooth, black, shiny, arising singly or more often aggregated to confluent. Epithecium olive-brown. Hymenium I+, K/I+ purple to brown red. Hypothecium brown. Asci (25–)26–35.5(–38) × (10–)13–21.5(–25) µm (n = 10). Ascospores hyaline, soleiform to narrowly ovate or lanceolate, 1-septate, usually constricted at the septum, with unequal cells (usually the upper one boader), (10.5–)11.5–14(–17) × (4–)4.5–6(–7.5) µm, l/b = (1.7–)2.1–2.9(–3.9) (n = 141). Found on phyllocladia, rarely on stems of Stereocaulon alpinum, S. arcticum, S. botryosum, S. capitellatum, S. depressum, S. glareosum, S. groenlandicum, S. intermedium, S. myriocarpum, S. paschale, S. rivulorum, S. saxatile and S. tomentosum. Mostly commensalistic, rarely causes slight bleaching of host tissues. Stereocaulon arcticum, S. intermedium, S. myriocarpum, S. paschale, S. saxatile and S. tomentosum are new host species. New to North America (except Greenland), Svalbard, Nenets Region, Sayan Mts., Baikal Siberia and mainland Yakutiya. SPECIMENS EXAMINED. – U.S.A. ALASKA: Great Kobuk Sand Dunes, 67º06’N, 159º01’W, alt. 50 m, lichen heath with sparse Picea, on S. glareosum, 11.viii.2000, M. Zhurbenko 00464 (LE 233527); Wrangell-Saint Elias Park and Preserve, Mt. George, 60º59’N, 141°28’W, alt. 1783 m, mountain tundra, on S. alpinum, 17.vii.2004, T. Jorgenson (LE 233427:a). CANADA. CANADIAN ARCTIC ARCHIPELAGO: Ellef Ringnes Is., Isachsen Bay, 78°47’N, 103°32’W, alt. 70 m, stony arctic desert, on S. groenlandicum, 25.vii.2005, N. Matveeva (LE 233750); on S. botryosum, 20.vii.2005, N. Matveeva (LE 233592). SVALBARD. Nordenskiöld Land, 3 km N of Barentsburg, 78º05’N, 14º13’E, alt. 100 m, wet scree, on S. rivulorum, 14.vii.2003, M. Zhurbenko 03268 (LE 233759); W coast of Grønfjorden, Aldegondabreen Glacier, 78º00’N, 14º12’E, alt. 100 m, fellfield in tundra, on S. botryosum (phyllocladia, rarely stems), 16.vii.2003, M. Zhurbenko 03264 (LE 233457). RUSSIA. MURMANSK REGION: Khibiny Mts., Akademicheskoe Lake, 67°43 N, 33°49 W, alt. 700 m, fellfield in tundra, on S. saxatile, 20.viii.2007, M. Zhurbenko 0748 (LE 210438). – NENETS REGION: Bol’shezemel’skaya Tundra, Ortin River, Vorgashor Stream, 67°50’N, 54°03’E, dwarf shrub tundra on sand, on S. arcticum, 29.vi.1999, O. Lavrinenko (LE 233437); on S. rivulorum, 29.vi.1999, O. Lavrinenko (LE 233484:b); Khar’yaga oilfield, 67º11’N, 56º30’E, alt. 60 m, tundra, on S. paschale, 24.vii.2007, M. Zhurbenko 0718 (LE 210330). SEVERNAYA ZEMLYA: Bol’shevik Is., Ostantsovaya River, 79°13’N, 102°02’E, alt. 40 m, rocks, on S. glareosum, 12.vii. 1996, M. Zhurbenko 96947:a (LE 233407:a); Mushketova Glacier, 79°11’N, 102°09’E, alt. 200 m, stony arctic desert, on S. botryosum, 11.vii.1996, M. Zhurbenko 96112:c (LE 233572:c). TAIMYR PEN.: Eniseyskii Gulf coast, Ragozinka River mouth, 72°48’N, 80°53’E, alt. 10 m, dwarf shrub tundra, on S. rivulorum, 11.vii.1990, M. Zhurbenko 90373 (LE 207709); same coast, Osipovka River, 72°42’N, 80°51’E, alt. 40 m, boulder field in tundra, on S. botryosum, 18.vii. 1990, M. Zhurbenko 90831:a (LE 233709:a), M. Zhurbenko 90828 (LE 233810); on S. groenlandicum, 18.vii.1990, M. Zhurbenko 90839:d (LE 233729:d); lower Pravaya Uboinaya River, "217" Mt., 73°26’N, 82°43’E, alt. 150 m, stony tundra, on S. groenlandicum, 5.viii.1990, M. Zhurbenko 90841 (LE 233830); on S. botryosum, 5.viii.1990, M. Zhurbenko 90832:b (LE 233930:b); Byrranga Mts., N of Levinson-Lessinga Lake, 74°31’N, 98°36’E, alt. 330 m, rocks in tundra, on S. groenlandicum, 27.vii.1994, M. Zhurbenko 94504 (LE 233547); S of Levinson-Lessinga Lake, 74°24’N, 98°46’E, alt.