Opuscula Philolichenum, 17: 368-373. 2018. *pdf effectively published online 12November2018 via (http://sweetgum.nybg.org/philolichenum/)

Contributions to the knowledge of lichenicolous fungi on Thamnolia

MIKHAIL P. ZHURBENKO1 AND YOSHIHITO OHMURA2

ABSTRACT. – Capronia thamnoliae, Cercidospora thamnoliae and Cercidospora thamnoliicola are reported new to ; Endococcus thamnoliae new to Papua New Guinea; Merismatium thamnoliicola new to Norway; Polycoccum vermicularium new to Argentina, , Colombia, Japan, Nepal and Papua New Guinea; Sphaerellothecium thamnoliae var. taimyricum new to Nepal; S. thamnoliae var. thamnoliae new to Australia, Falkland Islands, Japan and Papua New Guinea; Sphaeropezia santessonii new to Japan; Stigmidium frigidum new to Japan; Thamnogalla crombiei new to Australia, Bhutan, India, Japan, Nepal, North Korea and Papua New Guinea. Cornutispora ciliata is newly reported from Thamnolia. Material of Polycoccum vermicularium is shown to be morphologically heterogeneous.

KEYWORDS. – Biodiversity, biogeography, lichenicolous mycobiota.

INTRODUCTION

Thamnolia Ach. ex Shaer. (Icmadophilaceae) is among the host lichen genera with the highest diversity of known lichenicolous fungi (Diederich et al. 2018b). At least 26 species of these fungi have been documented from Thamnolia (Diederich et al. 2018a; Zhurbenko 2012, 2017; also herein). Although thamnoliicolous fungi have been specifically treated by Ihlen (1995) and Zhurbenko (2012), including an identification key presented in the latter, their distribution and frequency is still poorly known. Here we provide new distributional records, range extensions and taxonomic notes on thamnoliicolous fungi that were found in conjunction with examining material at several major herbaria.

MATERIALS AND METHODS

This study is based on 81 specimens of lichenicolous fungi found on Thamnolia deposited in H (33 specimens) and TNS (48 specimens). Microscopical examination was carried out using a Zeiss Axio Imager A1 microscope equipped with Nomarski differential interference contrast optics (DIC) in water, 10% KOH, Lugol’s iodine, directly or after a KOH pre-treatment or brilliant cresyl blue. The length, breadth and length/breadth ratio (l/b) of ascospores are given as: (min−){X−SD}−{X+SD}(−max), where “min” and “max” are the extreme observed values, X the arithmetic mean and SD the corresponding standard deviation. Measurements were taken from water mounts.

NOTES AND NEW DISTRIBUTIONAL RECORDS

Recent phylogenetic studies published by Onut-Brännström et al. (2018) have drastically changed the taxonomy of Thamnolia, which currently comprises three species: 1) T. subuliformis (Ehrh.) W.L.Culb. s.str. including two chemotypes with baeomycesic and squamatic acids (UV+) or thamnolic acid (UV−)

1MIKHAIL P. ZHURBENKO – Laboratory of the Systematics and Geography of Fungi, Komarov Botanical Institute, Russian Academy of Sciences, Professor Popov Street, 2, St. Petersburg, 197376, Russia. – e- mail: [email protected] 2YOSHIHITO OHMURA – Department of Botany, National Museum of Nature and Science, Amakubo 4-1-1, Tsukuba, Ibaraki, 305-0005, Japan. – e-mail: [email protected]

368 and exhibits a distribution that is circumpolar in the Arctic and in the alpine areas of both hemispheres; 2) T. tundrae Brännström & Tibell containing baeomycesic and squamatic acids (UV+) and known from the Eurasian Arctic and the Aleutian Islands; 3) T. vermicularis (Sw.) Schaer. s.str. containing thamnolic acid (UV−) and known from the high alpine regions of Central Europe (the Alps, Tatra Mountains and the Western Carpathian Mountains). These species are morphologically cryptic, in part sympatric, and can only be confidently identified using molecular data. As such the original identifications of Thamnolia host species for the taxa reported here should be revised. However, such revision is problematic because many specimens have not been sequenced. Due to this, and the fact that most of thamnoliicolous fungi species appear not to be restricted to a single host species, we omit below the species names of the hosts in the specimen citations below.

Capronia thamnoliae Zhurb. NOTE. – The species was formerly known from several finds in Canada, Norway, Russia and the U.S.A. (Zhurbenko 2012) and is here newly reported for Japan. Specimens examined (all on decaying podetia of Thamnolia spp.). ‒ JAPAN. : Shinano Province (Nagano Prefecture), Yatsugatake Mountains, Mount Yoko-dake, Chino-shi, 35°59′N 138°22′E, elev. 2700 m, 5.viii.1990, H. Shibuichi 8782a (TNS); Shinano Province (Nagano Prefecture), Yatsugatake Mountains, Mountain Ioh-dake, 36°00′N 138°22′E, elev. 2700 m, 9.viii.1988, H. Shibuichi 8427 (TNS). U.S.A. ALASKA: Juneau Icefield, summit of mountain above Taku Glacier, 58°42′N 134°13′W, elev. 1460 m, 19.vii.1962, H.A. Imshaug 28916a (TNS).

Cercidospora thamnoliae Zhurb. NOTE. – The species was formerly known from Norway and Russia (Ihlen 1995, Zhurbenko 2012) and is here newly reported for Japan. Specimen examined. ‒ JAPAN. HONSHU: Iwashiro Province (), Iide Mountains, , Yama-gun, 37°51′N 139°42′E, elev. 2000 m, 26.viii.1975, H. Shibuichi 5127a (TNS).

Cornutispora ciliata Kalb NOTES. – The species grows on various lichen genera (Diederich et al. 2018b) and is here newly reported from Thamnolia. The only species of Cornutispora previously known to occur on Thamnolia is C. intermedia Punith. & D.Hawksw. (Etayo 2010). Specimen examined. ‒ U.S.A. ALASKA: Alaska Range, Mountain Hayes Quadrangle, Gerstle River outwash gravel, 63°51′N 144°53′W, elev. 375 m, 23.vi.1966, L.A. Viereck 7971a (H).

Endococcus thamnoliae Etayo & R.Sant. NOTES. – Based on the material that we have examined, the range of variation in the size of the ascospores is somewhat larger than indicated in the protologue ((7.4‒)8.8‒11.0(‒13.2) × (3.2‒)3.7‒4.5(‒5.4) µm, l/b = (1.8‒)2.1‒2.7(‒3.4), n = 133 vs. 9‒12.5 × 3.5‒5 µm fide Etayo 2010). The species was formerly known from Peru (Etayo 2010) and is here reported for the first time outside South America from Papua New Guinea. Specimens examined. ‒ PAPUA NEW GUINEA. WESTERN HIGHLAND DISTRICT: between Pindaunde Lakes and summit of Mount Wilhelm, elev. 4100‒4200 m, 31.xii.1973, H. Kashiwadani 11006 (TNS), H. Kashiwadani 10831 (TNS). CENTRAL DISTRICT: Mount Albert Edward, elev. 3000‒3450 m, 25.x.1975, S. Kurokawa 9401 (TNS). PERU. JUNIN REGION: Jauja Province, 30 km (road distance) NNW of Jauja, 11°35′S 75°35′W, elev. 4100 m, 10.ii.1981, R. Santesson et al. s.n. (H); Tarma Province, between La Oroya and Junin, S of the turn towards San Pedro, 11°17′S 75°56′W, elev. 4200 m, 16.ii.1981, R. Santesson & R. Moberg s.n. (H).

Lichenopeltella thamnoliae R.Sant. NOTE. – The species is known only from South America, where it is rather widely distributed (Etayo 2010, 2017; Santesson 1998; Zhurbenko 2012). Specimens examined. ‒ COLOMBIA. BOYACÁ DEPARTMENT: Páramo de Pisva, 5 km E of Los Pinos, elev. 3750 m, 14.vi.1972, A.M. Cleef 4540 (H); Páramo de la Sarna between Sogamoso and Vado Hondo, 5 km NE of Laguna de Tota, elev. 3550 m, 29.iii.1973, A.M. Cleef 9203 (H). VENEZUELA. MÉRIDA: near Pico el Aguilo, elev. 4025 m, 18.viii.1989, R. Ornduff s.n. (H).

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Merismatium thamnoliicola Alstrup & E.S.Hansen NOTE. – The species was formerly known from Greenland and Russia (Alstrup & Hansen 2001, Zhurbenko 2012) and is here newly reported for Norway. Specimen examined. ‒ NORWAY. SÖR-TRÖNDELAG: Kongsvoll, vii.1933, E.P. Vrang s.n.[c] (H).

Polycoccum vermicularium (Linds.) D.Hawksw. NOTES. – The dimensions of the ascospores calculated from all the examined specimens vary considerably ((12.9‒)15.4‒18.8(‒22.0) × (6.4‒)7.3‒8.7(‒9.6) µm, l/b = (1.6‒)2.0‒2.4(‒2.8), n = 238). However, in the material from South America the length on average was somewhat shorter (14.6‒18.2 µm) than in the material from the northern Holarctic (16.4‒19.8 µm), which suggests that the material is taxonomically heterogeneous. It is also noteworthy that typically ascomata are aggregated in dense groups and often protruding in the ostiolar area, but in some well-developed specimens from South America (e.g., Cleef 1786, Imshaug 40468b, Imshaug 41387 & Harris, Imshaug 53198 & Ohlsson) they are exclusively dispersed and not protruding. The species is widely distributed in both hemispheres (Alstrup & Hawksworth 1990, Etayo 2010, Etayo & Sancho 2008, Flakus & Kukwa 2012, Hafellner 1994, Hawksworth & Diederich 1988, Santesson et al. 2004, Zhurbenko 2012) and is here newly reported for Argentina, Australia, Colombia, Japan, Nepal and Papua New Guinea. Specimens examined. ‒ ARGENTINA. ISLA DE LOS ESTADOS: Cabo San Bartolome, N slope of W end of peninsula, 54°54′S 64°42′W, 5.xi.1971, H. Imshaug 53198 & K. Ohlsson (H). AUSTRALIA. : , Mount McKay, 27.i.1967, R. Filson 9654 (H). CANADA. ONTARIO: Kenora District, Fort Severn, 56°00′N 87°28′W, 2.viii.1958, T. Ahti s.n.[b] (H). CHILE. MAGALLANES REGION: near base of Monte Aymond, 52°09′S 69°29′W, 9.x.1971, H. Imshaug 49809 & K. Ohlsson (H). COLOMBIA. BOYACÁ DEPARTMENT: Paramos al NW de Belen, vereda S. Jose de la Montana, alto de las Cruces y alrededores Cabeceras Q. El Toral, elev. 3830 m, 24.xi or ii.1972, A.M. Cleef 1786 (TNS). FALKLAND ISLANDS. East Falklands, Stanley, summit of Mount Kent, elev. 460 m, 14.i.1968, H.A. Imshaug 40468 & R.C. Harris (TNS); West Falklands, Port Howard, summit of Mount Maria, elev. 630 m, 28.i.1968, H.A. Imshaug 41387 & R.C. Harris (TNS). NEPAL. Himalaya, between Sangda and Kagbeini, elev. 4300 m, 12.v.1953, S. Nakao s.n. (TNS); Gandaki & Dhawalagiri Zones, Manang & Mustang Districts, Thorung Phedi (4370 m)‒Thorung La (5110 m)‒above Muktinath (3720 m), elev. 4850 m, 19.viii.1994, M. Mikage et al. 9460415-13a (TNS). JAPAN. HONSHU: Shinano Province (Nagano Prefecture), Yatsugatake Mountains, 35°58′N 138°22′E, 27.vii.1959, M. Togashi & S. Kurokawa s.n.[b] (TNS). NORWAY. SVALBARD: Spitsbergen, Ossian Sarsfjellet, Kongsfjorden, elev. 0‒100 m, 25.vii.1985, H. Kashiwadani 23219 (TNS). HORDALAND: parish Granvin, Mount Nesheimshorgen, elev. 1000 m, ix.1915, J.J. Havaas s.n. (H). PAPUA NEW GUINEA. WESTERN HIGHLAND DISTRICT: between Pindaunde Lakes and summit of Mount Wilhelm, elev. 4150 m, 31.xii.1973, H. Kashiwadani 11154a (TNS). PERU. ANCASH REGION: Huaraz Province, road Huaraz‒Casma, 31 km (road distance) WSW of Huaraz, 9°33′S 77°41′W, elev. 4100 m, 27.ii.1981, R. Santesson & R. Moberg s.n. (H). JUNIN REGION: border of Jauja and Tarma Provinces, km 37 of the road between Jauja and Tarma, 11°35′S 75°35′W, elev. 4140 m, 18.ix.1977, D. Vitt 21661 (H). SWEDEN. ÖLAND: Sandby, Ekelunda, northernmost part of Stora Alvaret, 31.v.2000, R. Skytén 6490 (H). JÄMTLAND: Åre parish, Mount Snasahögarna, elev. 730 m, 14.viii.1975, H. Vänskä 7529 (H).

Sphaerellothecium thamnoliae Zhurb. var. taimyricum Zhurb. NOTE. – This taxon was formerly known from Russia (Zhurbenko 2012) and is here newly reported from Nepal. Specimen examined. ‒ NEPAL. Gandaki & Dhawalagiri Zones, Manang & Mustang Districts, Thorung Phedi (4370 m)‒Thorung La (5110 m)‒above Muktinath (3720 m), elev. 4850 m, 19.viii.1994, M. Mikage et al. 9460415-13b (TNS).

Sphaerellothecium thamnoliae Zhurb. var. thamnoliae NOTES. – The taxon is widely distributed in the northern hemisphere (Zhurbenko 2012) and is here first reported for Australia, Falkland Islands, Japan and Papua New Guinea. In the southern hemisphere the species, without indication of variety, was formerly reported only from Ecuador (Etayo 2017).

370 Specimens examined. ‒ AUSTRALIA. AUSTRALIAN CAPITAL TERRITORY: Brindabella Range, 38 km SW of Canberra, Mount Franklin, 35°29′S 148°47′E, elev. 1640 m, 15.xii.1998, H. Streimann 63462a (H), H. Streimann 63462b (H). NEW SOUTH WALES: Ridge between Mountains Kelly and Scabby, 31 km ENE of Adaminaby, 35°43′S 148°52′E, elev. 1640 m, 22.i.1992, H. Streimann 49174 (H). FALKLAND ISLANDS. East Falklands, Darwin Settlement, Boca House on Brenton Lock, 11.i.1968, H.A. Imshaug 40278 & R.C. Harris (TNS). JAPAN. HONSHU: Rikuchu Province (), summit of , 39°33′N 141°29′E, elev. 1900 m, 27.vii.1967, S. Kurokawa 67149 (TNS), 3.ix.1959, S. Kurokawa 59360 (TNS). NORWAY. SÖR-TRÖNDELAG: Kongsvoll, vii.1933, E.P. Vrang s.n.[b] (H). PAPUA NEW GUINEA. Morobe Province, Mount Sarawaket, 4 km SE of Lake Gwam, 6°21′S 147°09′E, elev. 3300 m, 5.vii.1981, T. Koponen 31788 (H). WESTERN HIGHLAND DISTRICT: between Pindaunde Lakes and summit of Mount Wilhelm, elev. 4200 m, 31.xii.1973, H. Kashiwadani 11009 (TNS), H. Kashiwadani 11154b (TNS), H. Kashiwadani 11159 (TNS), elev. 4400 m, 31.xii.1973, H. Kashiwadani 11242 (TNS). CENTRAL DISTRICT: Mount Albert Edward, elev. 3600 m, 25.x.1975, S. Kurokawa 9400 (TNS). U.S.A. ALASKA: Juneau Icefield, summit of mountain above Taku Glacier, 58°42′N 134°13′W, elev. 1460 m, 19.vii.1962, H.A. Imshaug 28916b (TNS). COLORADO: Grand Co., Berthoud Pass, head of Current Creek, Continental Divide and cirque basin, elev. 3470‒3810 m, 6.vii.1960, S. Shushan et al. 24777a (H).

Sphaeropezia santessonii (Zhurb., Etayo & Diederich) Baloch & Wedin NOTE. – The species was formerly known from Canada, Iceland, Peru and Russia (Diederich et al. 2002, Zhurbenko 2012) and is here newly reported for Japan. Specimens examined. ‒ ICELAND. CENTRAL HIGHLANDS: S of Hofsjökull Glacier, Jökulkriki, elev. 620 m, 22.vii.1972, H. Kristinsson 24811 (H); Jokuldalur Valley, near Tungnafellsjokull Glacier, elev. 1050 m, 8.viii.1967, H. Kristinsson 23093a (TNS). JAPAN. HONSHU: Shinano Province (Nagano Prefecture), Yatsugatake Mountains, 27.vii.1959, M. Togashi & S. Kurokawa s.n.[a] (TNS). PERU. JUNIN REGION: Jauja Province, 30 km (road distance) NNW of Jauja, 11°35′S 75°35′W, elev. 4100 m, 10.ii.1981, R. Santesson et al. s.n. (H).

Stigmidium frigidum (Sacc.) Alstrup & D.Hawksw. NOTE. – The species is widely distributed in both hemispheres (Alstrup & Hawksworth 1990, Etayo 2010, Hafellner 1994, Hafellner & Mayrhofer 2007, Kukwa & Flakus 2009, Santesson et al. 2004, Zhurbenko 2012) and is here newly reported for Japan. Specimens examined. ‒ JAPAN. HONSHU: Shinano Province (Nagano Prefecture), Yatsugatake Mountains, Mount Yoko-dake, Chino-shi, elev. 2700 m, 5.viii.1990, H. Shibuichi 8782b (TNS); Kobushi- dake Mountains, Mount Sanpo, Minami-Saku-gun, elev. 2450 m, 12.vii.1979, H. Shibuichi 5862 (TNS).

Taeniolella sp. (putative asexual morph of Sphaerellothecium thamnoliae) NOTES. – The examined material matches the fungus described by Heuchert et al. (2018: Figs 73‒74) based on collections from Norway and Russia. This is the first report of this putative asexual morph from Japan. Specimens examined. ‒ JAPAN. HONSHU: Shinano Province (Nagano Prefecture), Jogo-sawa, Yatsugatake Mountains, Chino-shi, 35°59′N 138°22′E, elev. 2400 m, 27.viii.1997, H. Shibuichi 9804 (TNS). Musashi Province, Mount Sanpoh, Chichibu-gun, 35°55′N 138°44′E, elev. 2480 m, 2.viii.1974, H. Shibuichi 4967 (TNS). Kai Province (Yamanashi Prefecture), Tengu-one Ridge, Mount Kokushi-dake, Higashi-Yamanashi-gun, 35°52′N 138°40′E, elev. 2500 m, 26.vii.1972, H. Shibuichi 4686 (TNS); -dake, near Kotaro-goya, 35°41′N 138°14′E, elev. 3000 m, 4.viii.1967, M. Togashi s.n. (TNS).

Thamnogalla crombiei (Mudd) D.Hawksw. NOTE. – This species is widely distributed in both hemispheres (Etayo 2010, Etayo & Sancho 2008, Flakus & Kukwa 2012, Hafellner & Mayrhofer 2007, Hawksworth 1980, Kukwa & Flakus 2009, Santesson et al. 2004, Zhurbenko 2012, Zhurbenko & Daniëls 2003) and is here newly reported for Australia, Bhutan, India, Japan, Nepal, North Korea and Papua New Guinea. Specimens examined. ‒ AUSTRALIA. VICTORIA: between peaks 1 & 2, Mount Ida, i.1970, H.A. Morrison s.n. (H); Bogong High Plains, Mount Nelson, elev. 1880 m, 20.i.1966, R. Filson 8128 (TNS). BHUTAN. Lingshi (4100)‒Yale La (4800)‒Shodu (4000), 23.v.1967, H. Hara et al. 19803 (TNS); Laum Thang (3900)‒Singke La (4950)‒Chawa Gassar (4000), 19.v.1967, H. Hara et al.

371 20080 (TNS). CANADA. ALBERTA: SW of Calgary, 0.5 mi S of Savanna Creek, 3.25 mi ESE of the summit of Mount Pasque, elev. 2130 m, 28.ix.1963, C.D. Bird 9085 (H). ONTARIO: Kenora District, Fort Severn, 56°00′N 87°28′W, 2.viii.1958, T. Ahti s.n.[a] (H). INDIA. SIKKIM: Jongri Peak, elev. 4000 m, 21.v.1960, M. Togashi et al. s.n. (TNS). JAPAN. : Ishikari Province, Daisetsu Mountains, -un, 43°39′N 142°54′E, elev. 2100 m, 4.viii.1980, H. Kashiwadani 16253 & Y. Endo (TNS); Daisetsu Mountains, southern slope of Mount Akadake, 43°40′N 142°55′E, elev. 2000 m, 18.ix.1971, S. Kurokawa 71137 (TNS); Daisetsu Mountains, Mount Tomuraushi, 43°31′N 142°51′E, elev. 2000 m, 18.vii.1969, M. Togashi s.n.[b] (TNS); Daisetsu Mountains, 43°N 142°E, 26.vii.1937, Y. Asahina s.n. (TNS). Etchu Province (), Mount Yakushi, 36°28′N 137°33′E, 25.vii.1936, Y. Asahina s.n. (TNS); Mount Kaminotake, 36°25′N 137°31′E, 24.vii.1936, Y. Asahina s.n. (TNS). Iwashiro Province (Fukushima Prefecture), Mount Iide, Yama-gun, 37°51′N 139°42′E, elev. 2000 m, 26.viii.1975, H. Shibuichi 5127b (TNS), H. Shibuichi 5128 (TNS). Shinano Province (Nagano Prefecture), Mount Yatsugatake, Kita-Azumi-gun, 35°58′N 138°22′E, 11.viii.1952, S. Kurokawa 520431 (TNS). NEPAL. [without additional locality data], elev. 3900 m, 28.v.1953, D.D. Awasti 2334 (TNS). NORTH KOREA. Hamkyonnam-do Province, Mount Paekdu, 41°60′N 128°05′E, 26‒27.vii.1933, T. Ueda s.n. (TNS). NORWAY. TRØNDELAG: Oppdal Municipality, Kongsvoll, vii.1933, E.P. Vrang s.n.[a] (H). PAPUA NEW GUINEA. Western Highlands Province, between Pindaunde Lakes and summit of Mount Wilhelm, elev. 4150 m, 31.xii.1973, H. Kashiwadani 11159 (TNS). PERU. JUNIN REGION: Tarma Province, between La Oroya and Junin, S of the turn towards San Pedro, 11°17′S 75°56′W, elev. 4200 m, 16.ii.1981, R. Santesson s.n. & R. Moberg (H). CUZCO REGION: Paucartambo, road Oropesa‒Paucartambo, between Huancarane and Sayllapata, 13°25′S 71°45′W, elev. 3850 m, 26.iii.1981, R. Santesson et al. s.n. (H). SWEDEN. GOTLAND: Visby, 28.vi.1875, S.O. Lindberg s.n. (H). ÖLAND: Stora Alvaret, 4 km ESE of the Resmo church, 2.vii.1957, R. Santesson 11924 (H), 8.viii.1911, R. Sterner s.n. (H). U.S.A. ALASKA: Bering Strait District, Ukinyik Creek Drainage, 68°43‒47′N 165°45′‒166°12′E, elev. 600 m, 4.viii.1960, L. Viereck & A. Bucknell s.n.[b] (H). COLORADO: Grand Co., Berthoud Pass, head of Current Creek, Continental Divide and cirque basin, elev. 3470‒3810 m, 6.vii.1960, S. Shushan 24777a (H). Clear Creek Co., Mount Evans, elev. 4200 m, 30.vii.1961, A. Henssen 13095j (H). WYOMING: Albany Co., summit of Medicine Bow Peak, elev. 3660 m, 4.vii.1956, H.A. Imshaug 18901 (H).

ACKNOWLEDGEMENTS

The research of MZ was carried out within the framework of the research project of the Komarov Botanical Institute of the Russian Academy of Sciences “Biodiversity and spatial structure of fungi and myxomycetes communities in natural and anthropogenic ecosystems” (АААА-А18-118031290108-6) using equipment of its Core Facility Center “Cell and Molecular Technologies in Plant Science”, his visit to TNS was supported by JSPS Invitation Fellowship for Research in Japan (no. S16173). We are indebted to the curator of H herbarium Leena Myllys who assisted in processing relevant Thamnolia specimens for study and loan. Adam Flakus is thanked for his valuable remarks during the preparation of this paper.

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