Lecanora stanislai Guzow-Krzemińska, Łubek, Malíček & Kukwa, 2017

Guzow-Krzemińska, Beata, Łubek, Anna, Malíček, Jiří, Tønsberg, Tor, Oset, Magdalena & Kukwa, Martin, 2017, Lecanora stanislai, a new, sterile, usnic acid containing lichen species from Eurasia and North America, Phytotaxa 329 (3), pp. 201-211 : 205-208

Lecanora stanislai Guzow-Krzemińska, Łubek, Malíček & Kukwa sp. nov. MycoBank no 823496

Diagnosis: Characterized by a very thin and mostly endosubstratal in non-sorediate parts thallus composed of a more or less continuous layer of soredia without distinctly delimited soralia, the production of usnic acid and zeorin, by growing usually on smooth bark of trees in forests and by differences in ITS sequences.

Type:— POLAND. Równina Bielska, Białowieża Forest, Białowieża National Park, forest section No. 256, Querco-Piceetum, October 2014, on Carpinus betulus, M. Kukwa 13009 & A. Łubek (holotype UGDA!, isotype KTC!).

( Fig. 2A & B )

Thallus sorediate, indeterminate, small (c. 1 cm in diam.) or covering large parts of the substrate, very thin and in most cases completely endosubstratal in non-sorediate parts, rarely with small (c. 0.05 mm) convex, green or blue green areoles soon bursting to form soredia ; prothallus often dark brown to almost black, sometimes indistinct; soralia yellow-green to grey-green, sometimes with distinct bluish tinge due to a pigment in external soredia, developing from endosubstratal parts of thalli, effuse and forming more or less continuous layer over thalli, or rarely forming from areoles and then more or less discrete at first, but soon confluent; soredia simple, in older specimens covered with hyaline crystals, round, up to 35 μm in diam., rarely in irregular consoredia, up to 50 μm in diam.; soredial wall well developed, but not complete, without distinct projecting hyphae; apothecia and conidiomata unknown; photobiont trebouxioid, cells up to 10 μm in diam.

Chemistry: Usnic acid and zeorin, often with traces or minor amounts of additional terpenoids by TLC (all samples analyzed); usnic acid by LC-MS (two specimens analyzed: Kukwa 13010, 13011).

Etymology: The new species is dedicated to the eminent Polish lichenologist, Professor Stanisław (in Latin: Stanislaus) Cieśliński, who intensively studied lichens in the Białowieża National Park as well as in Northeastern Poland and greatly contributed to lichenology in the country.

Habitat and distribution: The new species grows in major cases on smooth bark of deciduous trees ( Carpinus betulus , Alnus spp. , Betula pendula , Quercus robur , Tilia cordata and others), Corylus avellana and Frangula alnus . Some specimens have been collected on coniferous trees ( Abies sp., Picea abies , Pinus sp. ) and two on smooth dry wood. It was usually found in shaded places in forests. In Poland, from where many records are known, the species is found usually in the forests protected in nature reserves and national parks with old-growth stands.

So far L. stanislai has been found in Eurasia ( Czech Republic, Norway, Poland, Russia, Ukraine) and western North America ( Canada, USA).

Notes: The new species was at first included in the variation range of L. compallens , however the morphological examination and the phylogenetic analyses showed that they are different. The main discriminating characters are the development of soralia and the thallus in non-sorediate parts. The esorediate thallus of L. stanislai is usually endosubstratal and not evident (the colour often resembling tree bark), but sometimes a few, small and indistinct areoles soon dissolving into soredia or soralia are developed directly from the endosubstratal thallus. In comparison the thallus of L. compallens is yellow-grey to whitish-grey, and often distinct in non-sorediate parts developing episubstratal areoles ( Fig. 2C ). The soralia in the latter species burst from the areoles and are at first well delimited ( Fig. 2C ), but soon they fuse to form a sorediate layer usually covering the whole thallus except for the marginal zone ( Fig. 2C & D ) (Herk & Aptroot 1999).

The new species can also be confused with L. expallens . The latter however, has externally yellowish to pale brownish soredia in sunny places and contains thiophanic acid (and other xanthones) in addition to usnic acid and zeorin ( Tønsberg 1992; Zduńczyk & Kukwa 2014a). However, the concentration of xanthones in L. expallens can frequently be very low and hardly visible on TLC plates; this is frequent in samples growing in shade, e.g. in forests. Therefore, it is necessary to extract substances from quite big thallus fragments and later carefully examine the TLC plates in UV before the application of sulphuric acid, when xanthones are best visible as dark red or orange spots. Although, the presence of xanthones in some collections from shady sites was revealed only by the more sensitive LC-MS method.

Lecanora thysanophora also contains usnic acid and zeorin, but this species has usually fibrous prothallus (sometimes only very locally developed) and additionally produces atranorin and unknown terpenoids called ‘thysanophora unknowns’ ( Harris et al. 2001).Another lichen with a similar chemistry and morphology, L. flavoleprosa , differs in the frequent production of an additional xanthone (so called ‘flavoleprosa unknown’), the areolate-sorediate thallus and in having a boreal-alpine distribution ( Tønsberg 1992; Czarnota et al. 2009).

The new species contains usnic acid like Lecanora strobilina ( Sprengel 1827: 300) Kieffer (1895: 74) and L. symmicta , but those two species, although with granular thalli in some specimens, are never truly sorediate and always fertile ( Śliwa & Wetmore 2000; Printzen 2001; LaGreca & Lumbsch 2013).

Several other species with leprarioid thalli also contain usnic acid, which in some of them is accompanied by zeorin. Lepraria ecorticata lacks zeorin (see above more discussion; Orange et al. 2017) and L. leuckertiana ( Zedda 2000: 108) L. Saag in Saag et al. (2009: 41) has obscurely and minutely lobed, cottony thallus with thick medulla ( Zedda 2000; Saag et al. 2009). Leprocaulon coriense ( Hue 1924: 386) Lendemer & Hodkinson (2013: 1011) and Septotrapelia usnica ( Sipman 2003: 179) Kalb & Bungartz in Bungartz et al. (2013: 11) have lobate thalli ( Sipman 2003; Saag et al. 2009; Bungartz et al. 2013; Lendemer & Hodkinson 2013). Leprocaulon calcicola Earl. -Benn. et al. in Orange et al. (2017: 184) has larger granules (60–120 μm in diam.; consoredia in Lecanora stanislai are up to 50 μm in diam.) and grows on mortar and overgrowing mosses ( Orange et al. 2017), whereas L. knudsenii Lendemer & Hodkinson (2013: 1011) grows on non-calcareous rocks in sheltered habitats and the thalli are leprose, not stratified and consist of aggregated granules ( Lendemer & Hodkinson 2013).

In the Polish localities in the Białowieża National Park the new species was sometimes accompanied by an morphologically similar, sterile, sorediate lichen, which also contained usnic acid. This species, however, lacked zeorin (but traces of other terpenoids were present), its thalli consisted only of soredia-like granules and no prothallus was detected. Similar specimens were found also in material collected in Canada (TØnsberg 12697 & 12692, BG). We have generated ITS sequences of two specimens with this characteristics (Kukwa 15649 & 15656, UGDA), however, they are not similar to any other sequences deposited in the GenBank ; such specimens are presently studied for another paper.

Additional specimens of L. stanislai examined. CANADA. British Columbia, Clearwater Valley, N of Clearwater along Stillwater Trail, Blackwater Creek, 51°57.0’N, 120°03.3’W, elev. 610–620 m, in a marsh, on Alnus incana , 9 May 2001, T. TØnsberg 29235, T. Goward ( BG); N of Vancouver, N of Brackendale, Paradise Valley, along the road W of Cheakamus River, 4.2 km from junction Hwy 99, 49°48’N, 123°09’W, elev. 50 m, on Alnus sp. , 17 September, 1989, T. TØnsberg 12924 ( BG). CZECH REPUBLIC. SILESIA, distr. Frýdek-Místek, Beskydy Protected Landscape Area, Horní Lomná, Mionší National Reserve, E facing slope of Mionší vrch, 49°32’21”N, 18°39’31”E, elev. 860 m, old-growth beech forest, on Fagus sylvatica , 9 September 2015, J. Malíček 8931, J. Kocourková, J. Vondrák (herb. Malíček); CENTRAL BOHEMIA, distr. Rakovník, Křivoklátsko Protected Landscape Area, Roztoky u Křivoklátu, Na Babě Nature Reserve, 50°01’N, 13°52’E, elev. 240 m, near the river, at the base of Carpinus betulus , 19 August 2011, J. Malíček 3675 (herb. Malíček). NORWAY. Aust-Agder, Valle, along Stavdalsbekken, UTM 32V, ML 1662, elev. 460 m, on Picea abies , 27 July 1987, T. TØnsberg 10186 ( BG); Buskerud, Krødsherad, NE of Ringneselva, UTM 32V, NM 3381, elev. 330 m, on Sorbus aucuparia , 15 June 1982, T. TØnsberg 6939 ( BG). POLAND. Pojezierze Iławskie, S of Szadowo, Liwa river valley, 53°46’24”N, 19°03’14”E, horn-beam forest, on Ulmus sp. , 12 April 2004, M. Kukwa 3054 ( UGDA); W of Mątki, forest section No. 207, 53°50’45”N, 18°58’37”E, on Carpinus betulus , 19 August 1996, M. Kukwa s.n. ( UGDA); W of Mątki, forest section No. 209, 53°50’44”N, 18°58’10”E, on Quercus sp. , 20 August 1996, M. Kukwa s.n. ( UGDA); Pojezierze Kaszubskie, Jar rzeki Raduni nature reserve, forest section no. 96, 54°18’N, 18°18’E, on Carpinus betulus , 10 August 1996, D. Królak s.n. ( UGDA); ibid., on Alnus incana , 19 September 1983, W. Fałtynowicz s.n. ( UGDA); Pojezierze Wschodniosuwalskie, Suwalski Landscape Park, S of Smolniki, (54°17’N / 22°52’E), SW part of lake Jaczno, 54°16’24”N, 22°52’02”E, humid forest, on bark of stump, 13 July 2000, K. Jando, M. Kukwa s.n. ( UGDA); Wybrzeże Słowińskie, Białogóra nature reserve, 54°49’43”N, 17°58’42”E, Empetro nigri-Pinetum, on Populus tremula , 23 June 2010, M. Kukwa 8163 ( UGDA); Równina Bielska, Białowieża Forest, S of Czerlonka, forest section no. 469C, 52°41’16”N, 23°43’02”E, on Carpinus betulus , 28 August 2016, M. Kukwa 17891, A. Łubek ( UGDA); W of Budy, forest section no. 335D, 52°44’03”N, 23°42’59”E, corticolous, on Carpinus betulus , 27 August 2016, M. Kukwa 17778, 17801, 17806, 17881, A. Łubek ( UGDA); Białowieża National Park, forest section No. 256, corticolous or lignicolous (1 specimen), 2014, M. Kukwa 12669, 12734, 12763, 12786, 12838, 12867, 12991, 13010, 13011, 13012, 13217, 13229, 13252, 13284, 13323, 13347, 13348, 13355, 13371, 13395, 13396, 13404, 13706, 13976, 14062, 14063, 14090, 14128, 14196, 14219, 14226, 14228, 14258, 14338, 14343, 14348, 14382, A. Łubek ( KTC, UGDA, herb. Malíček); ibid., corticolous or lignicolous (1 specimen), 2015, M. Kukwa 15650, 15652, 15654, 15657, 15662, 17124a, 17241, 17320, 17424, 17447, 17670, A. Łubek ( KTC, UGDA); Wysoczyzna Elbląska, Kadyny village, 54°17’44”N, 19°29’22”E, forest above the palace, on Fraxinus excelsior , 21 June 1956, T. Sulma s.n. ( UGDA); Wysoczyzna Żarnowiecka, Pużyckie Łęgi nature reserve, 54°38’N 17°51’E, on Carpinus betulus and Frangula alnus , 12 August 2015, M. Kukwa 17043, 17051 ( UGDA). RUSSIA. Caucasus Mts, Caucasian Biosphere Reserve, siliceous ridge 1 km SE of village Guzeripl, 43°59’12”N, 40°08’29”E, elev. 940 m, primeval mixed forest, on young Fagus orientalis , 7 June 2016, J. Malíček 10367 (herb. Malíček); Maykop, Guzeripl, protected area Kavkazskiy zapovednik, elev. 700 m, 43.98889°N, 40.12507°E, on bark of Alnus glutinosa , 8 June 2016, J. Vondrák 14920 ( PRA). UKRAINE. EAST CARPATHIANS, Zakarpattia Oblast Region, Khust Velyka Uhol’ka, valley of Velyka Uhol’ka river, c. 0.7 km NNE of the village, 48°15’02”N, 23°41’47”E, elev. 500 m, old-growth beech forest, on Fagus sylvatica , 13 May 2015, J. Malíček 8168 et al. (herb. Malíček). USA. ALASKA. Skagway, along and uphill from Klondyke Hwy, 59°32.68’N, 135°12.44’W, elev. 400 m, on Salix sp. , 20 Augut 2003, T. TØnsberg 32841 ( BG); ARKANSAS, Stone Co., Ozark Mts, NW of Mountain View, Blanchard springs Recreation Area, along North Sylamore Creek, 35°58’N, 92°10’W, elev. 115 m, on Pinus sp. , 2 October 1993, T. TØnsberg 19254 ( BG); CALIFORNIA, Del Norte Co., along U.S. Route 199 just S of the Oregon / California border, 41°59.6’N, 123°43.1’W, elev. 500–600 m, on Alnus rubra , 4 May 1991, T. TØnsberg 14701 ( BG); WASHINGTON, Chelan Co., along Old Blewett road (FR 7320) N of old Blewett Pass, 47°21.7’N, 120°39.6’W, elev. 1100 m, on Abies sp. and Acer glabratum , 24 September 1999, T. TØnsberg 28060, 28076 ( BG).

Selected specimens of L. compallens examined. CZECH REPUBLIC. SOUTH MORAVIA, Jihomoravský kraj, S of Lanžhot, Cahnov-Soutok National Nature Reserve, the protected area Cahnov , 48°39’22”N, 16°56’27”E, elev. 152 m, on fallen Quercus robur , 2 Apr. 2014, M. Kukwa 12477 ( UGDA) . ITALY. Sicily, Nebrodi Mts, Nicosia Riserva Naturale Orientata, Sambuchetti - Campanito: margin of pine forest in E part of protected area, 37°50’42’’N, 14°23’21’’E, elev. 1420 m, on bark of Pinus nigra , 24 September 2014, J. Malíček 7517 ( UGDA, herb. Malíček, fertile collections!) . NETHERLANDS. Prov. Drenthe, Drouwenerveen (between Gieten and Standskanaal ), 253,9/554,1 (6°50’E, 53°00’N), square 12-57, well-lit wayside Quercus robur trees, at exposed side of the trunk, 5 March 1998, C.M. van Herk 2345 (L; holotype) ; ibid., 8 August 2014, J. Malíček 6948 & L. Syrovátková (herb. Malíček; topotype) ; Roden , on Quercus sp. , 18 March 1998, C.M. van Herk s.n. ( UGDA) . POLAND. Równina Bielska, Białowieski National Park, forest section no 256, Carici elongatae-Alnetum, on bark of twigs of snag, Aug. 2014, M. Kukwa 13887, A. Łubek ( UGDA, KTC) .

Figures

FIGURE 2. Morphology of Lecanora stanislai (A & B) and L. compallens (C & D). A, central part of thallus (holotype); B, edge of thallus with prothallus (Kukwa 13404); C edge of thallus with episubstratal areoles and soralia (Malíček 7517); delimited and fusing soralia in the center of the thallus (Malíček 7517). Scale: A–D—0.5 mm.