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Phytotaxa 461 (1): 021–030
https://www.mapress.com/j/pt/
Copyright © 2020 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Christian Printzen: 28 Aug. 2020; published: 30 Sept. 2020
https://doi.org/10.11646/phytotaxa.461.1.3
21
Japewia aliphatica (Lecanoraceae, lichenized Ascomycota), a new acidophilous,
sorediate-blastidiate lichen from Europe
JIŘÍ MALÍČEK1,4*, ZDENĚK PALICE1,5, JAN VONDRÁK1,2,6 & TOR TØNSBERG3,7
1 Institute of Botany of the Czech Academy of Sciences, Zámek 1, CZ-252 43 Průhonice, Czech Republic;
2 Faculty of Biological Sciences, University of South Bohemia, Branišovská 31, CZ-370 05 České Budějovice, Czech Republic
3 Department of Natural History, University Museum, University of Bergen, Allégaten 41, P.O. Box 7800, NO-5020 Bergen, Norway;
4
�
jmalicek@seznam.cz; https://orcid.org/0000-0002-3119-8967
5
�
zdenek.palice@ibot.cas.cz; https://orcid.org/0000-0003-4984-8654
6
�
j.vondrak@seznam.cz; https://orcid.org/0000-0001-7568-6711
7
�
tor.tonsberg@uib.no; https://orcid.org/0000-0001-9081-0274
*Corresponding author:
�
jmalicek@seznam.cz
Abstract
Japewia aliphatica is described as a new species. It is characterized by the usually brown, often areolate, blastidiate to soredi-
ate-blastidiate, rarely fertile thallus containing unknown fatty acid(s). It grows on acidic bark of broad-leaved trees in mainly
in montane forests, and is known from Austria, the Czech Republic, Germany, Norway, Russia (European part of the Cau-
casus), Slovakia and Ukraine. The new species is well characterized by its morphological, chemical and molecular (nrITS,
mtSSU) traits. Systematic placement of Japewia is briefly discussed and its position within Lecanoraceae confirmed.
Keywords: Aliphatic acids, montane forests, sterile lichens
Introduction
Japewia Tønsberg (1990: 205; Lecanoraceae, lichenized Ascomycota) is a genus of lichenized fungi with crustose thalli,
a chlorococcoid photobiont, biatorine apothecia, reduced and weakly gelatinized exciple, frequently branched and
anastomosing paraphyses, asci with a distinct axial body, and simple, thick-walled, multilayered ascospores (Tønsberg
1990, Printzen 1999). According to the newest classification of Lecanoromycetes (Lücking et al. 2016), the genus has
been assigned to the family Ramalinaceae, but molecular data suggests that it rather belongs to Lecanoraceae (Schmull
et al. 2011, Miadlikowska et al. 2014). Two species have so far been described in this genus: Japewia subaurifera
Muhr & Tønsberg (Tønsberg 1990: 206) and Japewia tornoensis (Nyl.) Tønsberg. The third name, Japewia carrollii
(Coppins & P. James) Tønsberg, became the type species for the genus Japewiella (Printzen 1999) and is currently
known as Japewiella tavaresiana (H. Magn.) Printzen.
Japewia tornoensis, the type species of the genus (Tønsberg 1990), is characterized by its esorediate, dark brown
thallus with subglobose, red-brown apothecia. It is widely distributed in temperate and boreal regions of the Northern
Hemisphere as an epiphyte (see Czarnota 2009). In Antarctica it occurs on soil, bryophytes and rocks (Øvstedal &
Lewis Smith 2001). The second species, J. subaurifera, is rarely fertile and further characterized by the golden-brown
color of the soralia due to the production of the pigments secalonic acid X2 and Z, and eumitrin L (Elix & Tønsberg
1999) and by the accessory lobaric acid (Tønsberg 1990). That species is a widespread epiphytic lichen mainly in boreal
forests in the Northern Hemisphere (Czarnota 2009). The latter species has recently been reported from Tasmania
(Kantvilas 2011). However, as that material lacks lichens substances, it may prove to represent an undescribed taxon.
Since the nineties of the 20th century, one of us (TT) was aware of an undescribed taxon similar to Placynthiella
dasaea (Stirt.) Tønsberg. During recent field work in Central and Eastern European woodlands by JM, ZP, and JV, the
same lichen was repeatedly collected. Based on morphological, chemical and molecular characters, we describe this
distinctive taxon as a species new to science.
MALÍČEK ET AL.
22 • Phytotaxa 461 (1) © 2020 Magnolia Press
Material and methods
Sampling, morphology and chemistry
Collected specimens are deposited in BG, PRA, and the personal herbarium of J. Malíček. Microscopic descriptions
are based on hand-cut sections mounted in water. Lichen secondary metabolites were identified using thin
layer chromatography (TLC) in solvents A, B’ and C (Orange et al. 2010). The pictures were acquired by the (1)
stereomicroscope Olympus SZX 12 with the cooled color digital camera Olympus DP 70 (resolution 12.5 Mpx) in
the software QuickPHOTO MICRO 3.0 (Promicra), using an extended depth of field module Deep Focus and (2)
microscope Olympus BX 43 with color digital camera Promicra 3–5CP (resolution 5 Mpx) in the same software.
DNA extraction, PCR amplification and sequencing
The Invisorb Spin Plant Mini Kit (Invitek) and Chelex protocol (Ferencová et al. 2017) were used for DNA extractions.
The fungal ITS rDNA (henceforth ITS) and mitochondrial SSU (mtSSU) were amplified with the following primers:
ITS1F (Gardes & Bruns 1993) and ITS4 (White et al. 1990), mrSSU1, mrSSU2R and mrSSU3R (Zoller et al. 1999).
The algal ITS rDNA was amplified using the specific primer zeleny_F2 (Moya et al. 2018) and the universal primer
ITS4 (White et al. 1990), following methods by Moya et al. (2018). PCR reactions of nrITS and mtSSU were prepared
for a 20 µl final volume, containing 14 µl double-distilled water, 4 µl MyTaq polymerase reaction buffer, 0.2 µl MyTaq
DNA polymerase, 0.4 µl of each of the 25 mM primers, and 1 µl of the sample. Amplifications of both loci consisted
of an initial 1 min denaturation at 95 °C, followed by 35 cycles of 1 min at 95 °C, 1 min at 56 °C, 1 min at 72 °C, and a
final extension of 7 min at 72 °C. The PCR products were visualized on a 0.8% agarose gel and cleaned with ExoSAP-
IT™ PCR Product Cleanup Reagent (ThermoFisher Scientific), according to the manufacturer’s protocols. In total, 6
new ITS and 6 mtSSU sequences were generated (Table 1).
Sequence alignment and phylogenetic analysis
The newly produced sequences were edited in BioEdit 7.2.5 (Hall 1999). The final analyses included the newly
generated sequences, available sequences of the genus Japewia and selected closely related taxa according to Schmull
et al. (2011) and Miadlikowska et al. (2014). They all belong to Lecanoraceae, represent most of main lineages in
this family and both ITS and mtSSU sequences are available in the Genbank database. Additionally, Scoliciosporum
umbrinum was included based on its very close phylogenetic position to Japewia tornoensis in Zhao et al. (2016).
Lecidea nylanderi (Anzi 1860: 75) Th. Fr. (1874: 462) and Lecidea roseotincta Coppins & Tønsberg (1988: 415)
were selected as an outgroup because they form a sister clade to all other included species (see Schmull et al. 2011,
Miadlikowska et al. 2014). The ITS and mtSSU regions were aligned separately using MAFFT 7 (Katoh & Standley
2013) with L-INS-i method (Katoh et al. 2005). Ambiguous positions were excluded from the analysis using Gblocks
0.91b (Castresana 2000), with a less stringent selection, on the Phylogeny.fr server (Dereeper et al. 2008). Gaps were
coded in SeqState by simple coding (Simmons & Ochoterena 2000). The final ITS alignment contained 477 positions
and 24 sequences; the mtSSU alignment had 717 positions and 22 sequences.
Support values on nodes were checked for single-gene trees and no conflict among well-supported branches
(maximum likelihood bootstrap percentages >0.7) was detected. We concatenated the alignments and inferred a
phylogeny using MrBayes 3.2.6 (Huelsenbeck & Ronquist 2001; Ronquist et al. 2012). Results of jModelTest 2.1.10
(Darriba 2012) suggested the general time reversible model including gamma-distributed rates across sites modelled
with four discrete categories and a proportion of invariant sites (GTR+G+I) as the best substitution model for both
regions based on the AIC. Each analysis was performed with two runs, each with four MCMC chains (temperature
0.05). Trees were sampled every 500th generation. Analyses were stopped when the average standard deviation of the
split frequencies between the simultaneous runs was 0.00997. To eliminate trees sampled before reaching apparent
stationarity, the first 25% of entries were discarded as burn-in and the rest were used to compute a majority-rule
consensus tree with Bayesian posterior probabilities for the branches.
A maximum likelihood analysis was performed using RAxML-HPC v. 8.2.12 (Stamatakis 2014) using the
GTR+G+I model on the CIPRES Science Gateway (Miller et al. 2010). Non-parametric bootstrap analysis was
performed with 1000 bootstrap replicates. The maximum likelihood consensus tree is not shown, but bootstrap values
are indicated at branches in the Bayesian tree (Fig. 2). The alignment used in this study is publicly available in Zenodo
as doi: 10.5281/zenodo.3610084.
JAPEWIA ALIPHATICA (LECANORACEAE) Phytotaxa 461 (1) © 2020 Magnolia Press • 23
TABLE 1. GenBank accession numbers and voucher information of specimens used in this study. New sequences are
indicated in bold.
Taxon Source – Specimen ITS mtSSU
Frutidella caesioatra Norway, R. Haugan 10634 (O) MG925971 MG925872
Japewia aliphatica 1Czech Republic, Šumava Mts, J. Vondrák 18554 (PRA) MN547348 MN547347
Japewia aliphatica 2Czech Republic, Šumava Mts, J. Vondrák 18753 (PRA) MN547349 –
Japewia aliphatica 3Czech Republic, Šumava Mts, J. Vondrák 18728 (PRA) MN547350 MN547346
Japewia aliphatica 4Russia, Caucasus Mts, J. Vondrák 15548 (PRA) MN547351 MN547344
Japewia aliphatica 5Czech Republic, Beskydy Mts, J. Malíček 9764 (PRA; holotype) MN547352 MN547343
Japewia aliphatica 6Ukraine, Carpathians, J. Malíček 8238 (hb. J. Malíček) MN547353 MG773680
Japewia aliphatica 7Czech Republic, Hrubý Jeseník Mts, Z. Palice 19222 (PRA) –MN547345
Japewia subaurifera 1U.S.A., T. Spribille s.n. JN009716 –
Japewia subaurifera 2Japan, R. Haugan (O) MH481917 –
Japewia tornoensis 1Canada, C. Printzen (BG) HQ650656 HQ660559
Japewia tornoensis 2Canada, C. Printzen 5638 (BG) EF495163 –
Japewia tornoensis 3Sweden, M. Wedin (UPS) – DQ899301
Japewia tornoensis 4Norway, J. T. Klepsland (O) –MG925874
Lecanora allophana Finland, Kimito, J. Malíček 9491 (hb. J. Malíček) KY548051 KY502416
Lecanora argentata Russia, Caucasus Mts, J. Malíček 9620 (hb. J. Malíček) MK778604 MK778529
Lecanora caesiorubella Australia, H. T. Lumbsch 19974 JN943728 JQ782667
Lecanora cinereofusca U.S.A., North Carolina, J. Lendemer 35007 (NY) KP224471 KP224464
Lecanora intumescens Ukraine, Carpathians, J. Malíček 8203 (hb. J. Malíček) KY548039 KY502443
Lecanora polytropa U.S.A., Alaska, F. Lutzoni et al. (DUKE) HQ650643 DQ986807
Lecidea nylanderi U.S.A., T. Spribille 10020 (hb. Spribille) HQ650667 HQ660572
Lecidea polytrichina Sweden, Z. Palice s.n. (hb. Printzen) HQ650668 HQ660573
Lecidea roseotincta Norway, T. Tønsberg 34577 (hb. Schmull) HQ650670 HQ660575
Lecidella elaeochroma China, Xinjiang KT453749 KT453835
Lecidella stigmatea China, Xinjiang KT453758 KT453852
Protoparmeliopsis muralis Romania, J. S. Hur KP059048 KP059054
Scoliciosporum umbrinum Switzerland KX133008 –
Taxonomy
Japewia aliphatica Malíček, Palice, Tønsberg & Vondrák sp. nov. MycoBank no 832832
(Figs 1 & 2)
Diagnosis: The species is characterized by its brown, often areolate thallus composed of tiny blastidia that frequently develop into pale
soredia. Soralia usually irregular, rarely delimited and rounded, sometimes covering most of the thallus. Apothecia very rare, convex,
reddish-brown. Ascospores ellipsoid to subglobose, with a thick, multilayered wall. Pycnidia unknown. One or several fatty acids are
produced as lichen secondary metabolites.
Type: CZECH REPUBLIC. Beskydy Protected Landscape Area: Frenštát pod Radhoštěm, Kněhyně-Čertův mlýn National Nature Reserve,
old-growth spruce forest on E-facing slope of Mt Kněhyně (1257 m), 1190 m elev., 49°29’48’’N, 18°18’56’’E, on bark of Fagus
sylvatica, 31 August 2016, J.Malíček 9764 & Z.Palice (holotype: PRA; isotype: herb. Malíček).
Description: Thallus of sorediate aggregations of blastidia, covering patches up to 3(–5) cm or even larger areas when
intermixed with other lichens, when well-developed, often organized in areoles about 0.5 mm (up to 1 mm) in diam. and
up to 0.3(–0.6) mm thick; in shady places and also in specimens with delimited soralia occasionally with a brownish
to green-grey, thin (up to 100 μm high), unevenly verrucose to areolate-squamulose basal thallus. Prothallus absent.
Blastidia medium brown, chocolate brown at sun-exposed sites and brown-olive to pale grey-green at shady sites;
MALÍČEK ET AL.
24 • Phytotaxa 461 (1) © 2020 Magnolia Press
(20–)25–35(–45) μm in diam.; wall formed by brown (K+ dark brown) or rarely grey hyphae, composed of ellipsoid
to rounded cells, often with flexuous walls, up to 12 × 4(–7.5) μm. Soralia often missing, variable, mostly uneven in
shape, developing by disintegration of blastidia; becoming confluent and continuously covering the thallus or rarely
delimited and rounded, 0.1–0.2 mm in diam. Soredia farinose, whitish or less frequently pale brown to pale green-grey,
contrasting the brown blastidia, simple, 20–35 μm in diam., or in consoredia up to 80 μm in diam., in a mount delimited
by a colourless or pale brown to grey, more or less compact fungal sheath without projecting hyphae.
FIGURE 1. Morphology of Japewia aliphatica. A, thallus with blastidia developing soredia (PRA-Palice 18750); B, apothecium (PRA-
Palice 25176); C, blastidiate thallus (PRA, holotype); D, soralia developing from blastidiate thallus (PRA-Palice 19029). Scales: 1 mm
(A), 0,2 mm (B−D). Photo by J. Machač (A, B) and J. Malíček (C, D).
Apothecia very rare (known from two samples), lecideoid, convex, glossy, reddish-brown, epruinose, 0.4–0.5 mm
in diam. Proper exciple colourless in inner part, reddish-brown in outer part, K+ brown, HNO3–, composed of radiating
branched hyphae; cells 1.5–3.0 μm in diam. Hypothecium colourless, with oil droplets. Hymenium c. 100 μm high,
with oil droplets; epihymenium reddish-brown, K+ brown, HNO3–, without granules (POL–); paraphyses surrounded
by a gel coat, conglutinated, richly branched and anastomosing, 1.5–2.5 μm thick; tips swollen, up to 4 μm, in K with
more or less distinct, brown caps. Asci broadly clavate, initially apparently 8-spored, but mature asci (1–)3–6-spored;
ascospores broadly ellipsoid to subglobose, simple, colourless, (17–)18–23(–25) × (12–)13–16(–17) μm (n=25), wall
multilayered, 2–4(–6) μm thick. Pycnidia not observed.
Photobiont chlorococcoid, 5–9(–14) μm in diam., morphologically corresponds with the genus Symbiochloris
(Škaloud et al. 2016), but one sequenced specimen (Vondrák 15548; MT022442) was identified as Trebouxia sp., using
the BLASTN search (Altschul et al. 1990).
Chemistry: One or several fatty acids, which may belong to the apinnatic acid complex sensu Obermayer et al.
(2018), detected by TLC. A major fatty acid (A4, B’5, C5) visible in all samples (n=27). A double fatty spot often
visible in the A solvent. Spot reactions: K–, Pd–, C–, KC–; UV–.
Phylogeny: The new species as well as the genus Japewia are strongly supported as a distinct clade in the ITS
and mtSSU phylogeny (Fig. 3). Japewia subaurifera and J. tornoensis are not monophyletic based on our results
which may be caused by incomplete dataset of sequences (see Table 1) or a great variability among populations
within individual continents. Current classification of Lecanoromycetes (Lücking et al. 2016) placed this genus in
JAPEWIA ALIPHATICA (LECANORACEAE) Phytotaxa 461 (1) © 2020 Magnolia Press • 25
the family Ramalinaceae. This contradicts our results as well as the phylogenetic studies by Schmull et al. (2011) and
Miadlikowska et al. (2014). Therefore, the genus Japewia should be placed in Lecanoraceae.
FIGURE 2. Anatomy of Japewia aliphatica. A, apothecial section (PRA-Palice 27112); B, hymenium and ascus (PRA-Palice 27112); C,
exciple (PRA-Palice 27112); D, ascospores in KOH (PRA-Palice 27112); E−F, soredia (PRA-Palice 24323). Scales: 0,1 mm (C), 10 μm
(D−F, H) 20 μm (H). Photo by J. Malíček.
Etymology: The name refers to the diagnostic chemical constituent being an aliphatic acid.
Distribution and ecology: The new species is known from Austria, the Czech Republic, Germany, Norway, Russia,
Slovakia and Ukraine. It has already been published under provisional names Placynthiella aff. dasaea, Japewia sp.
or Japewia dasaea ined. from the Eastern Carpathians in Slovakia and Ukraine (Vondrák et al. 2015, 2018, Malíček et
al. 2018), the Czech Republic (Malíček et al. 2019) and as J. aliphatica from the Caucasus Mts (Urbanavichus et al.
2020).
Japewia aliphatica grows in various types of montane forests, where it prefers well-lit sites. In Norway, it has
been found on Alnus glutinosa in a boreo-nemoral rainforest close to sea-level. In Central Europe, it occurs mainly
above 1100 m elev. Rarely, it has been collected in river valleys at lower elevations (700–900 m). It prefers smooth
MALÍČEK ET AL.
26 • Phytotaxa 461 (1) © 2020 Magnolia Press
acidic bark on trunks of broad-leaved trees. Fagus sylvatica is the most frequent phorophyte, followed by Sorbus
aucuparia and Alnus incana. More rarely it was collected on Acer pseudoplatanus, Alnus glutinosa, Betula sp. and
Picea abies.
FIGURE 3. Phylogeny of the genus Japewia with selected members of the family Lecanoraceae. This is a Bayesian phylogenetic
reconstruction based on nrITS and mtSSU sequences. The new species, Japewia aliphatica, is indicated in bold. Branches with >0.95
Bayesian posterior probability values are indicated by thicker lines. Bayesian posterior probabilities (first value) and maximum likelihood
bootstrap percentages (second value) are indicated.
The most commonly co-occurring species are Parmeliopsis ambigua (Wulfen) Nyl., Buellia griseovirens (Turner
& Borrer ex Sm.) Almb., Lecanora pulicaris (Pers.) Ach., Hypogymnia physodes (L.) Nyl., Lepraria elobata Tønsberg,
Parmelia saxatilis agg. and Platismatia glauca (L.) W.L. Culb. & C.F. Culb. Other associated lichens include e.g.
Biatora efflorescens (Hedl.) Räsänen, Candelariella efflorescens agg., Lepraria rigidula (B. de Lesd.) Tønsberg,
Melanelixia glabratula (Lamy) Sandler & Arup, Toensbergia leucococca (R. Sant.) Bendiksby & Timdal and Violella
fucata (Stirt.) T. Sprib.
JAPEWIA ALIPHATICA (LECANORACEAE) Phytotaxa 461 (1) © 2020 Magnolia Press • 27
Notes: Although the new species is usually sterile, it can be recognized by the brown, often areolate thallus
composed of tiny blastidia and/or sorediate blastidia; and chemically by a fatty acid(s) as the diagnostic substance.
Japewia aliphatica and Placynthiella dasaea are usually morphologically indistinguishable. Chemically, however,
they are easy to hold apart containing aliphatic acids (C–) versus gyrophoric acid (C+ red), respectively. It may also
recall Japewia subaurifera but it is distinct in never having brightly coloured soralia ranging from yellow-, orange-
brown to chocolate-brown colour due to the presence of pigments (Elix & Tønsberg 1999). That species differs further
by the accessory presence of lobaric acid (Tønsberg 1992). The blastidiate thallus of J. aliphatica may also resemble
specimens of Scoliciosporum chlorococcum with a granular to partly sorediate thallus but that species is usually more
green and lacks secondary metabolites.
Additional specimens examined. AUSTRIA. Tyrol: Kitzbüheler Alpen, Kelchsau, Kurzer Grund, near river
Kurzer-Grund-Ache, along the tributary river Trattenbach, 1150–1160 m elev., 47°20’N, 12°09’E, on Alnus incana,
31 August 1996, T.Tønsberg 24336 (BG, FR, GZU, O, TRH, UPS), 24349, 24355 (BG); ibid.: 1110–1140 m elev.,
on a river bank, on Alnus incana, 31 August 1996, T.Tønsberg 24380 (BG); Stubaier Alpen, Stiglreith (Oberperfuss)
– Rosskogelhütte, in a swampy spruce forest, 1600 m elev., 47°14.5’N, 11°11.7’E, on twig of Picea abies, 29 August
1996, T.Tønsberg 24317 (BG).
CZECH REPUBLIC. Western Bohemia: Šumava Mts, WNW of Železná Ruda, Revier Eisenstein, valley of brook
Svarožná (Büchel-Bach), NE of the brook, 100–500 m E of German-Czech border, Picea abies forest, 900–970 m elev.,
49°09’N, 13°11’E, on Fagus sylvatica, 12 October 1999, T.Tønsberg 28148 (BG); Šumava Mts, S of Železná Ruda, just
E of forest road from Debrník to Ferdinand’s valley, 0.5–1 km S of Debrník, along the brook, ±swampy bank of stream
with Alnus incana and A. glutinosa, 720–750 m elev., 49°07.1’N, 13°14.1’E, on Alnus glutinosa and A. incana, 12
October 1999, T.Tønsberg 28186a, 28191a (BG); Železná Ruda, Mt Jezerní hora [1343], top plateau of W subsummit,
remnants of spruce forest just NE of open boulder scree, 1327 m elev., 49°10’22.5”N, 13°10’28.2”E, on bark of Sorbus
aucuparia, 18 September 2019, Z.Palice 27466 & P.Uhlík (PRA); Prášily, SW slope of Mt Ždánidla, 1180 m elev.,
49.0983908°N, 13.3497794°E, on Fagus sylvatica, 14 September 2017, J.Vondrák 18729 (PRA); ibid.: 1130 m elev.,
49°05’48.1”N, 13°21’12.5”E, on Fagus sylvatica, 18 September 2017, J.Vondrák 18554 (PRA); ibid.: 1120 m elev.,
49.0979886°N, 13.3429339°E, 14 September 2017, J.Vondrák 18728 (PRA); Southern Bohemia: Šumava Mts, Nová
Pec, managed beech forest on NE-facing slope of Mt Studničná (1160 m), 1130 m elev., 48°45’24”N, 13°54’10”E, on
Fagus sylvatica, 16 June 2017, J.Malíček 11274 & J.Vondrák 18750 (herb. Malíček, PRA); Nová Pec, Mt Hraničník
– NE slope, remnants of montane mixed forest, 1184 m elev., N48°45’07.2”, E013°54’33.1”, on branch of recently
wind-broken Fagus, 28 October 2014, Z.Palice 18307, 19029 (PRA); ibid.: N slope, old-growth montane mixed forest
with dominating Fagus, N48°45’14”, 1165 m elev., E013°54’16.5”, on bark of Sorbus aucuparia, 12 August 2017,
Z.Palice 24455 (PRA); ibid.: 1170 m elev., 48.75364°N, 13.90472°E, on Fagus sylvatica, 15 June 2017, J.Vondrák
18753, 18754 (PRA); ibid.: managed forest with beech predominant, 1130 m elev., 48°45’24”N, 13°54’10”E, on bark
of Sorbus aucuparia and Fagus sylvatica, 2 August 2017, Z.Palice 24323, 24381 (PRA); Nová Pec, Mt Plechý, NNW
slope, 1 km NW from the top, a belt of young spruce forest in valley of Ježový potok brook, 1285 m elev., 48°46’40.5”N,
13°50’51”E, on bark of Sorbus aucuparia, 21 August 2018, Z.Palice 25176 (PRA); ibid.: 21 August 2019, Z.Palice
27112 (PRA); N Moravia: E Sudetes, Jeseníky Mts, Mt Červená hora [1333], well-lit spruce forest at S-facing slope
just SE–SSE from the top, beneath a red-marked tourist trail, 1230 m elev., 50°08’23.5”N, 17°08’19.6”E, on bark of
Sorbus aucuparia, 18 August 2015, Z.Palice 19222 & J.Vondrák (PRA); Eastern Moravia: Beskydy Mts, Frenštát
pod Radhoštěm, Kněhyně-Čertův mlýn National Nature Reserve, SW-facing slope of Mt Čertův mlýn (1206 m), old-
growth beech-spruce forest, 1150 m elev., ca. 49°29’08’’N, 18°18’10’’E, 28 September 2013, J.Malíček 6110, 6112 &
J.Vondrák 11664 (herb. Malíček, PRA); ibid.: Mt Kněhyně, fragment of old-growth spruce forest with some beech and
sycamore, E-facing slope, 1190 m elev., 49°29’47.8”N, 18°18’56.1”E, on bark of Fagus, 31 August 2016, J.Malíček
& Z.Palice 21798 (PRA); ibid.: managed spruce forest at E facing slope, 1100 m elev., 49°29’47”N, 18°19’07.5”E, on
bark of Fagus, 1 September 2016, J.Malíček & Z.Palice 21838 (PRA).
GERMANY. Niederbayern: Böhmerwald, Regen, Mt Zwercheck [1334], open spruce forest 315 m E-ESE of
the top, just few metres from the state border with CR, near the saddle with Mt Jezerní hora [1343], 1317 m elev.,
49°10’24.8”N, 13°10’14.7”E, on bark of Sorbus aucuparia, 18 September 2019, Z.Palice 27533 & P.Uhlík (PRA)
[distributed also to other herbaria within the multiplicate by W. Obermayer: Dupla Graecensia Lichenum].
NORWAY. Hordaland: Lindås, along the brook W of Eidatræet, 1–10 m elev., 60°38’N, 5°28’E, on Alnus glutinosa,
5 September 1987, T.Tønsberg 10611 (BG).
RUSSIA. Republic of Adigea: Maykop, Guzeripl, protected area Kavkazskiy zapovednik, plateau Lagonaki, 1840
m elev., 44.07777°N, 40.01400°E, on Betula, 12 June 2016, J.Vondrák 15548 (PRA).
MALÍČEK ET AL.
28 • Phytotaxa 461 (1) © 2020 Magnolia Press
SLOVAKIA. Poloniny Mts: Ulič, Nová Sedlica, protected area Stužica, 800 m elev., 49°4’20”N, 22°32’6”E, on
Fagus sylvatica, 5 June 2014, J.Šoun & J.Vondrák 12187 (PRA), ibid.: 1150 m elev., 49°5’24”N, 22°32’57’’E, 7 June
2014, J.Šoun & J.Vondrák 12186 (PRA).
UKRAINE. East Carpathians: Khust, Velyka Uhol’ka, old-growth beech forest on a ridge 1.5 km SW of Mt
Manchul (1500 m), 6.5 km E of Zabrid, 1200 m elev., 48°17’52”N 23°39’59”E, on Fagus sylvatica, 17 May 2015,
J.Malíček 8238, Z.Palice 19774 & J.Vondrák 14134 (herb. Malíček, PRA); Shirokyi luh, old-growth beech forest
on a ridge 1.5 km E–ESE of Mt Menchul [1501], 5.5 km NW–NNW of Prihid’ village, 1250 m elev., 48°18’40”N,
23°41’59”E, on bark of old Fagus, 27 May 2019, J.Malíček, Z.Palice 26915 & J.Vondrák 21652 (herb. Malíček, PRA);
Gorgany Mts, Nadvirna, Bystrytsia, c. 5 km SE of village, 1050 m elev., 48.43071°N, 24.32381°E, on Fagus sylvatica,
23 August 2019, J.Vondrák 21903 (PRA).
Acknowledgements
Pavel Škaloud and Ondřej Peksa checked the identity of the photobiont. Ivana Černajová kindly provided a sequence
of the algal partner. Jiří Machač contributed with two photos of the new species. Both reviewers helped to improved
the manuscript. JM, ZP and JV have been supported by the long-term research development project RVO 67985939.
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