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Arthonia apatetica (Arthoniales, Arthoniaceae) new to northeastern Asia
Korean J. Microbiol. 2023;59(2):77-90
Published online June 30, 2023
© 2023 The Microbiological Society of Korea.

Beeyoung Gun Lee1* and Jae-Seoun Hur2

1Forest Bioresources Conservation Division, Baekdudaegan National Arboretum, Bonghwa 36209, Republic of Korea
2Korean Lichen Research Institute, Sunchon National University, Suncheon 57922, Republic of Korea
Correspondence to: *E-mail:; Tel.: +82-54-679-0679; Fax: +82-54-679-0970
Received April 19, 2023; Revised June 7, 2023; Accepted June 20, 2023.
Arthonia apatetica (A. Massal.) Th. Fr. is described as a lichen- forming fungus from South Korea, new to northeastern Asia. The minute and negligible microorganism is characterized by small, black apothecia with 1-septate ascospores, absence of crystals, pale brown hypothecium, periclinally arranged paraphysoids at epihymenium, and chlorococcoid photobiont among corticolous species. Molecular analyses employing the mitochondrial small subunit (mtSSU), the nuclear large subunit ribosomal RNA (LSU), and the RNA polymerase subunit II (RPB2) sequences strongly support the collected Korean specimens corresponding to A. apatetica in the Bryostigma clade of Arthonia. The newly produced genes are the first data of A. apatetica in Asia. An updated key is provided to assist in the identification of all 54 taxa of Arthonia in northeastern Asia.
Keywords : Arthoniaceae, biodiversity, molecular biology, phylogeny, taxonomy

A lichen is a symbiotic organism by association of a lichen-forming fungus and an alga and/or a cyanobacterium (Hawksworth, 1988). And the organism is regarded as a consortium of above members and lichenicolous fungi, endolichenic fungi, and other bacteria in a wide sense. Lichen taxonomy is based on the lichen-forming fungi and lichens are classified in the kingdom fungi. The number of lichen species is approximately 19,000 recorded (Lücking et al., 2017). The family Arthoniaceae is the second largest group of the lichen-forming fungi, and this study focuses on a species of the main genus Arthonia in the family.

The genus Bryostigma (syn. Arthonia) was first introduced in 1979 just with a single species B. leucodontis Poelt & Döbbeler (Poelt and Döbbeler, 1979). The authors characterized the genus by the hyphae reaction to red or blue in iodine, the undifferentiated excipulum, the Arthonia-type asci with or without amyloid ring structures, the small ascomata, and importantly the substrate preference to bryophytes (Poelt and Döbbeler, 1979) which is assumingly the key characteristic to name the genus.

However, Bryostigma have been challenged for natural classification. In specific, the genus with the type species B. leucodontis was reclassified into Arthonia s. lat. mainly by its similarity with the specimens of A. apatetica (Coppins, 1989). Both species share the important characteristics of paraphysoids, i.e., brown-walled paraphysoidal tips and periclinally extended paraphysoids at epihymenium, which are the taxonomic key distinguishing from a closely related species, A. patellulata Fée, representing non- or faintly pigmented paraphysoidal tips and erect paraphysoids at epihymenium (Coppins, 1989). The type species was moved into Arthonia as A. muscigena Th. Fr. (Santesson, 1993), and was classified into the Bryostigma clade of Arthonia based on molecular phylogeny (Frisch et al., 2014). The type species of Bryostigma is named B. muscigenum (Th. Fr.) Frisch & G. Thor at present (Frisch et al., 2014). Since Frisch et al. (2014), several new species were discovered in the Bryostigma clade, such as A. parietinaria Hafellner & Fleischhacker, A. lobariellae Etayo, A. toensbergii Holien & Frisch, A. dokdoensis S.Y. Kondr., Lőkös, B.G. Lee, J.J. Woo & Hur, and B. huriellae S.Y. Kondr. & Hur (Fleischhacker et al., 2016; Etayo, 2017; Frisch and Holien, 2018; Kondratyuk et al., 2019, 2020a). The old species A. glebosa is classified in the Bryostigma clade as well (Halıcı and Kahraman, 2021). However, the taxonomy of B. apotheciorum (A. Massal.) S. Y. Kondr. et J.-S. Hur (syn. A. apotheciorum [A. Massal.] Almq.) may not be convincing (Kondratyuk et al., 2020b) as the species was classified far from the Bryostigma clade in molecular phylogeny (Frisch et al., 2014; Lee and Hur, 2016). Overall 18 species are recorded in the Bryostigma clade at present as other possible species, A. fuscopurpurea (Tul.) R. Sant., A. patellulata, and lichenicolous species, are not yet included due mainly to lack of molecular results.

This study describes a lichen-forming fungus in the Bryostigma clade of Arthonia, new to northeastern Asia. Field surveys for the lichen biodiversity in the southern region of the Mts. Baekdudaegan and the forested wetlands of Korea were carried out during the spring and summer of 2021, and overall 25 specimens of Arthonia were collected from the barks of the broad-leaved deciduous trees (Fig. 1). The specimens were comprehensively analyzed in ecology, morphology, chemistry and molecular phylogeny and corresponded to A. apatetica. Arthonia apatetica was previously reported from Europe (Wirth, 1995), North America (Brodo, 1995), the northwest Siberia (Sedelnikova and Taran, 2000), and Mongolia (Hauck and Javkhlan, 2006). We report the specimens as a new record to northeastern Asia. The specimens are deposited in the herbarium of the Baekdudaegan National Arboretum (KBA), South Korea.

Fig. 1. Collection sites for the newly recorded Arthonia apatetica (black star marks).
The brackets indicate substrates, i.e., Ac, Acer tataricum subsp. ginnala; Al, Alnus incana subsp. hirsuta; Ca, Carpinus tschonoskii; Ce, Celtis sinensis; Di, Diospyros kaki.

Materials and Methods

Morphological and chemical analyses

Hand-cut sections were prepared with a razor blade under a stereomicroscope (Olympus optical SZ51; Olympus), examined under a compound microscope (Nikon Eclipse E400; Nikon) and imaged using a software program (NIS-Elements D; Nikon) and a DS-Fi3 camera (Nikon) mounted on a Nikon Eclipse Ni-U microscope (Nikon). The ascospores were investigated at 1000× magnification in water. The length and width of the ascospores were measured and the range of spore sizes was shown with average, standard deviation, and number of measured spores. Thin-layer chromatography (TLC) was performed using the solvent system C according to standard methods (Orange et al., 2001).

Molecular phylogeny

Isolation, DNA extraction, amplification, and sequencing: hand-cut sections of ascomata or thallus from all collected specimens were prepared for DNA isolation and DNA was extracted with a NucleoSpin Plant II Kit in line with the manufacturer’s instructions (Macherey-Nagel). PCR amplification for the mitochondrial small subunit (mtSSU), the nuclear large subunit ribosomal RNA (LSU), and the RNA polymerase subunit II (RPB2) genes was achieved using Bioneer’s AccuPower PCR Premix (Bioneer) in 20-μl tubes and primers mrSSU1 and mrSSU3R (Zoller et al., 1999), and LR0R and LR5 (Rehner and Samuels, 1994), and fRPB2-7cF and fRPB2-11aR (Liu et al., 1999), respectively. The PCR thermal cycling parameters used were 95°C (15 sec), followed by 35 cycles of 95°C (45 sec), 54°C (45 sec), and 72°C (1 min), and a final extension at 72°C (7 min) based on Ekman (2001). DNA sequences were generated by the genomic research company GenoTech.

Phylogenetic analyses: all mtSSU, LSU and RPB2 sequences were aligned and edited manually using ClustalW in Bioedit V7.2.6.1 (Hall, 1999). All missing and ambiguously aligned data and phylogenetically uninformative positions were removed and phylogenetically informative regions were finally analyzed in MEGA X (Stecher et al., 2020). The final alignment comprised 1347 (mtSSU), 2260 (LSU), and 2065 (RPB2) columns. In them, variable regions were 94 (mtSSU), 353 (LSU), and 32 (RPB2). Finally, the phylogenetically informative regions were 637 (mtSSU), 1113 (LSU), and 472 (RPB2). A concatenation was carried out for combining all mtSSU, LSU and RPB2 gene loci. They were manually combined for the informative regions. Two problematic sequences were removed when conflicting results occurred in the internal branches with the bootstrap values ≥ 70% and the posterior probabilities ≥ 95% in the concatenated tree. Phylogenetic trees with bootstrap values were obtained in RAxML GUI 2.0 beta (Edler et al., 2021) using the maximum likelihood method with a rapid bootstrap with 1,000 bootstrap replications and GTR GAMMA (TVM + G4 for both mtSSU and the concatenation of mtSSU, LSU and RPB2) for the substitution matrix. The posterior probabilities were obtained in BEAST 2.6.4 (Bouckaert et al., 2019) using the GTR 123421 (mtSSU) and the GTR 123121 (concatenation) models, as the appropriate model of nucleotide substitution produced by the Bayesian model averaging methods with bModelTest (Bouckaert and Drummond, 2017), empirical base frequencies, gamma for the site heterogeneity model, four categories for gamma, and a 10,000,000 Markov chain Monte Carlo chain length with a 10,000-echo state screening and 1,000 log parameters. Then, a consensus tree was constructed in TreeAnnotator 2.6.4 (Bouckaert et al., 2019) with the first 25% discard as a burn-in, no posterior probability limit, a maximum clade credibility tree for the target tree type, and median node heights. All trees were displayed in FigTree 1.4.2 (Rambaut, 2014) and edited in Microsoft Paint. Most sequences employed for the analyses are based on Frisch et al. (2014). Overall analyses in the materials and methods were accomplished based on Lee and Hur (2020).



Arthonia apatetica (Massal.) Th. Fr., Botaniska Notiser 56 (1866) (Fig. 2).

Fig. 2. Arthonia apatetica (KBA-L-0001570 for [A, B, D, & F–K]; KBA-L-0002050 for [C and D]) in morphology.
(A–E) Habitus and apothecia. (F) Adnate apothecia with chlorococcoid algae. (G) Interascal hyphae anastomosing with coarsely periclinally arranged paraphysoids at epihymenium. Paraphysoidal tips not pigmented but swollen a little. (H) Hymenium representing blue in iodine. (I) Asci clavate to widely clavate with 8-spores per ascus. (J–L) Ascospores consistently 1-septate with constriction at septum. Bars: (A), 2 mm; (B and C), 500 μm; (D and E), 200 μm; (F), 50 μm; (G–I), 20 μm; (J–L), 10 μm.

Thallus corticolous, crustose, smooth, olive-green, not developed or indistinct as getting old and mostly being disappeared; photobiont chlorococcoid, cells globose to subglobose, 5–20 μm diam. Prothallus absent. Apothecia simple, generally rounded, somewhat convex, dark brown to black, more brownish in water, not pruinose, 0.1–0.4 mm diam.; epihymenium brown to slightly grayish brown, 10–15 μm high; hymenium hyaline, 30–40 μm high; hypothecium pale brown, 30–40 μm high. Crystals or oil droplets not present. Interascal hyphae paraphysoidal, anastomosing, irregular but rather periclinally arranged at epihymenium, tips slightly swollen, not pigmented, branched, 1.5–2 μm thick, brown epihymenium not clearly pigmented by paraphysoidal tips. Asci 8-spored, clavate to narrowly clavate, 24–35 × 14–20 μm (n = 10). Ascospores constantly 1-septate, constricted at septum, upper cell wider and the lower longer, occasionally tales present on one or both ends, 11.5–17 × 4–6 μm (mean = 13.6 × 5.2 μm; SD = 1.1(L), 0.5(W); L/W ratio 2.0–3.4, ratio mean = 2.6, ratio SD = 0.3; n = 50), without epispore.

Chemistry: Epihymenium K+ greenish gray to blackish, hymenium I+ red-brown or blue, UV–. No lichen substance was detected by TLC.

Ecology and distribution: The species occurs on barks of diverse deciduous trees (e.g., Acer, Alnus, Carpinus, Celtis, or Diospyros). This species is distributed in Europe, North America, the northwest Russia, Mongolia, and Korea.

Remarks: Arthonia apatetica is characterized by small, black apothecia with 1-septate ascospores, absence of crystals, pale brown hypothecium, periclinally arranged paraphysoids at epihymenium, and chlorococcoid photobiont among corticolous species. Arthonia patellulata differs from A. apatetica by erect paraphysoids at epihymenium and darker hypothecium. Although B. muscigenum share the characteristic, i.e., the periclinally arranged paraphysoids at epihymenium, the former differs from the latter by narrower ascospores (2.5–4 μm) and darker hypothecium (Smith et al., 2009).

The Korean specimens are similar to those of Europe and North America with a little difference (Table 1). The Korean specimens are supposed to be classified into A. apatetica in morphology and molecular phylogeny and we regard that the difference in morphology is an infraspecific variation.

Infraspecific comparison of Arthonia apatetica among different continents
Specimens Korea Europe North America
Hymenium height (μm) 30–40 40–60 40–50
Hymenium color Hyaline Hyaline to pale red Pale brown
Hymenium in iodine Red-brown Red Red
Hypothecium color Pale brown Hyaline to straw-colored Pale brown
Paraphysoidal tip pigmentation Not pigmented Distinct dark brown
Paraphysoidal tip thickness (μm) 1.5–2 3–4 c. 1 (but variable)
Ascospores (μm) 11.5–17 × 4–6 12–15 × 4.5–5 10–13 × 4–5
Reference Present study Smith et al. (2009) Nash et al. (2007)

The morphological characteristics are referenced from the previous literatures. All information on A. apatetica of Korea is produced from collected specimens in this study.

Specimen examined: SOUTH KOREA, South Jeolla Province, Jinan, Jucheon-myeon, a forested wetland, 35°59’45.4”N, 127°21’58.2”E, 680 m alt., on bark of Acer tataricum subsp. ginnala, 10 May 2021, B.G. Lee & D.Y. Kim 2021-000098, with Lecanora sp., Phaeophyscia rubropulchra (KBA-L-0001570; GenBank ON116972 for mtSSU, ON115808 for LSU, and ON260924 for RPB2); SOUTH KOREA, South Jeolla Province, Gwangju, Gwangsan-gu, Dodeuk-dong, a forested wetland (Jang Valley), 35°08’55.1”N, 126°42’42.9”E, 60 m alt., on bark of Diospyros kaki, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000165, with Graphis scripta, Phaeophyscia rubropulchra, Porina hirsuta (KBA-L-0001637; ON116973 for mtSSU, ON115809 for LSU); same locality, on bark of Alnus incana subsp. hirsuta, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000182, with Hypotrachyna pseudosinuosa, Traponora varians (KBA-L-0001654); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000194, with Hypotrachyna pseudosinuosa, Phaeophyscia rubropulchra, Porina hirsuta (KBA-L-0001666); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000197, with Hypotrachyna pseudosinuosa, Phaeophyscia rubropulchra (KBA-L-0001669); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000199, with Hypotrachyna pseudosinuosa, Phaeophyscia rubropulchra (KBA-L-0001671); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000200, with Hypotrachyna pseudosinuosa, Porina hirsuta (KBA-L-0001672); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000201, with Phaeophyscia rubropulchra (KBA-L-0001673); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000202, with Phaeophyscia rubropulchra (KBA-L-0001674); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000203, with Phaeophyscia rubropulchra, Porina hirsuta (KBA-L-0001675); same locality, on bark of Celtis sinensis, 11 May 2021, B.G. Lee & D.Y. Kim 2021-000205, with Graphis scripta, Hypotrachyna pseudosinuosa, Phaeophyscia rubropulchra, Porina hirsuta (KBA-L-0001677); SOUTH KOREA, North Jeolla Province, Jangsu, Mt. Youngchi, 35°38’35.5”N, 127°36’59.7”E, 905 m alt., on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000565, with Bacidia ekmaniana, Lecidella euphorea (KBA-L-0002037); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000568, with Lecidella euphorea, Phaeophyscia adiastola (KBA-L-0002040); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000570, with Anisomeridium polypori, Lecidella euphorea (KBA-L-0002042); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000571, with Anisomeridium polypori, Pertusaria sp. (KBA-L-0002043); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000572, with Caloplaca flavorubescens, Lecanora hafelliana, Lecidella euphorea (KBA-L-0002044); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000573, with Bacidia ekmaniana, Lecanora aff. perflexuosa, Lecidella euphorea, Porina hirsuta (KBA-L-0002045); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000574, with Rinodina orientalis (KBA-L-0002046); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000575, with Graphis scripta, Rinodina orientalis (KBA-L-0002047); same locality, on bark of Carpinus tschonoskii, 08 Jun 2021, B.G. Lee & H.J. Lee 2021-000576, with Pyrenula laevigata, Rinodina orientalis, Scoliciosporum chlorococcum (KBA-L-0002048); SOUTH KOREA, North Jeolla Province, Jangsu, Mt. Jangan, 35°38’35.0”N, 127°36’57.8”E, 925 m alt., on bark of Carpinus tschonoskii, 09 Jun 2021, B.G. Lee & H.J. Lee 2021-000774, with Lecidella euphorea, Micarea aff. elachista (KBA-L-0002046); same locality, on bark of Carpinus tschonoskii, 09 Jun 2021, B.G. Lee & H.J. Lee 2021-000775, with Lecanora sp., Lecidella euphorea (KBA-L-0002047); same locality, on bark of Carpinus tschonoskii, 09 Jun 2021, B.G. Lee & H.J. Lee 2021-000776, with Rinodina orientalis (KBA-L-0002048; ON115810 for LSU, ON260925 for RPB2); same locality, on bark of Carpinus tschonoskii, 09 Jun 2021, B.G. Lee & H.J. Lee 2021-000778, with Caloplaca gordejevii, Lecanora hafelliana, Rinodina orientalis (KBA-L-0002050; ON260926 for RPB2); SOUTH KOREA, North Jeolla Province, Jangsu, Mt. Jangan, 35°38’36.1”N, 127°36’53.9”E, 950 m alt., on bark of Carpinus tschonoskii, 09 Jun 2021, B.G. Lee & H.J. Lee 2021-000788, with Lecanora sp., Lecidella euphorea, Myelochroa xantholepis, Pertusaria multipuncta, Rinodina archaea, Rinodina orientalis (KBA-L-0002060).

Molecular phylogeny

Two phylogenetic trees (mtSSU, and the concatenation of all mtSSU, LSU and RPB2) for the genus Arthonia and related genera were produced from overall 184 sequences (72 for mtSSU, 55 for LSU, and 57 for RPB2) of GenBank and eight new sequences (two for mtSSU, and each three for LSU and RPB2) from the new record (Table 2). The sequences of the new record were positioned in the Bryostigma clade in both the mtSSU and the concatenation trees (Figs. 3 and 4). Both trees describe that the sequences of the new record are located in the clade and between the tips of A. apatetica, supported by the bootstrap values of 98 (mtSSU) and 100 (concatenation), and the posterior probabilities of 1.0 (both) for the branches.

Species list and DNA sequence information employed for phylogenetic analysis
No. Species mtSSU LSU RPB2 Voucher
1 Arthonia apatetica KJ850992 KJ851045 KJ851125 UPS:Svensson 2017 (Europe)
2 Arthonia apatetica KJ850993 KJ851056 UPS:Hermansson 17868b (Europe)
3 Arthonia apatetica KJ850994 KJ851050 KJ851126 UPS:Svensson 1939 (Europe)
4 Arthonia apatetica ON116972 ON115808 ON260924 KBA-L-0001570
5 Arthonia apatetica ON116973 ON115809 KBA-L-0001654
6 Arthonia apatetica ON115810 ON260925 KBA-L-0002048
7 Arthonia apatetica ON260926 KBA-L-0002050
8 Arthonia apotheciorum KJ850970 KJ851148 UPS:Frisch 11/Se23
9 Arthonia biatoricola KJ850990 KJ851149 UPS:Thor 24350
10 Arthonia caesia FJ469671 FJ469668 FJ469670 AFTOL-ID 775
11 Arthonia calcarea EU704065 EU704029 Ertz 7540
12 Arthonia didyma EU704047 EU704083 EU704010 Ertz 7587
13 Arthonia didyma KJ851081 KJ851106 UPS:Frisch 11/Se41
14 Arthonia dispersa AY571383 AY571381 UPSC 2583
15 Arthonia eos KJ850987 KJ851053 KJ851134 UPS:Thor 26000
16 Arthonia glebosa MZ502372 JR 0.154
17 Arthonia granithophila KJ850981 KJ851049 KJ851107 UPS:Frisch 10/Se74
18 Arthonia graphidicola KJ850980 KJ851034 UPS:Frisch 10/Jp102
19 Arthonia ilicina KJ850982 KJ851069 UPS:McCune 31067
20 Arthonia incarnata KY983974 KY983982 Frisch 13/Jp215
21 Arthonia incarnata KY983976 KY983984 Thor 26035
22 Arthonia lapidicola KJ850997 KJ851070 KJ851119 UPS:Frisch 11/Se47
23 Arthonia lobariellae MT153920 MT153975 A.F.25324
24 Arthonia mediella KJ851014 KJ851032 KJ851133 UPS:Frisch 11/Se22
25 Arthonia mediella KJ851015 KJ851031 KJ851132 UPS:Thor 12/8
26 Arthonia molendoi KJ851000 KJ851051 KJ851117 UPS:Frisch 11/Se36
27 Arthonia molendoi MH177777 MH177781 MH177770 Frisch 17/No10
28 Arthonia neglectula KJ850989 KJ851037 KJ851118 UPS:Frisch 10/Se91
29 Arthonia parietinaria MH177778 MH177782 MH177771 Frisch 17/No25
30 Arthonia peltigerina KJ850998 KJ851122 UPS:Westberg Frisch 11/Se46
31 Arthonia phaeophysciae MN955428 MN963753 KT180627002
32 Arthonia physcidiicola KF707646 KF707657 UPS:Frisch 11/Ug318
33 Arthonia picea MG201836 MG201847 Frisch 13Jp107
34 Arthonia picea MG201837 MG201848 Frisch 13Jp124
35 Arthonia pinastri MN842780 MN842779 MFLU 17-3497
36 Arthonia punctiformis KJ850973 KJ851044 KJ851113 UPS:Thor 26158
37 Arthonia aff. punctiformis KJ850975 KJ851043 UPS:Thor 24702
38 Arthonia radiata EU704048 EU704011 Ertz s.n.
39 Arthonia radiata KJ850969 KJ851109 UPS:Frisch 11/Se25
40 Arthonia ruana MG495133 MG495139 KoLRI 044321
41 Arthonia ruana MG495134 MG495140 KoLRI 044324
42 Arthonia ruana MG495135 MG495142 MG589416 KoLRI 044331
43 Arthonia ruana MG495136 MG589417 KoLRI 044332
44 Arthonia ruana MG495141 MG589415 KoLRI 044325
45 Arthonia rubrocincta GU327684 Nelsen 4010
46 Arthonia sanguinaria MG201838 MG201846 Frisch13Jp59
47 Arthonia spadicea KJ851029 KJ851115 UPS:Frisch 11/Se31
48 Arthonia spadicea KJ851071 KJ851116 UPS:Thor 26200
49 Arthonia stereocaulina KJ850999 KJ851062 UPS:Westberg Frisch 11/Se48
50 Arthonia subfuscicola KJ850972 KJ851111 UPS:Frisch 11/Se15
51 Arthonia subfuscicola KJ851110 UPS:Thor 44866
52 Arthonia subfuscicola KJ850971 UPS:Thor 11/1
53 Arthonia thoriana MG207685 Sanderson 2174
54 Arthonia thoriana MG207687 Sanderson 2176
55 Arthonia thoriana MG207686 Sanderson 2175
56 Arthonia toensbergii MH177775 MH177779 Frisch N4-2-Pa4-1
57 Arthonia toensbergii MH177776 MH177780 Frisch N1-2-Pa6-1
58 Arthonia ulleungdoensis MG495138 MG495144 MG589418 KoLRI 044339
59 Arthonia sp. EU704049 EU704084 EU704012 Ertz 7775
60 Arthonia sp. EU704050 EU704013 Ertz 9090
61 Arthonia sp. KJ850985 KJ851064 KJ851129 UPS:Frisch 11/Ug212
62 Arthonia sp. KJ850986 KJ851068 KJ851131 UPS:Frisch 11/Ug218
63 Arthonia sp. KJ850995 KJ851046 KJ851120 UPS:Svensson 2148
64 Arthonia sp. KJ850996 KJ851121 UPS:Svensson 2295
65 Arthonia sp. ‘lobariicola’ KJ851002 KJ851036 KJ851128 UPS:Frisch 10/Jp124
66 Arthonia sp. KJ851019 KJ851082 KJ851123 UPS:Svensson 2324
67 Arthonia sp. KJ851025 KJ851039 KJ851100 UPS:Frisch 11/Ug41
68 Arthothelium galapagoense HQ454515 HQ454657 Ertz 11654
69 Arthothelium galapagoense HQ454516 HQ454658 Ertz 11790
70 Arthothelium norvegicum KJ851003 KJ851038 KJ851114 UPS:McCune 31061
71 Arthothelium orbilliferum KY983977 TRH-L-15449
72 Arthothelium spectabile KP870144 KP870160 TNS:Frisch 12Jp179a
73 Arthothelium spectabile MN955429 MN947670 Frisch 12Jp164
74 Bryostigma dokdoense MH510022 MH510019 SK L06
75 Bryostigma dokdoense MH510023 MH510021 SK L04
76 Bryostigma muscigenum KJ850991 KJ851052 KJ851124 UPS:Thor 26206
77 Opegrapha lithyrga EU704068 EU704096 EU704032 Ertz 8784
78 Opegrapha niveoatra EU704070 EU704098 EU704034 Ertz 7529
79 Opegrapha vermicellifera EU704077 EU704105 EU704041 Ertz 7562
80 Opegrapha vulgata EU704080 EU704108 EU704044 Ertz 7564
81 Opegrapha zonata EU704081 EU704109 EU704045 Vigneron 104
Overall 72 55 57

DNA sequences which were generated for the new record A. apatetica in this study, are presented in bold. All others were obtained from GenBank. The species names are followed by GenBank accession numbers and voucher information. mtSSU, mitochondrial small subunit; LSU, nuclear large subunit ribosomal RNA; RPB2, RNA polymerase subunit II; Voucher, voucher information.

Fig. 3. Phylogenetic relationships among available species in the genus Arthonia based on a maximum likelihood analysis of the dataset of mtSSU sequences.
The tree was rooted with Opegrapha sequences based on Lee and Hur (2016). Maximum likelihood bootstrap values ≥ 70% and posterior probabilities ≥ 95% are shown above internal branches. Branches with bootstrap values ≥ 90% are shown in bold. The sequences of newly recorded Arthonia apatetica specimens are presented in bold, and all species names are followed by the GenBank accession numbers. Table 2 provides the species related to the specific GenBank accession numbers and voucher information.

Fig. 4. Phylogenetic relationships among available species in the genus Arthonia based on a maximum likelihood analysis of the dataset of the concatenated dataset of all mitochondrial small subunit (mtSSU), nuclear large subunit ribosomal RNA (LSU), and RNA polymerase subunit II (RPB2) sequences.
The tree was rooted with Opegrapha sequences based on Lee and Hur (2016). Maximum likelihood bootstrap values ≥ 70% and posterior probabilities ≥ 95% are shown above internal branches. Branches with bootstrap values ≥ 90% are shown in bold. The sequences of newly recorded Arthonia apatetica specimens are presented in bold, and all species names are followed by the voucher information. Table 2 provides the species related to the specific GenBank accession numbers and voucher information.

Key to the species of Arthonia in northeastern Asia (54 taxa)

An updated taxonomic key is introduced for the species of Arthonia in northeastern Asia (Korea, Japan, and China) (Table 3). Based on the key of Lee and Hur (2016), eight species were additionally reported from Korea and Japan, i.e., A. almquistii Vain., A. apatetica, A. dokdoensis S.Y. Kondr., Lőkös, B.G. Lee, J.J. Woo & Hur, A. incarnata Kullh. ex Almq., A. lopingensis Zahlbr., A. picea Vain., A. sanguinaria Frisch & Y. Ohmura, and A. ulleungdoensis B.G. Lee & Hur (Zhurbenko et al., 2015; Zhurbenko et al., 2015, 2018; Kondratyuk et al., 2019; Lee and Hur, 2019). Although previously recorded in China, A. lopingensis was newly discovered in Japan with specific information (Frisch et al., 2018).

Key to the species of Arthonia in Northeast Asia (Korea, Japan and China)
1. Lichenicolous 2
Not lichenicolous 7
2. Hypothecium dark brown or blackish brown; epihymenium dark brown, on Rinodina species A. rinodinicola
Hypothecium hyaline to pale brown; epihymenium pale olive-brown or yellow-orange 3
3. Epihymenium yellow to orange; on Graphis species A. lopingensis
Epihymenium pale olive-brown to brown 4
4. On Phaeophyscia species; asci 35–45 × 16–20 μm A. phaeophysciae
On Rhizoplaca species; asci 28–33 × 14–17 μm A. clemens
On various species of Porpidiaceae and Agyriaceae, or Amygdalaria pelobotryon, esp. Trapelia species in NE Asia; asci 28–38 × 17–20 μm A. almquistii
On Caloplaca, Orientophila, or Xanthoria species 5
5. Ascospores 3-septate, 12–17 × 4.5–6 μm; on Xanthoria species, but usually on bark A. varia
Ascospores 1-septate, 8–13 × 4–6 μm 6
6. Ascospores 1-septate, 10–13 × 4–5 μm; on Orientophila species A. dokdoensis
Ascospores 1-septate, 8–11 × 4–6 μm; on Caloplaca or Xanthoria species A. molendoi
7. Foliicolous, saxicolous or terricolous 8
Corticolous 16
8. Foliicolous 9
Saxicolous or terricolous 13
9. Ascospores grayish brown when mature 10
Ascospores colorless when mature 12
10. Thallus K+ purple; ascospores 1-septate; usually on bark A. vinosa
Thallus K−; ascospores 2-septate 11
11. Apothecia blackish brown to black, usually rounded to oval, naked or with a very thin, indistinct, marginal pruina; ascospores 11–19 (–24) × 4–6 (–9) μm, 3–3.5 times as long as broad A. trilocularis
Apothecia blackish brown with a purplish tinge, irregularly angular, with a thin but distinct marginal pruina which sometimes spreads over the apothecial surface; ascospores 13–19 × 2.5–4.5 μm, 3–5 times as long as broad A. fuscocyanea
12. Apothecia light to dark brown, non-pruinose; ascospores consistently 2-septate, 11–16 × 3–5 μm A. accolens
Apothecia dark greenish gray but with a whitish pruina which gives them a conspicuous bluish tinge; ascospores variably 2- to 4-septate, 11–25 (–29) × 3.5–6.5 (–8) μm A. cyanea
13. Terricolous; thick thallus; dark brown hypothecium; ascospores single to 1-septate, hyaline A. glebosa
Saxicolous 14
14. Thallus foliose, thick A. rufidula
Thallus crustose, thin 15
15. Thallus rimose-cracked, thin; apothecia slit, lirellate, simple, branched or stellate, 0.6–1.2 (–2) × (0.1–) 0.15–0.3 mm; ascospores 3-septate, 16–20 (–24) × 4–5 (–6.5) μm; especially on limestone A. calcarea
Thallus partly immersed to irregularly scurfy granularverrucose, thin; apothecia globose, simple, convex, dark brown to black, not branched, (0.1–) 0.2–0.4 (–0.5) mm; ascospores 1-septate, 11–15 (–18) × 4–7 μm; on limestone or basalt A. lapidicola
16. Apothecia brownish black or black 17
Apothecia colored, not just brownish or blackish 45
17. Ascospores 1-septate 18
Ascospores 2-septate or multi-septate 22
18. Apothecia curved or flexuose, often branching A. excipienda
Apothecia rounded, not branching 19
19. Thallus white to pale fawn, K+ purple A. vinosa
Thallus green to dark gray, K− 20
20. Apothecia 0.1–0.4 mm; thallus olive-green A. apatetica
Apothecia 0.2–1.5 mm; thallus dark gray to black-gray 21
21. Apothecia 0.75–1 mm; ascospores 20–23 × 7–9 μm A. henoniana
Apothecia 0.2–1.5 mm; ascospores 7–11 × 3–4 μm A. spadicea
22. Ascospores 2-septate, sometimes 1-septate, 11–12 × 4.5–5 μm; apothecia sub-angular or irregularly stellate; thallus whitish macular A. biseptella
Ascospores 3-septate or more 23
23. Ascospores 3-septate 24
Ascospores more than 3-septate 30
24. Thallus bleached as whitish or grayish macular 25
Thallus not bleaching 27
25. Apothecia stellate, few radiate; ascospores bigger, 17–19 × 6–7 μm A. astropica
Apothecia linear, round or thinly stellate; ascospores smaller, 12–17 × 4.5–6 μm 26
26. Apothecia linear, round to irregular, flat, not branching; ascospores 12–17 × (3–) 4.5–6 μm; thallus light gray; on bark or sometimes on Xanthoria parietina A. varia
Apothecia divided or thinly stellate, erumpent, dendritically branching; ascospores 15–17 × 5–6 μm; thallus white macular; on bark only A. varia var. stenograpella
27. Locules unequal, apical cell enlarged; thallus cinnamon-gray, shiny A. schoepfiae
Locules equally divided; thallus whitish or grayish 28
28. Photobiont present, chlorococooid; apothecia rounded to subangular; ascospores 7.4–18 × 3.8–5.1 μm; I+ blue on apothecial section A. zelkovae
Photobiont absent; apothecia rounded to irregularly branched 29
29. Apothecia brown, distinctly pruinose; I+ blue turning red on apothecial section; hypothecium hyaline; paraphysoidal tips not thickened, without distinct pigment caps A. glaucella
Apothecia black, usually epruinose; I+ persistently blue on apothecial section; hypothecium tinged light brownish; paraphysoidal tip cells slightly thickened, with apical pigment caps A. pinastri
30. Ascospores up to 5-septate 31
Ascospores 5-septate or more 40
31. Apothecia rounded to substellate; ascospores less than 20 μm long 32
Apothecia mostly rounded or orbicular to angular, sometimes linear to stellate; ascospores over 20 μm long at maximum 35
32. Apothecia rounded; thallus yellowish gray; Ascospores 11–15 × 5–6 μm; on Rhus vernix A. vernicis
Apothecia various, rounded, lirellate or stellate; thallus whitish gray to greenish gray 33
33. Heavily blue-gray or whitish pruinose on apothecia; ascospore end cell noticeably enlarged A. cinereopruinosa
Epruinose or almost not pruinose on apothecia; ascospore cells equal and end cell not enlarged 34
34. Ascospores 3- to 4-septate, 15–20 × 4.5–6 μm; I+ blue on hymenium A. radiata
Ascospores 4- to 5-septate, 14–16 × 4–6 μm; I+ purplish red on hymenium A. taediosula
35. Locules unequal, end cells or center cells enlarged; spores larger, 16–32 × 6–12 μm 36
Locules equal, certain cells not enlarged; spores smaller, 13–23 × 4–7 μm 39
36. Ascospores usually 5-septate; apothecia rounded, angular or sometimes elongated 37
Ascospores usually 3- or 4-septate; apothecia irregularly rounded to elongated or stellate 38
37. Ascospores usually 5-septate, sometimes 4- or 6-septate, one or both spore end cells enlarged, 22–32 × (6–) 9–12 μm; apothecia round, somethimes elongated up to 1.2 mm, flat to convex, black with yellow tinge, 0.3–0.7 (–1.5) mm diam.; thallus smooth to rough and chinky, greenish gray to white A. complanata
Ascospores consistently 5-septate, two intermediate cells greater than else, 20–26 × 6.5–8 μm; apothecia orbicular or angular, not radiant, flat, black, 0.15–0.25 mm diam.; thallus smooth, gray-white macular formed A. gregantula
38. Apothecia irregularly rounded or elongated, 0.1–0.6 mm diam.; ascospores 3- or 4-(5-)septate, (16–)18–23 × 7–9 μm; pycnidia 60–80 μm, pycnoconidia 7–9 × 1 μm A. ilicinella
Apothecia linear, or more usually irregularly branched to stellate, up to 2 mm diam.; ascospores (2-) 3- or 4-septate, (13–) 16–22 (–24) × (5–) 6–7 (–9) μm; pycnidia 40–60 μm; pycnoconidia 4.5–5.5 × 0.5–1 μm A. stellaris
39. Ascospores usually 3-septate (sometimes up to 5-septate), 13–23 × 4–7 μm; I+ dark vinose-red on hymenium A. punctiformis
Ascospores mostly 4-septate, 15.5–21.3 × 4.5–6.3 μm; I+ blue on epihymenium and hymenium, and I+ vinose-red on ascus and spores A. coreana
40. Prothallus with black thin line; apothecia lirellate to sublinear 41
Prothallus indistinct or sometimes with brown line; apothecia rounded to substellate 42
41. Ascospores 5- to 6-septate, two center locules distinctly larger, 21–24 × 6–7 μm; asci 8-spored; thallus slightly olivaceous, shiny A. leioplacella
Ascospores 4- to 8-septate, end cell enlarged, 28–37 × 8–12 μm; asci 4- to 6-spored; thallus milky-white macular A. linearis
42. Hypothecium indistinct or ± hyaline; epihymenium blackish; ascospores 5- to 6-septate, (13–) 18–23 × 5–8 μm; apothecia 0.01–0.3 (up to 1–1.5) mm A. reniformis
Hypothecium brownish; epihymenium brownish; ascospores over 6-septate at the maximum 43
43. Ascospores 5- to 7-septate, 18–24 × 6–8 μm; apothecia 0.5–0.6 mm A. pertabescens
Ascospores 4 to 8 transeversely septate and 0 to 3 longitudinally septate, muriform; apothecia 0.7–2.6 mm 44
44. Apothecia irregular to stellate, punctiform; apothecial section 70–95 μm thick; hymenium 35–50 μm thick; asci 8-spored; ascospores brown when mature, 15–26 × 7–10.5 μm A. ruana
Apothecia rounded, non-punctiform; apothecial section 100–130 μm thick; hymenium 50–60 μm thick; asci 2- to 6-spored; ascospores colorless with dark septum when mature, 21–31 × 9–13 μm A. ulleungdoensis
45. Ascospores 9- to 13-septate, over 40 μm in length A. ochropallens
Ascospores 3- to 7-septate, less than 40 μm in length 46
46. Apothecia pruinose or epruinose, violet-brown or brown with purple-red pigment 47
Ascospores epruinose, cinnamon or pale yellow 50
47. Apothecia epruinose, dark red to dark brown-red 48
Ascospores pruinose, violet-brown to cinnabar red 49
48. Apothecia dark red; ascospores 4–5-septate, colorless with pale brown wall when mature, 17–19.5 × 6–7.5 μm; confluentic acid or ±psoromic acid present A. sanguinaria
Apothecia dark brown-red; ascospores 4–6-septate, brown when mature, 27–32.5 × 10–11.5 μm; no substance above A. picea
49. Apothecia violet-brown with grayish white pruina at margin; ascospores usually 5-septate, 21–24 (–28) × 7–8 μm A. adspersa
Apothecia brown to blackish with cinnabar-red pruina at margin and white pruina in center; ascospres 3- to 5-septate (up to 7-septate), 20–33 × 5.5–11 μm A. cinnabarina (Coniocarpon cinnabarinum)
50. Apothecia cinnamon or somewhat red-brown; anthraquinone (parietin) or depsidone(± psoromic acid, ± norstictic acid) present A. lopingensis
Apothecia pale straw-yellow; xanthone (arthothelin) or lichexanthone present or no substance 51
51. Pycnidia among apothecia, I+ purplish red on hymenium A. perpallens (A. antillarum f. spermogonifera)
Pycnidia unknown 52
52. Apothecia adnate and convex; hymenium I+ vinose-red or orange-red; UV– or UV+ white on thallus 53
Apothecia innate and almost flat; hymenium I+ blue; UV+ orange to red on apothecia 54
53. Prothallus pale yellow when present; asci 20–40 × 13–18 μm; ascospores generally 3-septate; xanthone present A. superpallens
Prothallus black when present; asci 40–50 × 14–20 μm; ascospores 2-septate; no substance A. incarnata
54. UV+ orange on apothecia (indicative of lichexanthone); apothecia smaller, 0.4–0.5 mm A. antillarum
UV+ red on apothecia (indicative of xanthone); apothecia larger and less branched, 0.3–0.7 mm A. parantillarum


Discrepancy between the genus Bryostigma and the Bryostigma clade

It should be noted that there is a discrepancy between the genus Bryostigma and the Bryostigma clade. Although most species in the Bryostigma clade including recently reported new species (Fleischhacker et al., 2016; Etayo, 2017; Frisch and Holien, 2018; Kondratyuk et al., 2019, 2020a) are classified into the genus Bryostigma (Kondratyuk et al., 2020b), many species in the clade do not correspond to the definition of the genus Bryostigma of Poelt and Döbbeler (1979). Although the authors emphasized bryophytes for the genus substrate when they defined the genus, just one single species, B. muscigenum, is muscicolous yet all other species in the clade are lichenicolous, corticolous, saxicolous or terricolous. In addition, there has been a wrong taxonomy for B. apotheciorum (Kondratyuk et al., 2020b). Arthonia apotheciorum was classified far from the Bryostigma clade (Frisch et al., 2014; Lee and Hur, 2016), and the species should be remained in Arthonia s. lat. Kondratyuk et al. (2020b) confused the species assumingly with A. apatetica for the genus Bryostigma. The definition of the genus Bryostigma should be emended for the Bryostigma clade based on a preliminary definition (Frisch et al., 2014; Frisch and Holien, 2018) or remained for further studies without just combinationing of species in the clade to the genus. Other species possibly belonging to the Bryostigma clade should be considered in the new definition (e.g., A. fuscopurpurea, A. patellulata, and many other lichenicolous species).

적 요

티끌별지의 [Arthonia apatetica (A. Massal.) Th. Fr.]가 북동아시아지역 미기록종으로서 한국에서 최초 발견되었다. 너무 작아서 간과되기 쉬운 이 미생물은, 작고 검은 자낭과 속에 1-격벽의 포자를 가지고 있고, 수정결정이 없으며, 옅은 갈색의 자낭하층을 보여주고, 자낭상층의 균사체가 수평으로 정렬되어있고, 클로로코코이드(chlorococcoid) 녹조류를 가진 수피착생형 지의류이다. 분자분석법을 통해 mtSSU, LSU, RPB2 유전자 염기서열을 이용 종합 분석한 결과, 한국에서 채집된 지의류 표본이 티끌별지의(A. apatetica)임을 증명하였다. 이번 연구를 통해 해당 미기록종의 유전정보를 아시아 최초로 확보하였다. 더불어 북동아시아 별지의(Arthonia)속 지의류 총 54분류군에 대한 분류검색표를 새롭게 마련하였다.


This work was supported by a grant from Korean Forest Service Program through the Korea National Arboretum (KNA-202003127AF-00) for the forested wetland conservation of Korea.

Conflict of Interest

The authors declare no conflicts of interest. All the experiments undertaken in this study comply with the current laws of the country where they were performed.

  1. Bouckaert RR and Drummond AJ. 2017. bModelTest: Bayesian phylogenetic site model averaging and model comparison. BMC Evol. Biol. 17, 42.
    Pubmed KoreaMed CrossRef
  2. Bouckaert R, Vaughan TG, Barido-Sottani J, Duchêne S, Fourment M, Gavryushkina A, Heled J, Jones G, Kühnert D, and De Maio NDe Maio N, et al. 2019. BEAST 2.5: an advanced software platform for Bayesian evolutionary analysis. PLoS Comput. Biol. 15, e1006650.
    Pubmed KoreaMed CrossRef
  3. Brodo IM. 1995. Lichens and lichinicolous fungi of the Queen Charlotte Islands, British Columbia, Canada: 1. Introduction and new records for B. C., Canada and North America. Mycotaxon 56, 135-173.
  4. Coppins BJ. 1989. Notes on the Arthoniaceae in the British Isles. Lichenologist 21, 195-216.
  5. Edler D, Klein J, Antonelli A, and Silvestro D. 2021. raxmlGUI 2.0 beta: A graphical interface and toolkit for phylogenetic analyses using RAxML. Methods Ecol. Evol. 12, 373-377.
  6. Ekman S. 2001. Molecular phylogeny of the Bacidiaceae (Lecanorales, lichenized Ascomycota). Mycol. Res. 105, 783-797.
  7. Etayo J. Hongos liquenícolas de Ecuador. Opera lilloana 50. Fundación Miguel Lillo, San Miguel de Tucumán, Argentina. ISSN 950-668-010-8.
  8. Fleischhacker A, Grube M, Frisch A, Obermayer W, and Hafellner J. 2016. Arthonia parietinaria-a common but frequently misunderstood lichenicolous fungus on species of the Xanthoria parietina-group. Fungal Biol. 120, 1341-1353.
    Pubmed CrossRef
  9. Frisch A, Grube M, Kashiwadani H, and Ohmura Y. 2018. Arthoniaceae with reddish, K+ purple ascomata in Japan. Phytotaxa 356, 19-33.
  10. Frisch A and Holien H. 2018. Arthonia toensbergii, a new lichenicolous fungus on Mycoblastus affinis from the boreal rainforests in Norway. Graph. Scr. 30, 34-43.
  11. Frisch A, Thor G, Ertz D, and Grube M. 2014. The Arthonialean challenge: restructuring Arthoniaceae. Taxon 63, 727-744.
  12. Frisch A, Thor G, Moon KH, and Ohmura Y. 2017. Arthonia incarnata (Arthoniaceae), a rare and poorly known old-growth forest lichen new to Asia. Nord. J. Bot. 35, 587-594.
  13. Halıcı MG and Kahraman M. 2021. DNA barcoding and morphological observations of three lichenized fungal species from James Ross Island (Antarctic Peninsula). Ukr. Antarct. J. 1, 123-148.
  14. Hall TA. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symp. Ser. 41, 95-98.
  15. Hauck M and Javkhlan S. 2006. Additions to the lichen flora of Mongolia: records from Khentey and Khangay. Willdenowia 36, 895-912.
  16. Hawksworth DL. 1988. The variety of fungal-algal symbioses, their evolutionary significance, and the nature of lichens. Bot. J. Linn. Soc. 96, 3-20.
  17. Kondratyuk S, Lőkös L, Halda J, Lee BG, Jang SH, Woo JJ, Park JS, Oh SO, Han SK, and Hur JS. 2019. Arthonia dokdoensis and Rufoplaca toktoana-two new taxa from Dokdo Islands (South Korea). Mycobiology 47, 355-367.
    Pubmed KoreaMed CrossRef
  18. Kondratyuk SY, Lőkös L, Oh SO, Kondratiuk TO, Parnikoza IY, and Hur JS. 2020a. New and noteworthy lichen-forming and lichenicolous fungi, 11. Acta Bot. Hung. 62, 225-291.
  19. Kondratyuk SY, Upreti DK, Mishra GK, Nayaka S, Ingle KK, Orlov OO, Kondratiuk AS, Lőkös L, Farkas E, and Woo JJWoo JJ, et al. 2020b. New and noteworthy lichen-forming and lichenicolous fungi 10. Acta Bot. Hung. 62, 69-108.
  20. Lee BG and Hur JS. 2016. Three new species and nine new records in the genus Arthonia from South Korea. Mycobiology 44, 202-216.
    Pubmed KoreaMed CrossRef
  21. Lee BG and Hur JS. 2019. Arthonia ulleungdoensis, a new lichenized fungus from Ulleung Island, South Korea. Microorganisms 7, 205.
    Pubmed KoreaMed CrossRef
  22. Lee BG and Hur JS. 2020. A new lichenized fungus, Lecanora baekdudaeganensis, from South Korea, with a taxonomic key for Korean Lecanora species. MycoKeys 70, 39-58.
    Pubmed KoreaMed CrossRef
  23. Liu YJ, Whelen S, and Hall BD. 1999. Phylogenetic relationships among ascomycetes: evidence from an RNA polymerse II subunit. Mol. Biol. Evol. 16, 1799-1808.
    Pubmed CrossRef
  24. Lücking R, Hodkinson BP, and Leavitt SD. 2017. The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota-Approaching one thousand genera. Bryologist 119, 361-416.
  25. Nash TH 3rd, Gries C, and Bungartz F. Lichen Flora of the Greater Sonoran Desert Region. Lichens Unlimited/Arizona State University, Tempe, Arizona, USA.
  26. Orange A, James PW, and White FJ. Microchemical Methods for the Identification of Lichens. British Lichen Society, London, UK.
  27. Poelt J and Döbbeler P. 1979. Bryostigma leucodontis nov. gen. et spec., a New Lichen With Almost Invisible Fruiting Bodies. Plant Syst. Evol. 131, 211-216.
  28. Rambaut A. FigTree v1.4.2. Institute of Evolutionary Biology. University of Edinburgh, Edinburgh, UK.
  29. Rehner SA and Samuels GJ. 1994. Taxonomy and phylogeny of Gliocladium analysed from nuclear large subunit ribosomal DNA sequences. Mycol. Res. 98, 625-634.
  30. Santesson R. The lichens and lichenicolous fungi of Sweden and Norway. SBT-förlaget, Lund, Sweden.
  31. Sedelnikova NV and Taran GS. 2000. The main characteristics of the lichenoflora of the Elizarovskiy zakaznik (lower Ob'River). Krylovia 2, 46-53.
  32. Smith CW, Aptroot A, Coppins BJ, Fletcher A, Gilbert OL, James PW, and Wolseley PA. The Lichens of Great Britain and Ireland. British Lichen Society, London, UK.
  33. Stecher G, Tamura K, and Kumar S. 2020. Molecular Evolutionary Genetics Analysis (MEGA) for macOS. Mol. Biol. Evol. 37, 1237-1239.
    Pubmed KoreaMed CrossRef
  34. Wirth V. Die Flechten Baden-Wurttembergs 1 and 2. Verlag Eugen Ulmer, Stuttgart, Germany.
  35. Zhurbenko MP, Frisch A, Ohmura Y, and Thor G. 2015. Lichenicolous fungi from Japan and Korea: new species, new records and a first synopsis for Japan. Herzogia 28, 762-789.
  36. Zoller S, Scheidegger C, and Sperisen C. 1999. PCR primers for the amplification of mitochondrial small subunit ribosomal DNA of lichen-forming ascomycetes. Lichenologist 31, 511-516.

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