Research Article |
Corresponding author: Beeyoung Gun Lee ( gitanoblue@koagi.or.kr ) Academic editor: Gerhard Rambold
© 2022 Beeyoung Gun Lee, Jae-Seoun Hur.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lee BG, Hur J-S (2022) A new species and four new records of Bacidia (Lecanorales, Ramalinaceae) from South Korea, with a key to Korean species. MycoKeys 93: 107-130. https://doi.org/10.3897/mycokeys.93.89283
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A new species, Bacidia fuscopallida Lee & Heo and four new records, B. ekmaniana R. C. Harris, Ladd & Lendemer, B. friesiana (Hepp) Körb., B. heterochroa (Müll. Arg.) Zahlbr. and B. suffusa (Fr.) A. Schneid., are described from South Korea. Bacidia fuscopallida differs from B. diffracta S. Ekman, the most similar species, by warted but non-granular thallus, paler and smaller apothecia without pruina, proper exciple without crystals, over 11-septate ascospores and smaller pycnidia and pycnoconidia. Bacidia ekmaniana is recorded new to Asia, B. heterochroa is reported new to northeastern Asia and B. friesiana and B. suffusa are new to Korea. Molecular analyses employing internal transcribed spacer (ITS) sequences strongly support the classification of the five species of Bacidia. A surrogate key is provided to assist in the identification of all 19 taxa in Bacidia of Korea.
biodiversity, corticolous, lichen, phylogeny, taxonomy
Bacidia has become a species-rich genus since
Bacidia is one of the least explored genera in Korea and the genus has just been reported since the 2010s. Since
This study describes a new species and four new records of the lichen genus Bacidia. Field surveys for the lichen biodiversity in the main mountains of Korea, i.e. Baekdudaegan, and several forested wetlands of South Korea were carried out during the spring to summer of 2019–2021 and 54 specimens of Bacidia were collected from barks of deciduous wide-leaved trees and shrubs (Fig.
Hand sections were prepared manually with a razor blade under a stereomicroscope (Olympus optical SZ51; Olympus, Tokyo, Japan), examined under a compound microscope (Nikon Eclipse E400; Nikon, Tokyo, Japan) and pictured using a software programme (NIS-Elements D; Nikon, Tokyo, Japan) and a DS-Fi3 camera (Nikon, Tokyo, Japan) mounted on a Nikon Eclipse Ni-U microscope (Nikon, Tokyo, Japan). The ascospores were examined 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 (SD), length-to-width ratio and the number of measured spores. Thin-layer chromatography (TLC) was performed using solvent system C according to standard methods (
Hand-cut sections of ten to twenty ascomata per collected specimen were prepared for DNA isolation (Table
Species | Bacidia fuscopallida | Bacidia ekmaniana | Bacidia friesiana | Bacidia heterochroa | Bacidia suffusa |
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Specimens | KBA-L-0001010 (isotype), KBA-L-0001037 (paratype), KBA-L-0001049 (paratype) | KBA-L-0000072, KBA-L-0002037 | KBA-L-0001910, KBA-L-0001913, KBA-L-0001914, KBA-L-0001917 | KBA-L-0000386, KBA-L-0002714, KBA-L-0002727, KBA-L-0002734 | KBA-L-0000358, KBA-L-0000359, KBA-L-0000368, KBA-L-0002776, KBA-L-0002778, KBA-L-0002835 |
Ascomata sections per specimen | 20 | 10 | 20 | 10 | 10 |
Ascomata sections per species | 60 | 20 | 80 | 40 | 60 |
An independent phylogenetic tree for the genus Bacidia was produced from 84 sequences from GenBank and 12 newly-generated sequences for the new species and the new records (Table
Species list and DNA sequence information employed for phylogenetic analysis.
No. | Species | ITS | Voucher |
---|---|---|---|
1 | Bacidia absistens | AF282085 | Ekman 3223 (BG) |
2 | Bacidia albogranulosa | MK158340 | J. Malicek 9622 |
3 | Bacidia albogranulosa | MK158342 | J. Vondrak 11888 (PRA) |
4 | Bacidia arceutina | AF282083 | Ekman 3110 (BG) |
5 | Bacidia arceutina | JQ796851 | LG DNA 579 |
6 | Bacidia areolata | MH048614 | M-0182592 |
7 | Bacidia auerswaldii | AF282122 | Johansson 20 (UPS) |
8 | Bacidia bagliettoana | AF282123 | Ekman 3137 (BG) |
9 | Bacidia bagliettoana | MG838190 | O-L-175215 |
10 | Bacidia beckhausii | AF282071 | Holien 6744 (TRH) |
11 | Bacidia beckhausii | JF714252 | MSSRF Lichen Herbarium |
12 | Bacidia biatorina | AF282079 | Knutsson 94–148 |
13 | Bacidia caligans | AF282096 | Johansson 21 (UPS) |
14 | Bacidia circumspecta | MH539764 | L-13006 |
15 | Bacidia circumspecta | AF282124 | Ekman L1330 (LD) |
16 | Bacidia cylindrophora | MG926005 | Kurokawa 1692 |
17 | Bacidia cylindrophora | MG926006 | Ohmura 7091 (GZU) |
18 | Bacidia diffracta | AF282090 | Wetmore 26401 (MIN) |
19 | Bacidia diffracta | MH048620 | Harris 46555-A |
20 | Bacidia ekmaniana | ON352611 | KBA-L-0002037 |
21 | Bacidia elongata | MH048626 | M-0182571 |
22 | Bacidia elongata | MH048629 | M-0182627 |
23 | Bacidia fraxinea | AF282088 | Johansson 1620 (BG) |
24 | Bacidia friesiana | ON352609 | KBA-L-0001910 |
25 | Bacidia friesiana | ON352610 | KBA-L-0001913 |
26 | Bacidia friesiana | MH539765 | L-13159 |
27 | Bacidia fuscopallida | ON352607 | KBA-L-0001010 |
28 | Bacidia fuscopallida | ON352608 | KBA-L-0001049 |
29 | Bacidia fuscoviridis | AM292665 | Nordin 5058 (UPS) |
30 | Bacidia gigantensis | MT425200 | MCM242 |
31 | Bacidia hemipolia | AF282072 | Toensberg 25091 (BG) |
32 | Bacidia heterochroa | ON352606 | KBA-L-0000386 |
33 | Bacidia heterochroa | ON352612 | KBA-L-0002727 |
34 | Bacidia heterochroa | ON352613 | KBA-L-0002734 |
35 | Bacidia hostheleoides | AF282081 | Seaward 108121 |
36 | Bacidia incompta | AF282092 | Ekman 3144 (BG) |
37 | Bacidia incompta | MG461697 | KoLRI Udo-32 |
38 | Bacidia kurilensis | MH048612 | M-0182622 |
39 | Bacidia kurilensis | MH048610 | M-0182620 |
40 | Bacidia kurilensis | MH048611 | M-0182621 |
41 | Bacidia laurocerasi | MH048609 | Galanina 424 |
42 | Bacidia laurocerasi subsp. laurocerasi | MN483106 | Spribille 26334 (KLGO) |
43 | Bacidia laurocerasi subsp. laurocerasi | AF282078 | Wetmore 74318 (MIN) |
44 | Bacidia lutescens | MG925952 | Ekman 3655 (BG) |
45 | Bacidia lutescens | AF282082 | Ekman L1161 (LD) |
46 | Bacidia medialis | AF282102 | Ekman L1193 (LD) |
47 | Bacidia polychroa | AF282089 | Knutsson 91–215 |
48 | Bacidia rosella | AF282086 | Ekman 3117 (BG) |
49 | Bacidia rubella | AF282087 | Ekman 3021 (BG) |
50 | Bacidia rubella | HQ650644 | AFTOL-ID 1793 |
51 | Bacidia rubella | JQ796852 | LG DNA 578 |
52 | Bacidia rubella | KX132984 | LIFU076–16 |
53 | Bacidia rubella | MG461695 | AFTOL-ID 1793 |
54 | Bacidia rubella | EU266078 | Hur H06122 |
55 | Bacidia rubella | MH048630 | M-0182581 |
56 | Bacidia rubella | MK158343 | J. Vondrak 12200 (PRA) |
57 | Bacidia sabuletorum | AF282069 | Ekman 3091 (BG) |
58 | Bacidia sachalinensis | MH048621 | M-0182619 |
59 | Bacidia sachalinensis | MH048625 | M-0182624 |
60 | Bacidia schweinitzii | AF282080 | Wetmore 72619 (MIN) |
61 | Bacidia schweinitzii | KX151766 | Lendemer 31230A (NY) |
62 | Bacidia scopulicola | AF282084 | Ekman 3106 (BG) |
63 | Bacidia sigmosporae | MW622004 | P.v.d. Boom 55090 |
64 | Bacidia sipmanii | JQ796853 | LG DNA 361 |
65 | Bacidia sorediata | KX151772 | Lendemer 33787 (NY) |
66 | Bacidia sorediata | KX151775 | Barton 658 (NY) |
67 | Bacidia squamulosula | MG925955 | Kalb & Kalb in Kalb, Lich. neotrop. 405 |
68 | Bacidia subareolata | MK499342 | MFLU 16-0573 |
69 | Bacidia subincompta | AF282125 | Ekman 3413 (BG) |
70 | Bacidia subincompta | KX098342 | WSL DF231 |
71 | Bacidia suffusa | ON352605 | KBA-L-0000359 |
72 | Bacidia suffusa | ON352614 | KBA-L-0002776 |
73 | Bacidia suffusa | ON352615 | KBA-L-0002778 |
74 | Bacidia suffusa | ON352616 | KBA-L-0002835 |
75 | Bacidia suffusa | AF282091 | Wetmore 74771 (MIN) |
76 | Bacidia suffusa | AY756456 | Andersen 99 (BG) |
77 | Bacidia suffusa | MH048615 | M-0182601 |
78 | Bacidia suffusa | MH048616 | M-0182593 |
79 | Bacidia suffusa | MH048617 | M-0182594 |
80 | Bacidia suffusa | MH048618 | M-0289887 |
81 | Bacidia suffusa | MH048619 | M-0289888 |
82 | Bacidia suffusa | MW728313 | LAH 36839 |
83 | Bacidia suffusa | MW788561 | LAH 36838 |
84 | Bacidia vermifera | AF282109 | Johansson 1619 (BG) |
85 | Bacidia vermifera | KX132992 | LIFU084-16 (versA) |
86 | Bacidia wellingtonii | MG925953 | Ziviagina s.n. |
87 | Bacidia sp. | AY756133 | KoLRI Udo-32 |
88 | Bacidia sp. | KX098339 | WSL DF223 |
89 | Bacidia sp. | KX098340 | WSL DF72 |
90 | Bacidia sp. | KX098341 | WSL DF80 |
91 | Bacidia sp. | MG773660 | Palice 19352 |
92 | Biatora bacidioides | MG773663 | Palice 19221 |
93 | Biatora bacidioides | MG773664 | Palice 19685 |
94 | Biatora pontica | KF650977 | C. Printzen 6114 (BG) |
95 | Biatora pontica | MK778588 | J. Malicek 10212 |
96 | Biatora printzenii | KF650978 | C. Printzen 6837 (BG) |
Overall | 96 |
The new species is positioned in the genus Bacidia in the ITS tree (Fig.
Phylogenetic relationships amongst available species in the genus Bacidia, based on a Maximum Likelihood analysis of the dataset of ITS sequences. The tree was rooted with the sequences of the genus Biatora, based on
Bacidia fuscopallida differs from B. diffracta by generally non-granular, olive-green thallus, pale yellow-orange apothecia without pruina, the absence of crystals in proper exciple, slightly narrower ascospores with up to 15-septation and smaller pycnidia and pycnoconidia.
South Korea, Gangwon Province, Gangneung, Okgye-myeon, Mt. Seokbyung, 37°34.45'N, 128°55.00'E, 271 m alt., on bark of Acer pictum var. mono (Maxim.) Maxim. ex Franch., 17 June 2020, B.G. Lee & H.J. Lee 2020-000811, with Porina hirsuta Aptroot & K.H. Moon (holotype: KBA-L-0001011!); same locality, on bark of Acer pictum var. mono, 17 June 2020, B.G. Lee & H.J. Lee 2020-000801 (isotype: KBA-L-0001001); same locality, on bark of Acer pictum var. mono, 17 June 2020, B.G. Lee & H.J. Lee 2020-000806, with Mikhtomia gordejevii (Tomin) S.Y. Kondr., Kärnefelt, Elix, A. Thell, Jung Kim, A.S. Kondr. & Hur, Straminella varia (Hoffm.) S.Y. Kondr., Lőkös & Farkas, Phaeophyscia limbata (Poelt) Kashiw., Porina hirsuta (isotype: KBA-L-0001006); same locality, on bark of Acer pictum var. mono, 17 June 2020, B.G. Lee & H.J. Lee 2020-000810 (isotype: KBA-L-0001010; GenBank ON352607 for ITS); South Korea, Gangwon Province, Gangneung, Okgye-myeon, Mt. Seokbyung, 37°34.39' N, 128°55.01'E, 349 m alt., on bark of Quercus mongolica Fisch. ex Ledeb., 17 June 2020, B.G. Lee & H.J. Lee 2020-000837, with Opeltia flavorubescens (Huds.) S.Y. Kondr. & Hur (paratype: KBA-L-0001037); South Korea, Gangwon Province, Gangneung, Okgye-myeon, Mt. Seokbyung, 37°34.28'N, 128°54.88'E, 438 m alt., on bark of Acer triflorum Kom., 17 June 2020, B.G. Lee & H.J. Lee 2020-000849, with Biatora pacifica Printzen, Tønsberg & G. Thor (paratype: KBA-L-0001049; GenBank ON352608 for ITS).
Thallus corticolous, crustose, areoles in young stage and soon coarsely continuous or warted on aging, often overlapping for each other, rarely granular, thin when not overlapping, olivish-green, margin indeterminate, 40–90 μm thick; cortex indistinct, hyaline, up to 5 μm thick; medulla a little shown as mycelia below algal layer; photobiont chlorococcoid, cells globose to subglobose, 5–15 μm thick, algal layer composing most part of thallus, 35–80 μm thick. Prothallus indistinct or whitish-grey and endosubstratal when present.
Apothecia numerous, solitary, marginate and flat in young stage and seeming immarginate and convex on aging (consistently marginate and flat on bark of Acer triflorum), 0.1–0.7 mm diam. (mean = 0.33; SD = 0.14; n = 105). Pruina absent. Disc biatorine, thalline exciple absent, pale yellow to pale orange in young stage and slightly more blackish generally around margin when mature (much more blackish on bark of A. triflorum and Q. mongolica from young stage). Proper exciple 65–80 μm wide laterally (SD = 5.7; n = 15), with radiating hyphae of 1–2.5 μm wide (SD = 0.5; n = 10) and outermost cell 2.5–4 μm wide (SD = 0.6; n = 10), hyaline to pale yellow around rim, but darker downwards (pale yellow to pale brown) and the dark colour extending to hypothecium. Epihymenium hyaline, with a little pigment of pale yellow to pale olive-brown locally, smooth and not granular, ca. 5 μm high. Hymenium hyaline, 70–100 μm high (SD = 8.9; n = 10). Hypothecium clearly pigmented, pale orange-brown to brown, prosoplectenchymatous (irregularly arranged), 70–130 μm high (SD = 18.9; n = 10). Crystals absent or a little present in upper hypothecium. Oil droplets absent. Paraphyses simple, rarely branched at tips, 1–1.5 μm wide, tips not or little swollen, not pigmented, 1.5–2 μm wide. Asci cylindrical to narrowly clavate, 8-spored, 49–72 × 11–14 μm (SD = 7.3 (L), 0.9 (W); n = 11). Ascospores 3- to 15-septate, acicular to filiform, 24–69 × 2–3.5 μm (mean = 52.8 × 2.6 μm; SD = 8.7 (L), 0.6 (W); L/W ratio = 3.8–30.5, ratio mean = 17.6, ratio SD = 5.0; n = 104). Pycnidia black, immersed and upper half only shown, globose, 60–65 μm high and 55–75 μm wide (SD = 2.4 (H), 8.2 (W); n = 5), with brownish wall, K–. Pycnoconidia hyaline, filiform, curved or almost straight, 6–17 × 0.3–0.5 μm (mean = 10.4 × 0.5 μm; SD = 2.9 (L), 0.1 (W); n = 53).
Thallus K– or K+ slightly yellow, KC–, C–, Pd–, UV–. Epihymenium K+ purple extending to outermost layers of proper exciple, C–. No lichen substance was detected by TLC.
The species occurs on barks of Acer pictum var. mono, A. triflorum and Quercus mongolica. The species is currently known from the type collections.
The species epithet indicates the pale brown colour of the lichen’s apothecia.
The new species is similar to B. diffracta and B. polychroa (Th. Fr.) Körb. in having colourless epihymenium with pale orange-brown pigment and K+ purple reaction, distinctly pigmented hypothecium with yellow, orange or brown, long ascospores generally with L/W ratio over 10 amongst corticolous species. However, B. diffracta differs from the new species by granular thallus, darker and larger apothecia with pruina, proper exciple with radiating clusters of minute crystals, slightly wider ascospores with up to 11-septation and larger pycnidia and pynoconidia (
Species | Bacidia fuscopallida | Bacidia diffracta | Bacidia hostheleoides | Bacidia polychroa | Bacidia purpurans |
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Thallus growth form | warted, rarely granular | finely granular | wrinkled or granular to subsquamulose | finely wrinkled to warted, sometimes areolate | areolate |
Thallus colour | olivish-green | pale grey, green-grey, yellow-grey to grey | pale grey to pale green-grey | white to grey or yellow-grey | pale grey-green to dark green |
Prothallus | white-grey around margin, endosubstratal | white-pale grey between granules, endosubstratal | absent | – | white, arachnoid |
Apothecia (mm in diam.) | 0.1–0.7 | 0.5–1.1 | 0.5–0.8 | 0.4–1.2 | – |
Disc colour | pale yellow to pale orange (young); more blackish (old) | brown-orange to dark brown | brown-orange | brown-orange to dark brown | dark purple-brown to brown |
Pruina | absent | white | absent | white | absent |
Crystals in proper exciple | absent | radiating clusters of minute crystals | absent | with or without radiating clusters of minute crystals | absent |
Crystals in hymenium | small crystals at bottom | – | – | – | absent |
Epihymenium colour | colourless with pale yellow-brown pigment | colourless with pale orange-brown pigment | very pale orange | colourless with brown-orange pigment | greyish |
Hymenium height (μm) | 70–100 | 70–100 | ca. 60 | 55–100 | ca. 100 |
Hypothecium colour | pale orange-brown to brown | pale brown to orange-brown | very pale orange | brown-orange to dark brown | orange-brown |
Hypothecium height (μm) | 70–130 | – | – | – | ca. 60 |
Ascospore (μm) | 24–69 × 2–3.5 | 32–69 × 1.9–4.1 | 16–25 × 2.9–5 | 31–74 × 1.9–5 | 50–75 × 2–4 |
Ascospore L/W ratio | 4–31 | 9–27 | 4–9 | 7–30 | – |
Ascospore septation | 3–15 | 3–11 | 3–5 | 2–15 | 3–15 |
Pycnidia (μm) | 55–75 | 150 | 50–100 | 100–170 | 150–200 |
Pycnoconidia | 6–17 × 0.3–0.5 | 10–15 × 0.5–0.6 | ①10–14 × 0.5 ②6–9 × 1.6–2 | 10–17 × 0.6–0.8 | 20–25 × 0.8 |
Substance | absent | atranorin, (trace of zeorin) | absent | (trace of atranorin) | atranorin |
Reference | this study |
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The new species is more similar to B. polychroa in having coarsely continuous or warted thallus. However, B. polychroa differs from the new species by greyish thallus, darker and larger apothecia often with pruina, proper exciple often with radiating clusters of minute crystals, wider ascospores and larger pycnidia and pycnoconidia (
The new species is quite similar to B. purpurans R. C. Harris, Ladd & Lendemer in having greenish thallus with areoles and K+ purple reaction in epihymenium. However, B. purpurans differs from the new species by arachnoid prothallus, darker apothecia, green excipular rim adjacent to epihymenium, greyish epihymenium, shorter hypothecium, absence of crystals, larger ascospores and larger pycnidia and pycnoconidia (
Bacidia fuscopallida (KBA-L-0001011, holotype for A–D, G–O KBA-L-0001049 for E, F KBA-L-0001010 for P, Q) in morphology A, B habitus and apothecia on bark of Acer pictum var. mono. Olive-green thallus and pale yellow-orange apothecia C vertical section of apothecia D prothallus present around margin of habitus (red arrows) E, F habitus and apothecia growing on bark of Acer triflorum G apothecial section H epihymenium colourless or a little pigmented I epihymenium K+ purple J small crystals (red arrows) present in upper hypothecium K proper exciple pigmented with pale or colourless margin. Radiating hyphae wider to margin L photobiont composing most part of thallus M, N asci cylindrical to narrowly clavate. Ascospores not twisted in ascus O ascospores acicular to filiform up to 15-septate P pycnidia globose with brown wall Q pycnoconidia curved or almost straight. Scale bars: 1 mm (A, E); 500 μm (B, C, F); 2 mm (D); 200 μm (G); 50 μm (H–J, P); 20 μm (K, L); 10 μm (M–O, Q).
The new species can be compared with B. hostheleoides in sharing non-pruinose apothecia and proper exciple without crystals. However, B. hostheleoides differs from the new species by greyish thallus, absence of prothallus, shorter hymenium, paler hypothecium and shorter ascospores with a few septa (
Thallus corticolous, crustose, somewhat granular when young and smoother when mature, grey, greenish-grey to pale grey, margin indeterminate. Prothallus generally not detected or whitish-grey when present.
Apothecia consistently flat or slightly convex when mature, marginate, without pruina, 0.4–1.4 mm diam. (mean = 0.75, SD = 0.23, n = 104). Disc biatorine, without thalline exciple, pale straw, light brown to brown, with a distinct proper margin which is smooth to rugose and becoming thinner but still distinct when mature. Proper exciple pale brown to red-brown, paler or colourless around rim and thicker downwards, 80–120 μm wide laterally. Epihymenium hyaline, smooth but not granular, ca. 5 μm high. Hymenium hyaline, 80–140 μm high. Hypothecium red-brown, prosoplectenchymatous (irregularly arranged), 120–250 μm high. Small crystals present a little in hypothecium, dissolving in K. Oil droplets absent. Asci narrowly clavate, 8-spored, 70–105 × 8–12 μm (n = 5). Ascospores acicular to filiform, cells near head sometimes irregularly swollen, 3- to 9-septate, 52–71 × 2–4.5 μm (n = 15). Pycnidia not detected.
Four new records of B. ekmaniana (KBA-L-0000412 for A–C), B. friesiana (KBA-L-0001914 for D–F), B. heterochroa (KBA-L-0000386 for G–I) and B. suffusa (KBA-L-0000359 for J–L) in morphology A habitus and apothecia. Granular thallus with green-grey pigment and straw-coloured apothecia B–C apothecial section with colourless epihymenium, red-brown hypothecium, and pale excipulum D habitus and apothecia. Thallus pale grey with slightly brownish pigment and pale pink apothecia E, F apothecial section with greenish epihymenium G habitus and apothecia. Thallus pale yellowish-grey and black apothecia with red pigment H, I apothecial section and proper exciple with dark margin J habitus and apothecia. Thallus whitish pale grey and pruinose apothecia K, L apothecial section with radiating clusters of crystals, which produce pruina on surface. Scale bars: 500 μm (A, D, G, J); 100 μm (B, E, H, K); 50 μm (C, I, L); 20 μm (F).
Thallus K–, C–. Apothecial section K–, C–. No lichen substance was detected by TLC.
Bacidia ekmaniana is easily confused with B. schweinitzii under the microscope, as well as in the field because both species often share their habitat and the habiti of both species look similar particularly when the ascomata of the latter are paler. Both species are often detected from one specimen under the microscope and those were frequently regarded as one species, i.e. B. schweinitzii. Generally, however, B. ekmaniana differs from the latter by paler ascomata. Bacidia ekmaniana has brown but not black apothecia when mature (
Bacidia ekmaniana is more similar to B. arceutina than B. schweinitzii in morphology in having pale ascomata. However, B. ekmaniana differs from B. arceutina by the colourless to pale excipular rim, colourless epihymenium and wider ascospores with more septation (
South Korea, North Gyeongsang Province, Bonghwa, Chunyang-myeon, Mt. Munsu, 36°59.28'N, 128°48.17'E, 1,058 m alt., on bark of Quercus mongolica, 29 August 2019, B.G. Lee 2019-000072 (KBA-L-0000072); South Korea, South Jeolla Province, Gokseong, Jukgok-myeon, Taeansa Temple, 35°08.06'N, 127°23.26'E, 297 m alt., on bark of Salix pierotii Miq., 25 May 2020, B.G. Lee 2020-000212, with Bacidia schweinitzii (KBA-L-0000412); same locality, on bark of Salix pierotii, 25 May 2020, B.G. Lee 2020-000227, with Bacidia schweinitzii, Coenogonium pineti (Ach.) Lücking & Lumbsch, Phaeophyscia rubropulchra (Degel.) Moberg, Porina melanops Malme (KBA-L-0000427); same locality, on bark of Idesia polycarpa Maxim., 25 May 2020, B.G. Lee 2020-000231, with Bacidia schweinitzii, Porina aff. melanops (KBA-L-0000431); same locality, on bark of Idesia polycarpa, 25 May 2020, B.G. Lee 2020-000232 (KBA-L-0000432); same locality, on bark of Taxicodendron vernicifluum (Stokes) F. A. Barkley, 25 May 2020, B.G. Lee 2020-000233, with Biatora aff. pacifica, Lecidea sp., Phaeophyscia rubropulchra, Rinodina sp., Traponora varians (Ach.) J. Kalb & Kalb (KBA-L-0000433); South Korea, North Gyeongsang Province, Bonghwa, Chunyang-myeon, Mt. Okseok, 37°00.91'N, 128°46.65'E, 1,085 m alt., on bark of Quercus mongolica, 15 September 2020, B.G. Lee & H.J. Lee 2020-001159, with Anisomeridium polypori (Ellis & Everh.) M.E. Barr, Bacidia schweinitzii, Rinodina sp. (KBA-L-0001359); same locality, on bark of Quercus mongolica, 15 September 2020, B.G. Lee & H.J. Lee 2020-001162, with Rinodina sp. (KBA-L-0001362); South Korea, North Jeolla Province, Jangsu, Mt. Youngchui, 35°38.59'N, 127°37.00'E, 907 m alt., on bark of Carpinus tschonoskii Maxim., 08 June 2021, B.G. Lee & H.J. Lee 2021-000563, with Lecanora megalocheila (Hue) H. Miyaw., Rinodina orientalis Sheard (KBA-L-0002035); same locality, on bark of Carpinus tschonoskii, 08 June 2021, B.G. Lee & H.J. Lee 2021-000565, with Arthonia apatetica (A. Massal.) Th. Fr., Lecidella euphorea (Flörke) Kremp. (KBA-L-0002037; GenBank ON352611 for ITS); same locality, on bark of Carpinus tschonoskii, 08 June 2021, B.G. Lee & H.J. Lee 2021-000569, with Anisomeridium polypori, Lecidella euphorea, Rinodina orientalis, Scoliciosporum sp. (KBA-L-0002041); same locality, on bark of Carpinus tschonoskii, 08 June 2021, B.G. Lee & H.J. Lee 2021-000573, with Arthonia apatetica, Lecanora aff. imshaugii Brodo, Lecidella euphorea, Porina hirsuta (KBA-L-0002045); South Korea, North Jeolla Province, Jangsu, Mt. Jangan, 35°38.58'N, 127°36.96'E, 925 m alt., on bark of Carpinus tschonoskii, 09 June 2021, B.G. Lee & H.J. Lee 2021-000759 (KBA-L-0002231); same locality, on bark of Carpinus tschonoskii, 09 June 2021, B.G. Lee & H.J. Lee 2021-000760, with Phaeophyscia adiastola (Essl.) Essl., Porina hirsuta, Rinodina orientalis, Scoliciosporum chlorococcum (Graewe ex Stenh.) Vězda (KBA-L-0002232); same locality, on bark of Carpinus tschonoskii, 09 June 2021, B.G. Lee & H.J. Lee 2021-000766, with Lecania sp., Phaeophyscia sp., Rinodina orientalis (KBA-L-0002238); South Korea, North Jeolla Province, Jangsu, Mt. Baegun, 35°36.76'N, 127°36.85'E, 661 m alt., on bark of Cornus walteri Wangerin, 10 June 2021, B.G. Lee & H.J. Lee 2021-000926 (KBA-L-0002398); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000927 (KBA-L-0002399); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000928 (KBA-L-0002400); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000929, with Phaeophyscia adiastola (KBA-L-0002401); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000930, with Phaeophyscia rubropulchra (KBA-L-0002402); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000931, with Lecanora sp., Phaeophyscia adiastola (KBA-L-0002403); same locality, on bark of Cornus walteri, 10 June 2021, B.G. Lee & H.J. Lee 2021-000932 (KBA-L-0002404).
Thallus corticolous, crustose, thin, little developed or indistinct, generally not continuous, minutely granular with contiguous granules when developed, pale grey with slightly brownish colour, margin indeterminate. Prothallus not detected.
Apothecia consistently flat or convex when mature, marginate, without pruina, 0.1–0.5 mm diam. (mean = 0.23, SD = 0.07, n = 107). Disc biatorine, without thalline exciple, pale pink to pale yellow when young and darker (particularly around margin) when mature. Proper exciple hyaline with or without pale brown pigment, the pigment slightly thicker close to hymenium or excipular rim, 40–50 μm wide laterally. Epihymenium bluish-green, ca. 5 μm high. Hymenium hyaline, 40–45 μm high. Hypothecium hyaline, 50–60 μm high; upper hypothecium paraplectenchymatous (globular to angular), lower hypothecium prosoplectenchymatous (periclinally or irregularly arranged). Crystals or oil droplets absent. Asci narrowly clavate, 8-spored, 39–41 × 10–12 μm (n = 3). Ascospores acicular to filiform, 3- or 7-septate, 28–38 × 1.5–2.5 μm (n = 14). Pycnidia not detected.
Epihymenium K–, C–. Hymenium K– or a few undeveloped asci K+ purplish. No lichen substance was detected by TLC.
Bacidia friesiana is similar to B. circumspecta (Norrl. & Nyl.) Malme and B. igniarii (Nyl.) Oxner (syn. Scutula igniarii (Nyl.) S. Ekman) in having epihymenium with green pigments, proper exciple without crystals and dark hypothecium amongst corticolous species. However, B. friesiana differs from the latter two by the excluded margin of apothecia and acicular ascospores. The latter species have a permanent margin of apothecia and bacilliform or clavate ascospores (
Phylogenetic analysis resulted in B. friesiana of Korea (ON352609 and ON352610) being nested with the sequences of Russia (MH539765), supported by a bootstrap value of 100 and a posterior probability of 1.00 for the branch (Fig.
South Korea, Gangwon Province, Yanggu, Nam-myeon, Dumu-ri, nearby a forested wetland, 38°02.12'N, 128°05.14'E, 421 m alt., on bark of Salix pierotii, 28 April 2020, B.G. Lee 2020-000164, with Mikhtomia gordejevii, Candelaria concolor (Dicks.) Arnold, Phaeophyscia adiastola, Porina cf. melanops, Rinodina cf. subminuta (KBA-L-0000364); South Korea, Gyeonggi Province, Yangpyeong, Cheongun-myeon, Dowon-ri, a forested wetland, 37°32.55'N, 127°48.60'E, 443 m alt., on bark of Salix pierotii, 31 May 2021, B.G. Lee & H.J. Lee 2021-000438, with Lecidella euphorea, Phaeophyscia adiastola, Rinodina orientalis (KBA-L-0001910; GenBank ON352609 for ITS); same locality, on bark of Aralia elata (Miq.) Seem., 31 May 2021, B.G. Lee & H.J. Lee 2021-000440, with Lecidella euphorea, Phaeophyscia adiastola, Traponora varians (KBA-L-0001912); same locality, on bark of Aralia elata, 31 May 2021, B.G. Lee & H.J. Lee 2021-000441, with Hyperphyscia adglutinata (Flörke) H. Mayrhofer & Poelt, Rinodina orientalis (KBA-L-0001913; GenBank ON352610 for ITS); same locality, on bark of Aralia elata, 31 May 2021, B.G. Lee & H.J. Lee 2021-000442, with Rinodina orientalis, Traponora varians (KBA-L-0001914); same locality, on bark of Aralia elata, 31 May 2021, B.G. Lee & H.J. Lee 2021-000443, with Hyperphyscia adglutinata, Rinodina orientalis, Traponora varians (KBA-L-0001915); same locality, on bark of Aralia elata, 31 May 2021, B.G. Lee & H.J. Lee 2021-000444, with Phaeophyscia adiastola, P. rubropulchra, Rinodina orientalis (KBA-L-0001916); same locality, on bark of Aralia elata, 31 May 2021, B.G. Lee & H.J. Lee 2021-000445 (KBA-L-0001917).
Thallus corticolous, crustose, continuous, wrinkled, or warted, pale yellowish-grey, margin indeterminate or determinate. Prothallus generally not present or locally present as blackish bordering a different lichen.
Apothecia flat, marginate, without pruina, 0.2–0.6 mm diam. (mean = 0.33, SD = 0.11, n = 72). Disc lecideine, without thalline exciple, blackish or reddish-black. Proper exciple hyaline with pale brown pigment dispersed, pigment slightly thicker close to hymenium, 80–100 μm wide laterally. Epihymenium brown to dark brown, ca. 10 μm high. Hymenium hyaline, 80–95 μm high. Hypothecium hyaline, 80–120 μm high, with a little pale yellow pigment. Crystals or oil droplets absent. Asci narrowly clavate to cylindrical, 8-spored, 42–48 × 12–13 μm (n = 3). Ascospores acicular to filiform, 9- or 10-septate, 36–67 × 2.5–4 μm (n = 11). Pycnidia not detected.
Epihymenium K+ purple or intensifying, extending to excipular rim. No lichen substance was detected by TLC.
Bacidia heterochroa is the most similar to B. laurocerasi (Delise ex Duby) Zahlbr. in having smooth thallus without granules, absence of crystals in exciple, epihymenium without green pigments, pale to colourless hypothecium, K+ purple in apothecial section and narrow ascospores less than 4 μm wide amongst corticolous species. However, B. heterochroa differs from B. laurocerasi by distinctly brown-pigmented paraphysial tips, less than 16-septate ascospores which are shorter but wider (less than 80 μm long but over 3.5 μm wide) and substrate preference to deciduous trees or shrubs (
Phylogenetic analysis resulted in B. heterochroa of Korea (ON352606, ON352612 and ON352613) being nested in a sister clade to B. laurocerasi, supported by a bootstrap value of 75 without a posterior probability as the Maximum Likelihood analysis did not match with the Bayesian Inference for the clade. The sequences of B. heterochroa were not compared with previous records due to the lack of data (Fig.
South Korea, Gangwon Province, Yanggu, Nam-myeon, Dumu-ri, a forested wetland, 38°02.12'N, 128°05.14'E, 421 m alt., on bark of Salix koriyanagi Kimura ex Goerz, 28 April 2020, B.G. Lee 2020-000186 (KBA-L-0000386; GenBank ON352606 for ITS); South Korea, South Jeolla Province, Damyang, Changpyeong-myeon, Oedong-ri, a forested wetland, 35°12.00'N, 127°00.88'E, 338 m alt., on bark of Fraxinus rhynchophylla Hance, 12 May 2021, B.G. Lee & D.Y. Kim 2021-000214 (KBA-L-0001686); South Korea, Gangwon Province, Jeongseon, Imgye-myeon, Gamok-ri, a forested wetland, 37°32.47'N, 128°57.72'E, 760 m alt., on bark of Acer tartaricum subsp. ginnala (Maxim.) Wesm., 17 June 2021, B.G. Lee & H.J. Lee 2021-001241, with Lecanora chionocarpa Hue (KBA-L-0002713); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001242, with Phaeophyscia adiastola (KBA-L-0002714); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001255, with Opeltia flavorubescens, Phaeophyscia adiastola (KBA-L-0002727; GenBank ON352612 for ITS); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001257, with Hyperphyscia adglutinata, Lecidella euphorea (KBA-L-0002729); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001262, with Lecidella euphorea, Phaeophyscia adiastola, Rinodina orientalis (KBA-L-0002734; GenBank ON352613 for ITS); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001263, with Opeltia flavorubescens, Phaeophyscia adiastola, Rinodina orientalis (KBA-L-0002735); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001267, with Lecidella euphorea, Porina hirsuta, Rinodina orientalis, Straminella varia (KBA-L-0002739); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001269, with Lecidella euphorea, Opeltia flavorubescens, Phaeophyscia rubropulchra, Rinodina orientalis (KBA-L-0002741).
Thallus corticolous, crustose, continuous, wrinkled, warted or subsquamulose, often granular locally, whitish pale grey. Prothallus generally not present or present as dark brown to black between different colonies.
Apothecia flat, marginate, with a little or heavy white pruina, generally more pruinose at margin, 0.3–1.7 mm diam. (mean = 0.75, SD = 0.28, n = 116). Disc lecideine, without thalline exciple, brown to dark brown. Proper exciple with radiating clusters of crystals produced around hypothecium and expanding to excipular rim and finally shown as pruina on surface, hyaline downwards but brown around rim, the brown concolorous or slightly paler to epihymenium, 80–100 μm wide laterally. Epihymenium brown to dark brown, ca. 10 μm high, with pruina (ca. 10 μm high) on surface. Hymenium hyaline, 70–80 μm high. Hypothecium hyaline, 80–100 μm high. Other small crystals present a few in upper hypothecium. Oil droplets absent. Asci cylindrical, 8-spored, 65–75 × 10–16 μm (n = 7). Ascospores acicular to filiform, up to 13-septate, 45–70 × 2.5–4.5 μm (n = 10). Pycnidia not detected.
Thallus K+ yellow, KC–, C–, Pd–, UV–. Epihymenium K–. Atranorin was detected by TLC.
Bacidia suffusa is the most similar to B. russeola (Kremp.) Zahlbr. in having dark apothecia, generally colourless epihymenium without green pigment, long ascospores with the L/W ratio over 11, pale or colourless hypothecium and K+ purple reaction on epihymenium and nearby excipular rim amongst corticolous species. However, B. suffusa differs from B. russeola by the presence of pruina on the disc and in proper exciple as radiating clusters of crystals and more than 10-septate ascospores (
Phylogenetic analysis resulted in B. suffusa of Korea (ON352605, ON352614, ON352615 and ON352616) being nested in a sister clade of the sequences of Pakistan (MW728313 and MW788561), Russia (MH048615, MH048616 and MH048617) or U.S.A. (MH048618 and MH048619). The molecular data of Korea converged into the previous data of B. suffusa, supported by a bootstrap value of 100 and a posterior probability of 1.00 for the branch (Fig.
South Korea, Gangwon Province, Yanggu, Nam-myeon, Dumu-ri, a forested wetland, 38°02.12'N, 128°05.14'E, 421 m alt., on bark of Salix pierotii Miq., 28 April 2020, B.G. Lee 2020-000158 (KBA-L-0000358); same locality, on bark of Salix pierotii, 28 April 2020, B.G. Lee 2020-000159 (KBA-L-0000359; GenBank ON352605 for ITS); same locality, on bark of Salix pierotii, 28 April 2020, B.G. Lee 2020-000168, with Candelaria concolor, Phaeophyscia adiastola, Phaeophyscia hirtuosa (Kremp.) Essl. (KBA-L-0000368); South Korea, Gangwon Province, Gangneung, Okgye-myeon, Mt. Seokbyung, 37°34.45'N, 128°55.01'E, 271 m alt., on bark of Acer pictum var. mono, 17 June 2020, B.G. Lee & H.J. Lee 2020-000799 (KBA-L-0000999); South Korea, Gangwon Province, Jeongseon, Imgye-myeon, Gamok-ri, a forested wetland, 37°32.47'N, 128°57.72'E, 760 m alt., on bark of Fraxinus chiisanensis Nakai, 17 June 2021, B.G. Lee & H.J. Lee 2021-001304, with Normandina pulchella (Borrer) Nyl., Phaeophyscia sp. (KBA-L-0002776; GenBank ON352614 for ITS); same locality, on bark of Fraxinus chiisanensis, 17 June 2021, B.G. Lee & H.J. Lee 2021-001305, with Anisomeridium polypori, Normandina pulchella, Phaeophyscia sp., Porina hirsuta (KBA-L-0002777); same locality, on bark of Fraxinus chiisanensis, 17 June 2021, B.G. Lee & H.J. Lee 2021-001306, with Normandina pulchella, Opeltia flavorubescens, Phaeophyscia adiastola (Essl.) Essl. (KBA-L-0002778; GenBank ON352615 for ITS); same locality, on bark of Fraxinus chiisanensis, 17 June 2021, B.G. Lee & H.J. Lee 2021-001308, with Phaeophyscia adiastola (KBA-L-0002780); same locality, on bark of Fraxinus chiisanensis, 17 June 2021, B.G. Lee & H.J. Lee 2021-001320, with Opeltia flavorubescens (KBA-L-0002792); same locality, on bark of Acer tartaricum subsp. ginnala, 17 June 2021, B.G. Lee & H.J. Lee 2021-001363 (KBA-L-0002835; GenBank ON352616 for ITS).
The key is composed of all 19 species in the genus Bacidia of Korea, including synonyms in Bacidina and Toniniopsis species.
1 | Epihymenium with green pigment | 2 |
– | Epihymenium colourless, yellow-brown, brown to dark brown, but without green pigment | 5 |
2 | Proper exciple with radiating clusters of coarse crystals (up to 7 μm wide); hymenium ca. 100 μm high; ascospores 40–68 × 2.5–3 μm; atranorin present | B. schweinitzii |
– | Proper exciple without crystals; hymenium less than 70 μm high; ascospores less than 50 μm long; without substance | 3 |
3 | Hypothecium colourless to pale blue-green; thallus pale grey to pale brown-grey without green colour | B. friesiana |
– | Hypothecium colourless to brown, dark red-brown; thallus grey-green to green-brown | 4 |
4 | Proper exciple with green pigment at rim, pale to colourless downwards; hypothecium K– or K+ green-brown; generally on rock or occasionally on bark or moss | B. egenula (Bacidina egenula) |
– | Proper exciple colourless at rim, red-brown to black-brown downwards; hypothecium K+ purple; on bark | B. subincompta (Toniniopsis subincompta) |
5 | On rock | 6 |
– | On bark or wood | 12 |
6 | Apothecia pruinose | 7 |
– | Apothecia not pruinose | 8 |
7 | Thallus coarsely granular without forming soredia; apothecia 0.7–1.2 mm diam.; hymenium 70–100 μm high; hypothecium colourless to pale yellow or pale orange; ascospores 40–70 × 2.5–3 μm, 3- to 7-septate | B. rubella |
– | Thallus granular with soredia; apothecia 0.3–0.7 mm diam.; hymenium 40–50 μm high; hypothecium orange-brown to dark red-brown; ascospores 24–46 × 1–2 μm, 1- to 3-septate | B. arnoldiana (Bacidina arnoldiana) |
8 | Disc brown, red-brown to black; hypothecium pale brown to dark brown | 9 |
– | Disc pale yellow, pale orange to dark brown; hypothecium colourless to pale yellow or pale orange | 10 |
9 | Proper exciple dark coloured; ascospores 25–35 × 6–10 μm, with L/W ratio less than 10 | B. hakonensis |
– | Proper exciple colourless to pale brown; ascospores 24–46 × 1–2 μm, with L/W ratio over 10 | B. arnoldiana (Bacidina arnoldiana) |
10 | Thallus rimose, wrinkled or warted, but not granular; disc pale yellow or pale grey; epihymenium K– | B. chloroticula (Bacidina chloroticula) |
– | Thallus granular; disc pale to dark brown; epihymenium K+ purple | 11 |
11 | Thallus granular forming isidia- or coral-like structures; prothallus absent; apothecia flat; ascospores 25–34 × 1.1–1.9 μm; occasionally on old wood | B. egenuloidea (Bacidina egenuloidea) |
– | Thallus granular-warted; white prothallus present on border; apothecia flat to convex; ascospores 24–43 × 2–2.5 μm | B. inundata (Bacidina inundata) |
12 | On wood. Thallus granular forming isidia- or coral-like structures; disc pale orange to dark purple-brown; proper exciple orange-brown to brown at rim; on old wood, but generally on rock | B. egenuloidea (Bacidina egenuloidea) |
– | On bark | 13 |
13 | Proper exciple with radiating clusters of crystals; white pruina present; atranorin present as a major compound or a trace | 14 |
– | Proper exciple without crystals; pruina absent; without substance | 17 |
14 | Hypothecium brown-orange to dark brown; apothecial section K+ purple-red | B. polychroa |
– | Hypothecium colourless to pale yellow or pale orange; apothecial section K– | 15 |
15 | Thallus generally coarsely granular, pale grey to green-grey; prothallus white to pale grey when present; ascospores up to 9-septate | B. rubella |
– | Thallus smooth, wrinkled, warted or granular locally, white-grey to grey; prothallus absent; ascospores up to 13-septate | 16 |
16 | Thallus grey; disc not pruinose generally, but sometimes white-pruinose; proper exciple with radiating clusters of minute crystals (ca. 0.5 μm wide); epihymenium without distinct colour; ascospores 50–85 × 2.6–3.4 μm | B. fraxinea |
– | Thallus whitish pale grey; disc light to heavily pruinose; proper exciple with radiating clusters of coarse crystals (up to 10 μm wide); epihymenium brown to dark brown; ascospores 45–70 × 2.5–4.5 μm | B. suffusa |
17 | Thallus granular with soredia-like goniocysts | 18 |
– | Thallus smooth, wrinkled, warted or rarely granular, but without soredia | 19 |
18 | Hypothecium colourless; conidia curved without hook | B. delicata (Bacidina delicata) |
– | Hypothecium orange-brown to dark red-brown; conidia hooked | B. sulphurella (Bacidina sulphurella) |
19 | Disc purple-brown to black or slightly blackish when mature; epihymenium K+ purple | 20 |
– | Disc pale yellow, pale grey or pale brown; epihymenium K– | 22 |
20 | Proper exciple colourless to pale yellow at rim; thallus olive-green; apothecia generally pale yellow to pale orange with slightly blackish pigment; epihymenium colourless with a little pale yellow-brown pigment | B. fuscopallida |
– | Proper exciple dark brown to black-brown at rim; thallus white to pale grey; apothecia purple-brown to black; epihymenium brown to dark brown | 21 |
21 | Brown pigment of epihymenium deposited in caps of paraphysial tips; thallus wrinkled or warted, but not squamulose; prothallus blackish on border when present; ascospores 32–67 × 2.5–4.5 μm, 3- to 15-septate | B. heterochroa |
– | Brown pigment of epihymenium distributed in upper hymenial jelly; thallus wrinkled or warted, sometimes squamulose to varnish-like crust; prothallus white between areoles; ascospores 45–80 × 2–3.5 μm, 7- to 28-septate | B. laurocerasi |
22 | Thallus rimose, wrinkled or warted; apothecia ca. 0.2 mm diam.; hypothecium colourless; ascospores 24–28 × 1–1.2 μm, 0- to 3-septate; occasionally on rock | B. chloroticula (Bacidina chloroticula) |
– | Thallus granular to smooth; apothecia 0.4–1.4 mm diam.; hypothecium straw, yellow-brown to red-brown; ascospores 45–70 × 1.5–4 μm, 3- to 15-septate | 23 |
23 | Proper exciple yellow-brown to brown at rim; epihymenium yellow-brown; ascospores 1.5–2.5 μm wide, 3- to 7-septate | B. arceutina |
– | Proper exciple colourless to pale yellow at rim; epihymenium colourless; ascospores 2–4.5 μm wide, 3- to 15-septate | B. ekmaniana |
This work was supported by a grant from the Korean Forest Service Program through the Korea National Arboretum (KNA-202003127AF-00) for the forested wetland conservation of Korea.