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Research Article
Two new Rinodina lichens from South Korea, with an updated key to the species of Rinodina in the far eastern Asia
expand article infoBeeyoung Gun Lee, Jae-Seoun Hur§
‡ Baekdudaegan National Arboretum, Bonghwa, Republic of Korea
§ Sunchon National University, Suncheon, Republic of Korea
Open Access

Abstract

Rinodina salicis Lee & Hur and Rinodina zeorina Lee & Hur are described as new lichen-forming fungi from forested wetlands or a humid forest in South Korea. Rinodina salicis is distinguishable from Rinodina excrescens Vain., the most similar species, by its olive-gray thallus with smaller areoles without having blastidia, contiguous apothecia, non-pruinose discs, paler disc color, wider ascospores in the Pachysporaria-type II, and the absence of secondary metabolites. Rinodina zeorina differs from Rinodina hypobadia Sheard by areolate and brownish thallus, non-pruinose apothecia, colorless and wider parathecium, narrower paraphyses with non-pigmented and unswollen tips, longer and narrower ascospores with angular to globose lumina, and the absence of pannarin. Molecular analyses employing internal transcribed spacer (ITS) sequences strongly support the two new species to be unique in the genus Rinodina. An updated key is provided to assist in the identification of all 63 taxa in Rinodina of the far eastern Asia.

Keywords

Biodiversity, corticolous, phylogeny, Physciaceae, taxonomy

Introduction

Rinodina, the largest genus in the family Physciaceae, comprises about three hundred species worldwide (Sheard et al. 2017; Wijayawardene et al. 2020). Several infrageneric groups have been studied since Malme (1902) introduced the ascospore-type concept for the groups in Rinodina (Poelt 1965; Grube and Arup 2001). Although the classification based on different ascospore types has been coarsely accepted, the variety of ascospores does not always correspond to the infrageneric classification. As the pattern of ascospore ontogeny is considered more important than the spore type itself, it is understood that the ascospore types should be respected in developmental stages of a spore (Giralt 1994; Grube and Arup 2001; Sheard 2010; Resl et al. 2016).

The Rinodina has been studied in Europe (Mayrhofer and Poelt 1979; Giralt et al. 1995; Giralt 2001; Mayrhofer and Moberg 2002), North America (Sheard and Mayrhofer 2002; Sheard 2004, 2010, 2018; Sheard et al. 2011, 2012; Lendemer et al. 2012, 2019; Morse and Sheard 2020), islands of South America (Bungartz et al. 2016), Australia to New Zealand (Mayrhofer 1983, 1984b; Kaschik 2006; Elix 2011; Elix et al. 2020), Asia to Russian Far East (Mayrhofer 1984a; Galanina et al. 2011; Lendemer et al. 2012; Sheard et al. 2017; Galanina et al. 2018; Galanina and Ezhkin 2019; Zheng and Ren 2020; Galanina et al. 2021; Kumar et al. 2021), and South Africa (Matzer and Mayrhofer 1996; Mayrhofer et al. 2014). Molecular works have been accomplished over the continents (Grube and Arup 2001; Wedin et al. 2002; Nadyeina et al. 2010; Resl et al. 2016).

Sheard et al. (2017) achieved the first and comprehensive study on the genus Rinodina of the far eastern Asia (Korea, Japan, and Russian Far East). Several studies announced further more species in the genus, such as R. badiexcipula Sheard, R. convexula H. Magn., R. occulta (Körb.) Sheard, R. oxneriana S.Y. Kondr., Lőkös & Hur and R. tephraspis (Tuck.) Herre from South Korea (Kondratyuk et al. 2016, 2017; Yakovchenko 2018; Kondratyuk et al. 2020) and R. colobinoides (Nyl.) Müll. Arg., R. herrei H. Magn., R. laevigata (Ach.) Malme, and R. parasitica H. Mayrhofer & Poelt from the Kuril Islands and the Magadan region, Russian Far East (Galanina and Ezhkin 2019; Galanina et al. 2021). Among them, R. oxneriana was discovered as a new species and other eight species were reported as new records to the far eastern Asia. The species of Rinodina in the far eastern Asia are mainly corticolous and the main genera of the substrate trees are Quercus, Picea, Salix, Betula and Alnus (Fig. 1) (Lendemer et al. 2012; Sheard et al. 2012; Joshi et al. 2013; Kondratyuk et al. 2013, 2016, 2017, 2020; Aptroot and Moon 2014; Sheard et al. 2017; Yakovchenko et al. 2018; Galanina and Ezhkin 2019; Gananina et al. 2021). Those main substrates vigorously grow in a humid forest, a valley or a wetland, and particularly the genera Salix and Alnus often inhabit the water. Inhabiting those tree barks, diverse Rinodina species are easily detected in shaded forests and forested wetlands in which are one of the representative lichens of the ecosystems.

Figure 1. 

Substrates of Rinodina species in the far eastern Asia. Rinodina species of the far eastern Asia occur mainly on bark, and the genera Quercus, Picea, Salix, Betula and Alnus are the main substrates for corticolous Rinodina species of the far eastern Asia.

This study describes two new lichen-forming fungi in the genus Rinodina. Field surveys for the lichen biodiversity in the forested wetlands of South Korea were carried out during the summer of 2020, and a couple of specimens of Rinodina were collected from barks of Quercus and Salix, the most common genera of the substrates for corticolous Rinodina species in the far eastern Asia, in a humid forest and a forested wetland on mountains (Fig. 2). The specimens were comprehensively analyzed in ecology, morphology, chemistry and molecular phylogeny and did not correspond to any previously known species. We describe them as new species, Rinodina salicis and R. zeorina, and this discovery contributes to the taxonomy with overall 63 taxa in the genus Rinodina of the far eastern Asia. The type specimens are deposited in the herbarium of the Baekdudaegan National Arboretum (KBA, the herbarium acronym in the Index Herbariorum), South Korea.

Figure 2. 

Specific collection sites for two new species A habitat/landscape for R. salicis B habitat/landscape for R. zeorina C location for R. salicis (a black star); locations for R. zeorina (two black diamonds).

Materials and methods

Morphological and chemical analyses

Hand sections were prepared manually with a razor blade under a stereomicroscope (Olympus optical SZ51; Olympus, Tokyo, Japan), scrutinized under a compound microscope (Nikon Eclipse E400; Nikon, Tokyo, Japan) and pictured using a software program (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 systems A and C according to standard methods (Orange et al. 2001).

Isolation, DNA extraction, amplification, and sequencing

Hand-cut sections of ten to twenty ascomata per collected specimen were prepared for DNA isolation and DNA was extracted with a NucleoSpin Plant II Kit in line with the manufacturer’s instructions (Macherey-Nagel, Düren, Germany). PCR amplifications for the internal transcribed spacer region (ITS1-5.8S-ITS2 rDNA) RNA genes were achieved using Bioneer’s AccuPower PCR Premix (Bioneer, Daejeon, Korea) in 20-μl tubes with 16 μl of distilled water, 2 μl of DNA extracts and 2 μl of the primers ITS5 and ITS4 (White et al. 1990). 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). The annealing temperature was occasionally altered by ±1 degree in order to get a better result. PCR purification and DNA sequencing were accomplished by the genomic research company Macrogen (Seoul, Korea).

Phylogenetic analyses

All ITS sequences (Table 1) were aligned and edited manually using ClustalW in Bioedit V7.2.6.1 (Hall 1999). All missing and ambiguously aligned data and parsimony-uninformative positions were removed and only parsimony-informative regions were finally analyzed in MEGA X (Stecher et al. 2020). The final alignment comprised 974 bp in which 167 variable regions were detected. The phylogenetically informative regions were 523. Phylogenetic trees with bootstrap values were obtained in RAxML GUI 2.0 beta (Edler et al. 2019) using the maximum likelihood method with a rapid bootstrap with 1000 bootstrap replications and GTR GAMMA for the substitution matrix. The posterior probabilities were obtained in BEAST 2.6.4 (Bouckaert et al. 2019) using the GTR 121343 model, 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 1000 log parameters. Then, a consensus tree was constructed in TreeAnnotator 2.6.4 (Bouckaert et al. 2019) with no discard of burnin, 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. The bootstrapping and posterior probability analyses were repeated three times for the result consistency and no significant differences were shown for the tree shapes and branch values. The phylogenetic trees and DNA sequence alignments are deposited in TreeBASE under the study ID 28192. Overall analyses in the materials and methods were accomplished based on Lee and Hur (2020).

Table 1.

Species list and DNA sequence information employed for phylogenetic analysis.

No. Species ID (ITS) Voucher
1 Amandinea lignicola JX878521 Tønsberg 36426 (BG)
2 Amandinea punctata HQ650627 AFTOL-ID 1306
3 Buellia badia MG250192 TS1767 (LCU)
4 Buellia boseongensis MF399000 KoLRI 041680
5 Buellia numerosa LC153799 CBM:Watanuki:L01034
6 Rinodina afghanica MT260860 500103 (XJU-L)
7 Rinodina alba GU553290 GZU 000272655
8 Rinodina albana GU553297 GZU 000272651
9 Rinodina anomala MN587028 Sipman 62934
10 Rinodina archaea DQ849292 H. Mayrhofer 15752 (GZU)
11 Rinodina atrocinerea AF540544 H. Mayrhofer 13.740 & U. Arup (GZU)
12 Rinodina balanina KY266842 O-L-195705
13 Rinodina bischoffii DQ849291 M. Lambauer 0031 (GZU)
14 Rinodina cacaotina DQ849295 H. Mayrhofer 10770 (HO)
15 Rinodina calcarea GU553292 GZU 000272654
16 Rinodina cana MN587029 Sipman 63008
17 Rinodina capensis DQ849296 W. Obermayer 09230 (GZU)
18 Rinodina confragosa DQ849297 W. Obermayer 09091 (GZU)
19 Rinodina confragosula DQ849298 M. Lambauer 0044 (GZU)
20 Rinodina degeliana KX015681 Tønsberg 42631
21 Rinodina destituta KT695382 BIOUG24047-H02
22 Rinodina disjuncta MK812529 TRH-L-15387
23 Rinodina efflorescens KX015683 Malicek 5462
24 Rinodina exigua GU553294 GZU 000272652
25 Rinodina gallowayi DQ849299 M. Lambauer 0125 (GZU)
26 Rinodina gennarii AJ544187 B44435
27 Rinodina glauca GU553295 GZU 000272662
28 Rinodina herteliana DQ849300 M. Lambauer 0177 (GZU)
29 Rinodina immersa DQ849301 M. Lambauer 0129 (GZU)
30 Rinodina interpolata AF250809 M263
31 Rinodina jamesii DQ849303 H. Mayrhofer 10810 (GZU)
32 Rinodina lecanorina AF540545 H. Mayrhofer 13.120 (GZU)
33 Rinodina lepida AY143413 Trinkaus 137
34 Rinodina luridata DQ849304 H. Mayrhofer 12122 (GZU)
35 Rinodina luridescens AJ544183 B42835
36 Rinodina metaboliza MT260864 20080224 (XJU-L)
37 Rinodina milvina GU553299 KW 63379
38 Rinodina mniaroea KX015689 Spribille 20101 (GZU)
39 Rinodina mniaroea KX015691 V. Wagner, 15.07.06/1 (GZU)
40 Rinodina mniaroea KX015692 Spribille 20391 (GZU)
41 Rinodina moziana DQ849307 H. Mayrhofer 6729 (GZU)
42 Rinodina moziana var. moziana DQ849305 M. Lambauer 0214 (GZU)
43 Rinodina nimisii AJ544184 B42685
44 Rinodina obnascens AJ544185 B42477
45 Rinodina oleae DQ849308 M. Lambauer 0178 (GZU)
46 Rinodina oleae GU553301 GZU 000272565
47 Rinodina olivaceobrunnea AF540547 J. Romeike 2.090300 (GOET)
48 Rinodina orculata DQ849309 H. Mayrhofer 15754 (GZU)
49 Rinodina orientalis MW832807 BDNA-L-0000284
50 Rinodina orientalis MW832808 BDNA-L-0000653
51 Rinodina orientalis MW832809 BDNA-L-0000774
52 Rinodina oxydata DQ849313 H. Mayrhofer 11406 (GZU)
53 Rinodina plana AF250812 E34
54 Rinodina pyrina AF540549 P. Bilovitz & H. Mayrhofer 483 (GZU)
55 Rinodina ramboldii DQ849315 G. Rambold 5094 (M)
56 Rinodina reagens DQ849316 M. Lambauer 0218 (GZU)
57 Rinodina roboris MK811851 O-L-206765
58 Rinodina roscida DQ849317 S. Kholod plot515 (GZU)
59 Rinodina salicis MW832810 BDNA-L-0000558
60 Rinodina salicis MW832811 BDNA-L-0000560
61 Rinodina septentrionalis GU553303 GZU 000272561
62 Rinodina sheardii MK778639 J. Malicek 10238
63 Rinodina sheardii MK778640 J. Vondrak 15298 (PRA)
64 Rinodina sophodes AF540550 P. Bilovitz 968 (GZU)
65 Rinodina teichophila GU553305 GZU 000272659
66 Rinodina trevisanii KX015684 de Bruyn s.n. 2011 (GZU)
67 Rinodina tunicata AF540551 H. Mayrhofer 13.749 & R. Ertl (GZU)
68 Rinodina turfacea AF224362 Moberg 10422
69 Rinodina vezdae DQ849318 H. Mayrhofer 15757 (GZU)
70 Rinodina zeorina MW832812 BDNA-L-0000642
71 Rinodina zeorina MW832813 BDNA-L-0000646
72 Rinodina zeorina MW832814 BDNA-L-0000650
73 Rinodina zeorina MW832815 BDNA-L-0000651
74 Rinodina zeorina MW832816 BDNA-L-0000668
75 Rinodina zeorina MW832817 BDNA-L-0000933
76 Rinodina zwackhiana AF540552 H. Mayrhofer 13.848 (GZU)
77 Rinodinella controversa AF250814 M281
78 Rinodinella dubyanoides AF250815 E29
Overall 78

Results and discussion

Phylogenetic analyses

An independent phylogenetic tree for the genus Rinodina and related genera was produced from 67 sequences from GenBank and 11 newly generated sequences for the two new species and related species (Table 1). The two new species were positioned in the genus Rinodina in the ITS tree. The ITS tree describes that R. salicis, a new species, is coming alone in a single clade. Several species such as R. mniaroea (Ach.) Körb., R. roscida (Sommerf.) Arnold, R. bischoffii (Hepp) A. Massal., R. luridata (Körb.) H. Mayrhofer, Scheid. & Sheard, R. metaboliza Vain., R. albana (A. Massal.) A. Massal., R. afghanica M. Steiner & Poelt, R. zwackhiana (Kremp.) Körb., R. calcarea (Hepp ex Arnold) Arnold, R. immersa (Körb.) J. Steiner, R. tunicata H. Mayrhofer & Poelt, Rinodinella controversa (A. Massal.) H. Mayrhofer & Poelt, and R. dubyanoides (Hepp) H. Mayrhofer & Poelt, are situated close to the new species; this particular clade lacks statistical support (bootstrap value of 58 and a posterior probability of 0.75). Rinodina zeorina, the other new species, was located in a clade with R. sheardii Tønsberg, represented by a bootstrap value of 89 and a posterior probability of 0.88 (not shown) for the branch (Fig. 3).

Figure 3. 

Phylogenetic relationships among available species in the genus Rinodina based on a maximum likelihood analysis of the dataset of ITS sequences. The tree was rooted with the sequences of the genera Amandinea and Buellia. Maximum likelihood bootstrap values ≥ 70% and posterior probabilities ≥ 95% are shown above internal branches. Branches with bootstrap values ≥ 90% are shown as fatty lines. Two new species, R. salicis and R. zeorina are presented in bold as their DNA sequences were produced from this study. All species names are followed by the Genbank accession numbers.

Taxonomy

Rinodina salicis B.G. Lee & J.-S. Hur, sp. nov.

MycoBank No: MB839186
Fig. 4

Diagnosis

Rinodina salicis differs from R. excrescens by olive-gray thallus with smaller areoles without blastidia, contiguous apothecia, the absence of pruina on disc, paler disc color, wider ascospores in the Pachysporaria-type, and the absence of secondary metabolites.

Type

South Korea, Gangwon Province, Gangneung, Seongsan-myeon, Eoheul-ri, a forested wetland, 37°43.61'N, 128°48.13'E, 212 m alt., on bark of Salix koreensis Andersson, 02 June 2020, B.G.Lee & H.J.Lee 2020-000358 (holotype: BDNA-L-0000558; GenBank MW832810 for ITS); same locality, on bark of Salix koreensis, 02 June 2020, B.G.Lee & H.J.Lee 2020-000360, with Caloplaca gordejevii (Tomin) Oxner, Lecanora sp., and Phaeophyscia sp. (paratype: BDNA-L-0000560; GenBank MW832811 for ITS).

Thallus corticolous, crustose, minutely bullate, some developing to conglomerate and continuous, rarely lobulated, thin, grayish-green to olive green, margin indeterminate, vegetative propagules absent, areoles 0.1–0.2 mm diam., 85–90 μm thick; cortex hyaline, 10 μm thick, cortical cells 5–9 μm diam.; medulla 60–65 μm thick, intermixed with algal cells, without crystals (PL–); photobiont coccoid, cells globose, 5–15 μm. Prothallus absent.

Apothecia abundant, rounded, often contiguous, emerging on the surface of thallus and sessile when mature, constricted at the base, 0.2–1.3 mm diam. Disc flat, not pruinose, pale brown or dark brown from early stages, 220–260 μm thick; margin persistent, prominent, generally entire or somewhat flexuous, a little crenulate, thalline margin concolorous to thallus but proper margin near disc distinctly pale brown. Amphithecium well-developed, with small crystals in both cortical layer and the algal-containing medulla, crystals extending to the base, not dissolving in K, 60–70 μm wide laterally, algal layers continuous to the base or solitary, algal cells 5–15 μm diam., cortical layer hyaline, 10–20 μm thick. Parathecium hyaline but light brown at periphery, 45–50 μm wide laterally and 70–80 μm wide at periphery. Epihymenium brown, not granular, pigment slightly paler in K but not diluted, 5–10 μm high. Hymenium hyaline, 70–90 μm high. Hypothecium generally hyaline, with pale yellow pigment, prosoplectenchymatous (irregular), 70–80 μm high. Oil droplets are present mainly in hypothecium and a little in hymenium. Paraphyses septate, anastomosing, 1–1.5 μm wide, simple or branched at tips, tips swollen, pigmented, epihymenium pigmented by paraphysial tips, 4.5–7.5 μm wide. Asci clavate, 8-spored, 68–90 × 20–25 μm (n = 5). Ascospores ellipsoid, 1-septate, Pachysporaria-type II, rarely Physcia-type, Type A development, hyaline when young and light brown to brown in mature, 14–24 × 8–13.5 μm (mean = 18.2 × 10.5 μm; SD = 2.12(L), 1.19(W); L/W ratio 1.2–2.4, ratio mean = 1.7, ratio SD = 0.2; n = 105). Pycnidia not detected.

Figure 4. 

Rinodina salicis (BDNA-L-0000558, holotype) in morphology A–D habitus and apothecia. Thallus olive-gray composed of tiny areoles and non-pruinose apothecia E well-developed amphithecium and algal layer extending to the base F asci clavate with eight spores G ascospores simple in the beginning and developed 1-septate, Pachysporaria-type II, rarely Physcia-type at mature. Scale bars: 1 mm (A–D); 200 μm (E); 10 μm (F, G).

Chemistry

Thallus K–, KC–, C–, Pd–. Hymenium I+ purple-blue. UV–. No lichen substance was detected by TLC.

Distribution and ecology

The species occurs on the bark of Salix koreensis. The species is currently known from the type collections.

Etymology

The species epithet indicates the lichen’s substrate preference, namely the substrate tree Salix koreensis.

Notes

The new species is similar to R. excrescens and R. bullata Sheard & Lendemer in having bullate thallus. However, the new species differs from R. excrescens by olive-gray thallus with smaller areoles without having blastidia, contiguous apothecia, the absence of pruina on disc, paler disc color, ascospore type, larger ascospore, and the absence of secondary metabolites (Sheard 1966; Sheard et al. 2012).

The new species is closer to R. bullata in having small bullate areoles without having blastidia. However, the new species differs from the latter by olive-gray thallus, contiguous and larger apothecia, proper margin with pale brown color, crystals present in both cortex and medulla in amphithecium, larger ascospores, K– reaction on thallus, and the absence of lichen substance (Sheard et al. 2012, 2017).

The new species is comparable to R. granulans Vain. as the latter represents thallus with minute areoles. However, the new species differs from the latter by thallus color, slightly smaller areoles without blastidia, abundance of apothecia without pruina, Pachysporaria-type II ascospores, K– reaction on thallus, and the absence of lichen substance (Giralt et al. 1994; Galanina et al. 2011). Reference Table 2 provides the key characteristics distinguishing R. salicis from the compared species above.

Table 2.

Comparison of Rinodina salicis with closely-related species.

Species Rinodina salicis Rinodina bullata Rinodina excrescens Rinodina granulans
Thallus growth form bullate without blastidia bullate without blastidia bullate with blastidia bullate with blastidia, forming leprose crust
Areoles (mm in diam.) 0.1–0.2 0.1–0.15(–0.2) up to c. 1.98 (0.1–)0.2–0.3(–0.5)
Thallus color olive-gray light gray gray gray to gray-brown
Apothecia (mm in diam.) 0.2–1.3 0.3–0.6 up to c. 1.26 up to 0.3
Apothecia contiguation often contiguous not contiguous not contiguous not contiguous
Apothecia abundance abundant abundant abundant very rare
Pruina absent on disc often present on disc often present on disc
Disc color pale to dark brown brown brown to black reddish brown
Proper margin pale brown indistinct indistinct
Crystals in amphithecium present in medulla and cortex present in cortex present
Ascospore type Pachysporaria-type II Pachysporaria-type II Physcia-type Physcia-type to Milvina-type
Ascospores (μm) 14–24 × 8–13.5 14.5–16.5 × 8–9 17.5–19.5 × 8.5–9.5 18–25 × 10–14
Spot test thallus K– thallus K+ yellow thallus K– thallus K+ faint yellow
Substance absent atranorin pannarin, (rarely zeorin) pannarin
Reference BDNA-L-0000558 (holotype), BDNA-L-0000560 (paratype) Sheard et al. 2012, 2017 Sheard 1966; Sheard et al. 2012, 2017 Giralt et al. 1994; Galanina et al. 2011

Rinodina zeorina B.G. Lee & J.-S. Hur, sp. nov.

MycoBank No: MB839187
Fig. 5

Diagnosis

Rinodina zeorina differs from R. hypobadia by areolate, brownish thallus, apothecia without pruina, hyaline and wider parathecium, narrower paraphyses with hyaline and unswollen tips, longer and narrower ascospores with just angular to globose lumina, and the absence of pannarin.

Type

South Korea, North Gyeongsang Province, Bonghwa-gun, Seokpo-myeon, Mt. Cheongok, 37°01.89'N, 128°58.65'E, 1,104 m alt., on bark of Quercus mongolica, 16 June 2020, B.G. Lee & H.J. Lee 2020-000733, with Biatora sp., Lecidella euphorea (Flörke) Kremp., Pertusaria multipuncta (Turner) Nyl., and Sagiolechia sp. (holotype: BDNA-L-0000933; GenBank MW832817 for ITS).

Thallus corticolous, crustose, areolate, rimose to continuous, thin, light gray to light brownish gray, margin indeterminate or determinate with prothallus, vegetative propagules absent, 160–250 mm diam., 80–170 μm thick, areoles 0.1–0.5 mm diam.; cortex brown, 5–8 μm thick, with epinecral layer, hyaline, 3–7 μm thick; medulla 35–40 μm thick, intermixed with algal cells, without crystals (PL–); photobiont coccoid, cells globose, 5–9 μm. Prothallus absent or brownish black when present.

Figure 5. 

Rinodina zeorina (BDNA-L-0000933, holotype for A–G; BDNA-L-0000668 for H–K) in morphology A–C habitus and apothecia on bark of Quercus mongolica. Thallus brownish and areolate and non-pruinose apothecia D well-developed amphithecium and pigmented hypothecium E epihymenium with brown pigment which extending to the cortical layer of amphithecium. Parathecium light brown at periphery F hypothecium with light (olive-)brown pigment G ascospores 1-septate, Dirinaria-type but lumina angular to globose H habitus and apothecia on bark of Tilia amurensis. Thallus more grayish I apothecial section representing well-developed amphithecium and pigmented hypothecium J asci clavate with eight spores K ascospores 1-septate, Dirinaria-type but lumina angular to globose. Scale bars: 1 mm (A–C); 200 μm (D); 50 μm (E, F); 10 μm (G); 1 mm (H); 200 μm (I); 10 μm (J, K).

Apothecia abundant, rounded, erumpent in the beginning and sessile when mature, constricted at the base, 0.2–0.6 mm diam. Disc flat, not pruinose but epinecral debris shown in water, black to dark brown from early stages, 150–200 μm thick; margin persistent, prominent, generally entire or a little crenulate, concolorous to thallus. Amphithecium well-developed, with small crystals in the algal-containing medulla and particularly near the base, dissolving in K, 70–90 μm wide laterally, algal cells evenly distributed from periphery to base, 10–15 μm diam., cortical layer brownish, cortical cells granular, 2–3 μm diam., with epinecral layer, up to 5 μm thick. Parathecium hyaline but light brown at periphery, 5–10 μm wide laterally and 20–50 μm wide at periphery. Epihymenium red-brown, small granules not dissolving in K, 8–10 μm high. Hymenium hyaline, 90–95 μm high. Hypothecium brown with olive pigment in upper part, prosoplectenchymatous (irregular), 60–65 μm high. Oil droplets present a little in hypothecium. Paraphyses septate, anastomosing, 0.5–1 μm wide, simple or branched at tips, tips generally not swollen or little swollen, not pigmented, epihymenium pigmented by small granules, not by paraphysial tips, up to 1.5 μm wide. Asci clavate, 8-spored, 60–75 × 15–21 μm (n = 3). Ascospores ellipsoid, 1-septate, Dirinaria-type but lumina angular to globose, Type B development not detected, septum inflated a little or not, without a torus, hyaline when young and generally brown or dark brown in mature, 11–20 × 5–8.5 μm (mean = 15.4 × 7.1 μm; SD = 1.77(L), 0.70(W); L/W ratio 1.5–3.4, ratio mean = 2.2, ratio SD = 0.3; n = 105). Pycnidia raised, asymmetric, 175–225 μm wide. Pycnoconidia bacilliform, 3–4 × 0.5 μm.

Chemistry

Thallus K–, KC–, C–, Pd–. Hymenium I+ blue. UV–. Zeorin was detected by TLC.

Distribution and ecology

The species occurs on the bark of Quercus mongolica, Tilia amurensis Rupr., and Maackia amurensis Rupr. & Maxim. The species is currently known from a humid forest and a forested wetland of two mountainous sites.

Etymology

The species epithet indicates that the lichen’s substance, zeorin, is a major compound.

Notes

The new species is similar to R. hypobadia, R. sheardii, and R. sp. A in having a pigmented hypothecium. However, the new species differs from R. hypobadia by areolate, brownish thallus, apothecia without pruina, hyaline and wider parathecium, narrower paraphyses with hyaline and unswollen tips, longer and narrower ascospores with just angular to globose lumina, and the absence of pannarin (Sheard et al. 2017).

The new species differs from Rinodina sheardii by the absence of vegetative propagules, and Dirinaria-type ascospores in smaller size (Sheard et al. 2017).

The new species differs from Rinodina sp. A by wider parathecium, narrower paraphyses with swollen tips, smaller ascospores Dirinaria-type, and the absence of pannarin (Sheard et al. 2017).

The new species can be compared with R. manshurica and R. aff. oleae in having erumpent apothecia, small ascospores(<21 μm long) with swollen septum among corticolous species. However, the new species differs from R. manshurica by crystals present in the amphithecium, wider parathecium, narrower paraphyses without swollen tips, pigmented hypothecium, and longer and narrower ascospores (Tønsberg 1992; Sheard et al. 2017).

The new species is distinguished from R. aff. oleae by narrower ascospores, and pigmented hypothecium (vs. hyaline hypothecium) (Sheard et al. 2017). Reference Table 3 provides the key characteristics distinguishing R. zeorina from the compared species above.

Table 3.

Comparison of Rinodina zeorina with closely-related species.

Species Rinodina zeorina Rinodina hypobadia Rinodina manshurica Rinodina sheardii Rinodina aff. oleae Rinodina sp. A
Thallus growth from areolate, rimose to continuous rimose, not areolate rimose, rimose-areolate ±areolate to ±continuous, sorediate continuous, rimose-areolate continuous to areolate
Thallus color light gray to light brownish gray light to dark gray gray-brown yellow, yellow-brown, or pale brown or greenish (dark gray to olive-green) dark gray to gray-brown
Pruina absent, but epinecral debris shown in water slightly pruinose absent absent (absent)
Parathecium color hyaline and light brown at periphery red-brown red-brown to brown (hyaline to brownish)
Parathecium at periphery (μm) 20–50 10–20 c. 20 c. 30 (up to 30) c. 25
Paraphyses (μm) up to 1.5 2–2.5 2.0 2.0 (1–2) 3.0
Paraphysial tips not or little swollen, not pigmented 3–4 μm, lightly pigmented c. 3 μm, light pigmented c. 3 μm c. 4.5 μm, pigmented
Hypothecium color brown with olive pigment reddish or chestnut brown hyaline dilute brown to red-brown hyaline light brown
Crystals in amphithecium present in medulla present in both cortex and medulla absent present present in medulla
Ascospore type Dirinaria-type with angular-globose lumina Dirinaria-type with Physcia- or Physconia-like lumina Dirinaria-type, with Physcia-like lumina Pachysporaria-type I Dirinaria-type with Physcia-like lumina Pachysporaria-type I
Ascospores (μm) 11–20 × 5–8.5 12.5–18.5 × 6.5–10 14–16.5 × 7.5–8.5 16–35 × 8–17 15.5–19 × 6.5–9.5 22–28.5 × 10.5–15.5
Pycnidia 175–225 up to 300
Pycnoconidia (μm) 3–4 × 0.5 3.5 × 1.0 (4–5 × 1)
Substance zeorin pannarin, zeorin absent zeorin (absent) pannarin, zeorin
Reference BDNA-L-0000933 (holotype), BDNA-L-0000642, BDNA-L-0000646, BDNA-L-0000650, BDNA-L-0000651, BDNA-L-0000668 Sheard et al. 2017 Sheard et al. 2017 Tønsberg 1992; Sheard et al. 2017 Joshi et al. 2013; Smith et al. 2009; Sheard et al. 2017 Sheard et al. 2017

The new species is compared further with other Rinodina species having the substance zeorin, R. ascociscana (Tuck.) Tuck., R. buckii Sheard, R. efflorescens Malme, R. luteonigra Zahlbr., R. subalbida (Nyl.) Vain., R. subminuta H. Magn., and R. willeyi Sheard & Giralt. However, all of them are different from the new species because those species represent larger ascospores in Physcia- to Physconia-type for R. ascociscana; sorediate thallus, mostly light brown hypothecium and Teichophila-type ascospores and the presence of pannarin for R. buckii; sorediate thallus, colorless hypothecium, Pachysporaria-type ascospores and the presence of pannarin and secalonic acid A for R. efflorescens; colorless hypothecium, larger ascospores in Pachysporaria-type and the presence of thiomelin for R. luteonigra; larger spores in Pachysporaria-type and the presence of pannarin for R. subalbida; larger spores in Physcia-type for R. subminuta; sorediate thallus and the presence of pannarin for R. willeyi (Sheard et al. 2012, 2017).

Additional specimens examined

South Korea, Gangwon Province, Pyeongchang-gun, Daegwallyeong-myeon, Heonggye-ri, a forested wetland, 37°46.00'N, 128°42.33'E, 1,047 m alt., on bark of Maackia amurensis, 03 June 2020, B.G. Lee & H.J.Lee 2020-000442, with Buellia disciformis (Fr.) Mudd, Buellia sp., Catillaria nigroclavata (Nyl.) J. Steiner, Lecanora megalocheila (Hue) H. Miyaw., Lecanora symmicta (Ach.) Ach., Lecidella euphorea, and Lambiella cf. caeca (J. Lowe) Resl & T. Sprib. (BDNA-L-0000642; GenBank MW832812 for ITS); same locality, 37°46'0.02"N, 128°42'19.58"E, 1,047 m alt., on bark of Maackia amurensis, 03 June 2020, B.G. Lee & H.J.Lee 2020-000446 (BDNA-L-0000646; GenBank MW832813 for ITS); same locality, 37°46.00'N, 128°42.33'E, 1,047 m alt., on bark of Maackia amurensis, 03 June 2020, B.G. Lee & H.J.Lee 2020-000450 (BDNA-L-0000650; GenBank MW832814 for ITS); same locality, 37°46.00'N, 128°42.33'E, 1,047 m alt., on bark of Maackia amurensis, 03 June 2020, B.G. Lee & H.J.Lee 2020-000451 (BDNA-L-0000651; GenBank MW832815 for ITS); same locality, 37°46.00'N, 128°42.33'E, 1,047 m alt., on bark of Tilia amurensis, 03 June 2020, B.G. Lee & H.J.Lee 2020-000468, with Amandinea punctata (Hoffm.) Coppins & Scheid., Bacidia aff. beckhausii Körb., Catillaria sp., Micarea prasina Fr., Phaeophyscia limbata (Poelt) Kashiw., Rinodina cf. oleae Bagl., Traponora aff. varians (Ach.) J. Kalb & Kalb (BDNA-L-0000668; GenBank MW832816 for ITS).

Key to the species of Rinodina from the far eastern Asia (63 taxa)

Eleven more species have been recorded since Sheard et al. (2017), such as Rinodina badiexcipula, R. colobinoides, R. convexula, R. herrei, R. laevigata, R. occulta, R. oxneriana, R. parasitica, R. tephraspis, and two new species from this study (Kondratyuk et al. 2016, 2017; Yakovchenko et al. 2018; Galanina and Ezhkin 2019; Kondratyuk et al. 2020; Galanina et al. 2021). Particularly, R. laevigata of Aptroot and Moon (2014) was rejected by Sheard et al. (2017), but Galanina et al. (2021) confirmed the species in the far eastern Asia. This key includes all above species except for R. convexula because the species was just announced for a new record to Korea without any specific description for reference (Kondratyuk et al. 2020). Rinodina confragosa (Ach.) Körb., R. milvina (Wahlenb.) Th. Fr., and R. olivaceobrunnea C.W. Dodge & G.E. Baker were reported from Korea and Russian Far East (Kondratyuk et al. 2016; Galanina et al. 2021) as expected to occur (Sheard et al. 2017). All expected species are remained with an asterisk mark(*).

Overall, 63 taxa of Rinodina are currently recorded or expected to the far eastern Asia (Korea, Japan and Russian Far East).

1 Substratum rock 2
Substratum bark, wood, soil, decaying ground vegetation, bone or other lichens 17
2 Thalli with vegetative propagules 3
Thalli lacking vegetative propagules 4
3 Thallus effigurate, typically with isidia; when fertile spores belong to the Physconia-type; associated with seabird colonies; northern R. balanina
Thallus not effigurate, vegetative propagules blastidia with budding soredia; spores Pachysporaria-type II; not coastal; southern R. placynthielloides
4 Always maritime, typically on coastal rocks; spores Dirinaria-type R. gennarii
Generally inland or occasionally maritime; spores belonging to a different type 5
5 Medulla orange, K+ red-violet; spores Pachysporaria-type I, ultimately developing satellite apical lumina R. cervina
Medulla not orange, not K+ red-violet; spores of various types but never developing apical lumina 6
6 Thallus and apothecium margins K+ yellow, atranorin in cortex 7
Thallus and apothecium margins K−, atranorin absent 10
7 Spores with angular lumina, walls thickened at septum and apices, Physcia-type; proper exciple hyaline throughout, or if lightly pigmented not aeruginose (N−); thalline margin never pigmented 8
Spores with ‘hourglass’-shaped lumina, Mischoblastia-type; proper exciple typically aeruginose at periphery (N+ red under microscope); thalline margin often becoming pigmented 9
8 Apothecia 0.1–0.3 mm diam., hymenium 80–100 μm high, hypothecium 65–135 μm high, asci 75–80 × 16–19 μm, spores 17–27 × 8–13 μm R. confragosa
Apothecia 0.6–1.5 mm diam., hymenium 55–85 μm high, hypothecium 10–55 μm high, asci 45–50 × 13–20 μm, spores 11–16 × 5–9 μm R. occulta
9 Thallus plane; spores averaging <21 μm in length, rarely swollen at septum R. oxydata
Thallus verrucose; spores averaging >21 μm in length, often swollen at septum when mature R. moziana (syn. R. destituta)
10 Spores elongately ellipsoid, l/w ratio c. 2.0, Pachysporaria-type R. cinereovirescens
Spores broadly ellipsoid, l/w ratio <2.0, belonging to various types 11
11 Spores >20 μm long at maximum, Teichophila-type, often swollen at septum, more so in KOH 12
Spores <20 μm long, never swollen at septum, belonging to another type 13
12 Spores 18.5–25 × 10–12.5 μm R. tephraspis
Spores 20–32 × 11–19 μm R. teichophila
13 Spores with broad pigmented band around septum, Bischoffii-type R. bischoffii *
Spores lacking a broad pigmented band around septum, belonging to another type 14
14 Spores with Physcia-like lumina when immature, becoming rounded especially at the apices, lateral walls thin 15
Spores with rounded lumina from beginning, lateral walls relatively thick 16
15 Thallus thick, dark brown; spores constricted at septum when mature, Milvina-type; secondary metabolites absent R. milvina
Thallus thin, gray to light brown; spores Physconia-type; thalline margin C+ red (under microscope), gyrophoric acid in medulla R. sicula
16 Apothecial discs pruinose; spores Pachysporaria-type R. compensata
Apothecial discs not pruinose; spores Pachysporaria- to Milvina-like R. kozukensis
17 On soil, decaying ground vegetation, wood, bone or lichenicolous 18
Strictly corticolous or lignicolous 27
18 Spores 1-septate 19
Spores 3-septate or submuriform 20
19 Spores Teichophila-type R. herrei
Spores Physcia-type, rarely with apical satellite lumina 21
20 Spores strictly 3-septate, type B development (apical wall thickened prior to septum formation); secondary metabolites absent R. conradii
Spores 3-septate at first, typically becoming submuriform, type A development (apical wall thickening after septum formation); deoxylichesterinic acid present R. intermedia
21 Strictly lichenicolous, on Aspicilia or Rhizocarpon R. parasitica
Generally not lichenicolous 22
22 Sphaerophorin crystals in medulla (sometimes lichenicolous) R. turfacea
Sphaerophorin lacking in medulla (never lichenicolous) 23
23 Cortex K+ yellow or medulla orange, K+ red 24
Cortex reaction absent 25
24 Thallus light gray; K+ yellow, atranorin in cortex R. mniaroeiza *
Thallus a shade of brown; medulla orange, K+ red, skyrin or other anthraquinones present R. cinnamomea *
25 Spores averaging <23 μm in length R. olivaceobrunnea
Spores averaging >23 μm in length 26
26 Thallus and apothecia not pruinose; apothecial discs becoming convex, thalline margin then excluded; spores averaging 24.5–25.5 μm in length, l/w ratio 2.0–2.2 R. mniaroea
Thallus and apothecia typically pruinose; apothecial discs plane or concave, not convex, thalline margin never excluded; spores averaging 30–32 μm in length, l/w ratio 2.2–2.5 R. roscida
27 Vegetative propagules present 28
Vegetative propagules absent 37
28 Thallus typically golden yellow 29
Thallus a shade of gray or brown 30
29 Thallus with small, dense isidia; very rarely with apothecia; spores Pachysporaria-type I R. chrysidiata
Thallus with marginal, labriform soralia, sometimes becoming pustulate; frequently, but not always, with apothecia; spores Physcia-type R. xanthophaea
30 Phyllidia present R. oxneriana
Blastidia or soredia present 31
31 Thallus mainly blastidiate, blastidia 35–60 μm diam. R. colobinoides
Thallus generally not blastidiate, but sorediate or sometimes blastidiate 32
32 Blastidia present at margin, no substance, spores Teichophila-type R. herrei
Soredia and/or blastidia present, atranorin or pannarin present, spores in various types 33
33 Thallus light gray; soralia labriform at first, soredia whitish; K+, P+ yellow, cortical atranorin present, pannarin absent R. subparieta (syn. R. degeliana)
Thallus darker gray; soredia never whitish; K−, P+ cinnabar, atranorin absent, pannarin present 34
34 Thallus usually of convex to bullate areoles; blastidia often present, sometimes breaking into soredia; zeorin typically absent, when fertile pannarin also in epihymenium R. excrescens
Thallus never consisting of bullate areoles; soredia always present; zeorin typically present, pannarin never in epihymenium 35
35 Soredia typically yellowish, secalonic acid A present; spores Physcia-type when fertile, averaging <20 μm in length R. efflorescens
Soredia never yellowish, secalonic acid A absent; spores not Physcia-type, averaging >20 μm in length 36
36 Thallus minutely verrucose, verrucae central on areoles, quickly forming raised soralia, later spreading over thallus surface; soredia >40 μm diam.; spores Teichophila-type R. buckii
Thallus with plane areoles, soredia developing marginally on areoles, never raised centrally on verrucae, later spreading over thallus surface; soredia <40 μm diam.; spores Pachysporaria-type I R. willeyi
37 Ascospores 3-septate or submuriform 38
Ascospores 1-septate, rarely with satellite apical cells 39
38 Spores strictly 3-septate, type B development (apical wall thickened prior to septum formation); secondary metabolites absent R. conradii
Spores 3-septate at first, becoming submuriform, type A development (apical wall thickening after septum formation); deoxylichesterinic acid present R. intermedia
39 Thallus brightly pigmented; xanthone present, UV+ orange 40
Thallus a shade of gray or brown; xanthone absent, UV− 41
40 Thallus citrine, thiomelin present; spores averaging 31.0–34.5×16.0–17.5 μm, Pachysporaria-type I; not sorediate; subtropical, Tsushima Island, Japan R. luteonigra
Thallus golden yellow, secalonic acid A present; spores averaging 23.5–28.5×2.0–15.0 μm, Physcia-type; frequently sorediate; temperate, widely distributed R. xanthophaea
41 Thallus K+ yellow or P+ cinnabar, atranorin or pannarin present 42
Thallus K−, P−, both atranorin and pannarin absent 49
42 Thallus K+ yellow, atranorin present, pannarin absent 43
Thallus P+ cinnabar, pannarin present, atranorin absent 45
43 Spores averaging >33 μm long, Pachysporaria-type I R. megistospora
Spores averaging <33 μm long, Physcia- or Physconia-type 44
44 Spores averaging >26 μm long, strictly Physcia-type; never sorediate; distribution limited to coastal foreshores R. macrospora
Spores averaging <26 μm long, Physcia- to Physconia-type; most frequently sorediate; distribution inland R. subparieta (syn. R. degeliana)
45 Hypothecium pigmented dark reddish brown; spores Dirinaria-type, (12–)14–16.5(–18)× (6.5–)7.0–8.5(–9.5) μm, lightly pigmented R. hypobadia
Hypothecium never strongly pigmented; spore type otherwise 46
46 Spores averaging <20 μm in length, Physcia-type; thallus becoming bullate, often with minute blastidia R. excrescens
Spores averaging >20 μm in length, not Physcia-type; thallus sometimes verrucate but never bullate or blastidiate 47
47 Thallus persistently plane; epihymenium lacking crystals, P−; spores averaging >29 μm R. tenuis (syn. R. adirondackii)
Thallus becoming verrucate; epihymenium with or without crystals, P+ or P−; spores averaging <29 μm 48
48 Epihymenium typically possessing pannarin crystals, P+ cinnabar; spores lacking apical canals; widely distributed in Japan and adjacent mainland R. subalbida
Epihymenium lacking pannarin crystals, P−; spores with very obvious apical canals; Cheju Island, Korea Rinodina sp. A
49 Spores 16 per ascus R. polyspora
Spores 4–8 per ascus 50
50 Medulla with sphaerophorin crystals, PL+ 51
Medulla lacking sphaerophorin crystals, PL− 52
51 Thallus dark gray, typically dark brown; areoles becoming contiguous, plane, 0.40–0.55 mm wide; spores averaging 26.5–27.5 × 13.5–14.5 μm R. badiexcipula
Thallus light gray, sometimes brownish; areoles remaining discrete, convex, 0.20–0.30 mm wide; spores averaging 23.0–25.5 × 11.5–13.5 μm R. cinereovirens
52 Spores swollen at septum, more so in KOH, type B development (apical wall thickening prior to septum formation), Dirinaria-type 53
Spores not swollen at septum, even in KOH, type A development (apical wall thickening after septum formation), various types 59
53 Spores averaging >21 μm long R. endospora
Spores averaging <21 μm long 54
54 Spores lacking wall thickening at maturity (septal and apical thickenings may be present briefly in immature spores) 55
Spore lumina Physcia-like, with persistent apical wall thickening 56
55 Thallus gray to ochraceous, rugose, areoles to 0.7 mm wide; apothecia to 0.8 mm in diam., discs plane, never convex; spores averaging 15.5–18.0 × 8.0–8.5 μm, l/w ratio 1.9–2.1 R. mongolica
Thallus gray, never ochraceous, continuous to rimose; apothecia to 0.30–0.50 mm in diam., discs often becoming convex; spores averaging 12.5–13.5 × 5.5–6.0 μm, l/w ratio 2.1–2.4 R. pyrina *
56 Apothecia not erumpent; spores averaging 17.5–21.5 × 9–11 μm R. metaboliza
Apothecia erumpent; spores smaller 57
57 Hypothecium pigmented with brown, spores 11–20 × 5–8.5 μm, zeorin present R. zeorina
Hypothecium colorless, spores 15.5–18 × 8–9 μm, no substance 58
58 Spores averaging 15.5–16.0 μm in length R. manshurica
Spores averaging 16.5–18.0 μm in length R. aff. oleae
59 Spores averaging >22 μm in length 60
Spores averaging <22 μm in length 61
60 Margins of apothecia often radially cracked; spores Physcia- to Physconia-type R. ascociscana (syn. R. akagiensis, R. melancholica)
Margins of apothecia not radially cracked; spores Pachysporaria-type I R. dolichospora
61 Spores Pachysporaria-type II R. salicis
Spores Physcia- or Physconia-type 62
62 Spores Physcia- to Physconia-type, some lumina becoming rounded at apices, at maturity thin-walled 63
Spores strictly Physcia-type, apical walls remaining thick 67
63 Thallus dark brown, spores darkly pigmented at maturity, torus prominent; oro-arctic to coastal 64
Thallus a shade of gray, sometimes brownish, spores typically pigmented at maturity, torus present but not prominent; boreal 66
64 Thallus inconspicuous; apothecia mostly crowded, typically broadly attached R. olivaceobrunnea *
Thallus of dispersed or contiguous areoles; apothecia mostly dispersed, narrowly or broadly attached 65
65 Ascospores 20–21.5 × 10–11.5 μm, thallus well-developed, flat, scurfy or thick rugose areolate, apothecia broadly attached in the beginning then becoming narrow and even stipitate, discs convex when mature R. sibirica
Ascospores 18.5–19.5 × 8.5–9.0 μm, thallus poorly developed, evanescent, thin or scabrid, sometimes areolate, apothecia broadly attached to thallus, discs typically flat R. laevigata
66 Thallus thick, rugose, areolate; apothecia crowded, discs persistently plane, thalline margins persistent R. archaea *
Thallus thin, plane, continuous or rimose-areolate; apothecia dispersed, discs becoming convex, often excluding thalline margin R. trevisanii *
67 Spores averaging >18 μm long, zeorin present R. subminuta
Spores averaging <18 μm long, zeorin absent 68
68 Apothecia erumpent at first, discs often becoming strongly convex; spores with lightly pigmented tori at maturity R. orientalis
Apothecia never erumpent, discs persistently plane; spores with very dark, prominent tori at maturity 69
69 Apothecia crowded, broadly attached; thalli associated with leaf scars or other mesic microhabitats; areoles plane, contiguous, to >0.2 mm in diam. R. freyi
Apothecia mostly scattered, narrowly attached; thalli typically in more xeric microhabitats; areoles convex, scattered, to 0.2 mm in diam. R. septentrionalis

Acknowledgements

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.

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