Taiwania 67(1): 155‒163, 2022
DOI: 10.6165/tai.2022.67.155

NOTE

Noteworthy foliicolous lichens collected from Iriomote Island, southernJapan
Kento MIYAZAWA1,*, Yoshihito OHMURA2, Yuichi YAMAOKA3
1. Degree Program in Life and Earth Sciences, Graduate School of Science and Technology, University of Tsukuba, 1-1-1 Tennodai,
Tsukuba, Ibaraki, 305-8572, Japan. 2. Department of Botany, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba,
Ibaraki, 305-0005, Japan. 3. Faculty of Life and Environmental Sciences, University of Tsukuba, 1-1-1 Tennodai, Tsukuba, Ibaraki,305-8572, Japan. *Corresponding author’s email: tentomushi.3838@gmail.com(Manuscript received 16 August 2021; Accepted 18 January 2022; Online published 25 January 2022)
ABSTRACT: The following noteworthy species were identified based on foliicolous lichen material collected in 2011 and 2019 on
Iriomote Island: Byssoloma chlorinum, B. vanderystii and Gallaicolichen pacificus, reported as new to Japan, and B. minutissimum
and Fellhanera rhapidophylli, reported as new to Iriomote Island. Hyphophores and diahyphae of Tricharia kashiwadanii were
described for the first time, based on the original material and a collection from Iriomote Island. The mtSSU sequences of B.
chlorinum, Porina kamerunensis and T. kashiwadanii were obtained for the first time as well as additional sequences of B. annuum,
B. vanderystii, Eugeniella micromata. The mtSSU sequence was obtained from the specimens were compared in terms of homology
with those registered in GenBank.
KEY WORDS: Ascomycota, lichenized fungi, mtSSU, mycota, red-listed taxa, taxonomy, the Tropics, threatened species.

INTRODUCTION

MATERIALS AND METHODS

There are more than 800 species of foliicolous lichensworldwide, distributed mainly in tropical regions (e.g.,
Santesson, 1952; Lücking, 2008). In Japan, 85 taxa offoliicolous lichens have been reported (Thor et al., 2000;
Suto and Ohtani, 2015; Miyazawa et al., 2020). Amongthem, 14 taxa are categorized as “Near Threatened” (NT)and 14 taxa as “Data Deficient” (DD) in the Red List forJapanese species (Ministry of the Environment, 2020).
Iriomote Island, one of the Ryukyu Islands, is locatednear the southernmost part of Japan (N24°17′33″,
E123°51′43″) and about 200 km east of Taiwan(Matsuzawa, 2019). Approximately 90% of the island iscovered with subtropical rainforest, forming a semiclosed ecosystem with little influence fromanthropogenic activities (Matsuzawa, 2019), and a largepart of island is well-preserved old-growth forest locatedwithin the national park. Sixty-five taxa of foliicolous
lichens were reported from Iriomote Island before thepresent study (Thor et al., 2000; Ohmura, 2012;
Miyazawa et al., 2020). Although this represents thehighest number of taxa among the foliicolous lichens
reported in Japan, further investigations are expected tofind additional taxa (Thor et al., 2000).
The aim of this study was to conduct taxonomic
examinations of the foliicolous lichens collected from
Iriomote Island and to report additional information on
the foliicolous lichen mycota of Japan.

Collections
The field investigations were performed by Y.
Ohmura and G. Tanaka on 8–11 March 2011 and by K.
Miyazawa and Y. Ohmura on 18–21 November 2019 on
Iriomote Island, and approximately 370 specimens offoliicolous taxa ware collected. All voucher specimens
are housed in the herbarium of the National Museum of
Nature and Science (TNS), Tsukuba, Japan.
Morphology and chemistry
Morphological observations were performed using a
dissecting microscope (MZ-8 [Leica, Wetzlar, Germany]or SZ-PT [Olympus, Tokyo, Japan]) and a differential
interference contrast microscope (BX51; Olympus).
Anatomical examination was performed using hand-cut
sections mounted in a glycerin/ethanol/water (1:1:1)
solution (Asahina, 1936). Photographs were taken using
a digital camera (COOLPIX P6000 [Nikon, Melville, NY,
USA] or NEX-5 [Sony, New York, NY, USA]), and theimages obtained were subjected to depth stacking using
CombineZP (https://combinezp.software.informer.com/)(Fig. 2A). Spore size is expressed as follows: (minimum–)range of mean ± standard deviation (–maximum) (n =number of measurements).
Ascus amyloidity was assessed using with 5–10%
KOH solution and Lugol's solution. Crystals wasevaluated by polarization under differential microscopy.
Secondary substances from the thallus were examined byhigh-performance thin-layer chromatography (HPTLC)
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Table 1. BLAST results of mtSSU sequences obtained in this study. If the homology of the top-hit sequence was greater than 95%,
three sequences were presented, but if it was less than 95%, only the top-hit sequence was presented.
Species/ Voucher
Byssoloma chlorinum
K. Miyazawa 372 & Y. Ohmura (TNS)
K. Miyazawa 549 & Y. Ohmura (TNS)
K. Miyazawa 566 & Y. Ohmura (TNS)
K. Miyazawa 567 & Y. Ohmura (TNS)
K. Miyazawa 569 & Y. Ohmura (TNS)
K. Miyazawa 570 & Y. Ohmura (TNS)
K. Miyazawa 575 & Y. Ohmura (TNS)
B. annuum
K. Miyazawa 451 & Y. Ohmura (TNS)GenBank No.a BLAST Resultb
LC648410
LC648417
LC648419
LC648420
LC648421
LC648422
LC648423

737/795 (93%), 1136, 0.0 (MN105612, Byssoloma leucoblepharum)
731/787 (93%), 1133, 0.0 (MN043721, Byssoloma leucoblepharum)
770/832 (93%), 1182, 0.0 (MN105612, Byssoloma leucoblepharum)
777/839 (93%), 1195, 0.0 (MN105612, Byssoloma leucoblepharum)793/862 (92%), 1201, 0.0 (MN043695, Byssoloma cf. leucoblepharum)
763/822 (93%), 1181, 0.0 (MN105612, Byssoloma leucoblepharum)792/861 (92%), 1199, 0.0 (MN043695, Byssoloma cf. leucoblepharum)

LC648414

819/824 (99%), 1496, 0.0 (MN043716, Byssoloma annuum)
816/821 (99%), 1491, 0.0 (MN043716, Byssoloma annuum)
817/843 (97%), 1496, 0.0 (MN105600, Byssoloma brunneodiscum)
806/807 (99%), 1496, 0.0 (MN043726, Byssoloma annuum)
788/789 (99%), 1452, 0.0 (MN043716, Byssoloma annuum)
786/808 (97%), 1367, 0.0 (MN105603, Byssoloma brunneodiscum)
806/809 (99%), 1476, 0.0 (MN043726, Byssoloma annuum)
788/791 (99%), 1443, 0.0 (MN105603, Byssoloma annuum)
787/809 (97%), 1367, 0.0 (MN105603, Byssoloma brunneodiscum)759/762 (99%), 1389, 0.0 (MN043726, Byssoloma annuum)
759/762 (99%), 1389, 0.0 (MN043716, Byssoloma annuum)
741/762 (97%), 1286, 0.0 (MN105603, Byssoloma brunneodiscum)

K. Miyazawa 501 & Y. Ohmura (TNS)

LC648415

K. Miyazawa 558 & Y. Ohmura (TNS)

LC648418

K. Miyazawa 583 & Y. Ohmura (TNS)

LC648424

B. vanderystii
K. Miyazawa 400 & Y. Ohmura (TNS)

LC648411

Eugeniella micromata
K. Miyazawa 437 pr.p. & Y. Ohmura (TNS) LC648413
Porina kamerunensis
K. Miyazawa 504 & Y. Ohmura (TNS)
LC648416
Tricharia kashiwadanii
K. Miyazawa 405 & Y. Ohmura (TNS)
LC648412

795/796 (99%), 1465, 0.0 (MN043715, Byssoloma vanderystii)
795/796 (99%), 1465, 0.0 (MN043713, Byssoloma vanderystii)
794/796 (99%), 1459, 0.0 (MN043719, Byssoloma vanderystii)
783/792 (99%), 1413, 0.0 (MK957161, Eugeniella micrommata)611/660 (93%), 935, 0.0 (DQ168394, Porina subnitidula)
663/741 (89%), 929, 0.0 (KF833340, Echinoplaca sp.)a. GenBank accession number for mtSSU.
b. BLAST result with GenBank sequences. Identity; score; e-value accession number, species) of the top 3 sequences (maximum) ofrelated taxa are shown.

following the method of Schumm and Elix (2015).
The detailed morphologies of peltidiangia andpeltidia were observed by scanning electron microscopy(SEM) (S-4200; Hitachi, Tokyo, Japan). A fragment ofthallus containing peltidiangia was cut and attached to asample stand with double-sided tape. The sample wascoated with platinum palladium by an ion sputtering
device (E1030; Hitachi) for SEM.
DNA extraction, PCR amplification and sequencing
DNA extraction from selected samples was
conducted following the method of Izumitsu et al. (2012).
One to three apothecia were picked from a colony or afragment of thallus, ground using sterile glass slides, andplaced in a 1.5-mL tube containing 100 μL TE buffer. The
tubes were heated in a microwave oven (600 W) for 1
min., kept at room temperature for 30 s, and heated again
in a microwave oven (600 W) for 1 min. The heatedmicrotubes were immediately stored in a freezer at −20°C
for approximately 1 day, and 2 μL of solution from eachmicrotube was used as a DNA template for PCR.
The partial sequence of the fungal mitochondrial small
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subunit (mtSSU) was amplified using the primer set
mrSSU1 and mrSSU3R (Zoller et al., 1999) according to
the following protocol. PCR was performed in a 25 μL
reaction volume containing 2 μL DNA template, 12.5 μL
GenRED PCR Mix Plus (Nippon Gene, Tokyo, Japan), 2.5
μL each primer (2 pmol/μL), and 5.5 μL distilled water.
The PCR conditions were according to Wang et al. (2020)
using a TaKaRa PCR Thermal Cycler Dice® Touch(TaKaRa, Tokyo, Japan). The PCR products were checkedby electrophoresis on a 1.5 % agarose gel stained with
Midori Green Direct DNA Stain (Nippon Genetics, Tokyo,
Japan) and visualized using WSE-5200 Printgraph 2 M(ATTO Corporation, Tokyo, Japan). The PCR products
were purified using a FastGene™ Gel/PCR extraction kit(Nippon Genetics) following the manufacturer’s
instructions. Sequences were obtained by a DNA
sequencing service (Eurofins Genomics, Tokyo, Japan).
The obtained sequences were analyzed by the
GenBank BLAST search (blastn) using the default
settings. The voucher specimen data and GenBank
accession numbers for the obtained mtSSU sequences areshown in Table 1.

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RESULTS AND DISCUSSION
Taxonomic examination of 209 of the 370 specimens
identified 51 taxa at the species level (Supplement, Table
S1). Among them, 3 species were new to Japan and an
additional 2 species were new to Iriomote Island but
known from other localities in Japan; 13 species are thoselisted in the Red List for Japanese species (Ministry of theEnvironment, 2020) (Table S1).
For the species newly recorded in Japan, fulldescriptions are given based on the Iriomote Island
specimens. For the species newly recorded on Iriomote
Island and the red-listed species, including the DD
species, newly obtained taxonomic and ecologicalinformation are noted. The mtSSU sequences obtained
from the specimens were compared with those registeredin GenBank in terms of homology.
Species new to Japan
Byssoloma chlorinum (Vain.) Zahlbr. Lich. Univ. 8: 233.
1932.
Fig. 1A, B
Thallus continuous, 15–30 mm across, 10–50 μm
thick, minutely farinose to granulose, light green,sometimes whitish green. Apothecia rounded, up to 0.6
mm diam., 100–250 μm tall; margin byssoid, reduced towell-developed, persistent or rarely evanescent,occasionally spreading laterally over thallus surface, 50–
100 μm broad, white, composed of loosely woven
colorless to pale brown hyphae (2–3 μm wide); disc plane
to slightly convex, brownish black; epithecium indistinct;
hymenium 40–100 μm tall, colorless to pale brown;
hypothecium 40–70 μm tall, dark brown, K–; apothecialbase brownish black to aeruginous, K–; paraphyses
branched, 0.8–1.0 μm wide, not widened apically. Asciclavate, I+ blue, tholus amyloid (Byssoloma-type), 40–45
× 10–12 μm. Ascospores oblong-ellipsoid, 3-septate(rarely 4-septate), with or without slight constrictions at
septa, colorless, (8.0–)9.7–12.7(–16.0) × (2.5–)2.8–3.8(–4.0) μm (n = 40). Pycnidia subglobose to barrel-shaped,
80–180 μm diam., grey to black with white margin, walls
aeruginous. Conidia pyriform, non-septate, colorless,(3.0–)3.3–4.3(–5.0) × (1.0–)1.2–1.5(–1.5) μm (n = 20).
Photobiont trebouxioid, globose to subglobose,(5.0–)5.1–7.5(–9.0) × (4.0–)5.0–6.8(–8.0) (n = 20).
Chemistry: no secondary substances were detected by
HPTLC.
Byssoloma chlorinum is characterized by a light green
farinose thallus (Fig. 1A), blackish apothecia with whitish
developed byssoid margin (Fig. 1A), and 3-septate
ascospores (Fig. 1B).
The ascospore size in the materials in the Iriomote
Island collections (see above) (Fig. 1B) largely
overlapped with the range (9.6–15 × 2.0–4.3 μm) of the
protologue (Vainio, 1924) (10–12 × 2–2.5 μm), thedescription by Lücking (2008) (10–15 × 2.5–3.5 μm), andour measurements for an exsiccata (Lücking: Lich. Fol.

Exs. 132, TNS) [(9.6–)10.0–11.9(–12.6) × (3.0–)3.3–
4.1(–4.3) μm (n=14)].
Hypothecium of the Iriomote Island materials is welldeveloped and much thicker (40–70 μm tall) than themeasurement (15–25 μm) reported by Lücking (2008),
causing the disc shape to be plane to slightly convex.
The byssoid excipulum varies from distinctly reduced
to well-developed spreading laterally over the thallus
surface. Such variations were observed even within thesame thallus in some specimens.
The BLAST results for the mtSSU sequences of the
Iriomote Island samples showed less than 93% identitywith those of other Byssoloma species in GenBank (Table1). It is because no sequence of B. chlorinum wasregistered in GenBank before this study.
Byssoloma chlorinum resembles B. vanderystii Sérus.,
but the latter is readily distinguished by chocolate-browndiscs and 7-sptate ascospores. Among foliicolous taxa, B.
chlorinum may be confused with B. leucoblepharum(Nyl.) Vain. However, B. chlorinum is distinguished fromthe latter by having a light green farinose thallus and dark
apothecia with a whitish byssoid margin, while B.
leucoblepharum has a dark smooth thallus and apothecia
with a chamois-colored byssoid margin (Lücking, 2008).
Further taxonomic research is needed for these species
because the features are variable and sometimes
overlapping (see Lücking, 2008).
Byssoloma chlorinum is from Asia (India [Gupta andSinha, 2017], Indonesia [Santesson, 1952], Thailand[Buaruang et al., 2017], and Vietnam [Aptroot andSparrius, 2006]), Oceania (Australia [Lücking et al.,2001], French Polynesia [type locality], and Papua New
Guinea [Aptroot and Sipman, 1993]), North and SouthAmerica (Bermuda [Berger and LaGreca, 2014], andother Neotropical regions [Lücking, 2008]), Europe(Azores [Breuss, 2018]), and Africa (Kenya [Lücking and
Kalb, 2002] and Seychelles [Seaward and Aptroot, 2009]),
showing a pantropical distribution (Lücking, 2008). This
species is new to Japan.
Exsiccati examined. COSTA RICA. Puntarenas Province:
Cocos Island National Park, ca. 540 km SW of Puntarenas in the NW
Pacific Ocean (N05°33′, W87°04′), on leaves of Calophyllum brasiliense,
0 m elev., April 1992, Lücking 92-1020 (Lücking: Lich. Fol. Exs. 132,
TNS); Limon Province: Cahuita National Park, “Kelly Creek” section,
140 km ESE of San José and 45 km SE of Limon at the Atlantic coast(N09°44', W82°50'), on leaves of undetermined dicotyledon, 0 m elev.,
October 1991, Lücking 91-2956 (Lücking: Lich. Fol. Exs. 133, TNS).
Specimens examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): Haemida Beach, Iriomote Island, Yaeyama Islands(N24°16′24″, E123°49′54″) on Arenga engleri, 10 m elev., 21 November
2019, K. Miyazawa 575 & Y. Ohmura (TNS); along the stream at SE footof Mt. Goza, Iriomote Island, Yaeyama Islands (N24°17′, E123°49′), on
broad-leaf tree, 10 m elev., 11 March 2011, Y. Ohmura 8037 & G. Tanaka(TNS); ditto, on Arenga engleri, Y. Ohmura 8058 & G. Tanaka (TNS);
along trail of Ohtomi-yuhodo, Iriomote Island, Yaeyama Islands(N24°17′58″, E123°51′45″), on Arenga engleri, 50 m elev., 18 November
2019, K. Miyazawa 372 & Y. Ohmura (TNS), K. Miyazawa 549 & Y.
Ohmura (TNS), K. Miyazawa 566 & Y. Ohmura (TNS), K. Miyazawa 567& Y. Ohmura (TNS), K. Miyazawa 569 & Y. Ohmura (TNS), K. Miyazawa
570 & Y. Ohmura (TNS), K. Miyazawa 577 & Y. Ohmura (TNS), K.

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Fig. 1. Byssoloma chlorinum and B. vanderystii. A. Thallus and apothecia of B. chlorinum (K. Miyazawa 372 & Y. Ohmura, TNS).
B. Ascospore of B. chlorinum (K. Miyazawa 372 & Y. Ohmura, TNS). C. Thallus and apothecia of B. vanderystii (K. Miyazawa 400& Y. Ohmura, TNS). D. Photobionts of B. vanderystii. E. Ascospores of B. vanderystii. Scales: A = 0.5 mm; B = 10 μm; C = 1 mm;
D–E = 20 μm.

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Miyazawa 580 & Y. Ohmura (TNS), K. Miyazawa 581 & Y. Ohmura(TNS); along the trail by Urauchi River, ca. 0.5 km NW of Maryudo
Waterfall, Iriomote Island, Yaeyama Islands (N24°21′, E123°48′), on deadleaf of Arenga engleri, 80 m elev., 9 March 2011, G. Tanaka 364 & Y.
Ohmura (TNS), Y. Ohmura 7966 & G. Tanaka (TNS).

Byssoloma vanderystii Sérus., Lichenologist 11: 181.
1979.
Fig. 1C–E
Thallus crustose, irregular in shape, continuous, ca. 25mm across, 15–25 μm thick, smooth to minutely farinose,
pale to dark greenish grey. Apothecia sessile to adnate,rounded, 0.2–0.6 mm diam., 150–200 μm tall; margin welldeveloped, densely byssoid, persistent, spreading laterally
over thallus surface, 80–200 μm broad, white, composedof loosely woven colorless hyphae; disc slightly to strongly
convex, chocolate-brown, with paler marginal zone;
epithecium indistinct; hymenium 70–100 μm tall, colorless;hypothecium 30–40 μm tall, dark brown, K–; apothecialbase dark brown, K–. Asci clavate, I+ blue, tholus amyloid(Byssoloma-type), 40–50 × 10–13 μm. Ascospores 8 per
ascus, cylindrical, (6–)7-septate, with or without slight
constrictions at septa, colorless, (22.0–)25.4–30.6(–31.0) ×(2.0–)1.9–2.5(–3.0) μm (n = 20). Pycnidia subglobose, 60–
140 μm diam., brownish black to black, covered by whitish
loose tissue. Conidia bacillar, non-septate, colorless,(4.0–)4.1–4.7(–5.3) × (1.0–)1.1–1.3 μm (n = 30).
Photobiont trebouxioid, ellipsoid, (4.6–)6.5–6.7(–8.1) ×(2.2–)2.8–4.2(–5.3) μm (n = 30).
Chemistry: no secondary substances were detected by
HPTLC.
Byssoloma vanderystii is characterized by convex
apothecia with a well-developed byssoid margin
spreading laterally over the thallus surface (Fig. 1C) and7-septate ascospores (Fig. 1E).
The morphology of the Iriomote Island material was
fundamentally consistent with the protologue and thedescription provided by Lücking (2008), except the sizeof the asci and photobiont. The ascus size of the Iriomote
Island material (40–50 × 10–13 µm) is approximately halfthat (70–90×12–15 µm) reported by Lücking (2008).
Although the ascus size is not mentioned in the
protologue, it was estimated to be 50–60 × 14–18 μm
based on the illustrations of the type specimen (Sérusiaux,
1979; Vězda, 1987). The ascus size seems to be highlyvariable in this species. However, whether this variabilitylies within the species or is attributed to B. vandersytii
comprising different species is uncertain based on thecurrent data. This study treats these differences as
variation within a single species. The photobiont of
Iriomote Island material is ellipsoid in shape (Fig. 1D),
while that described in the protologue is spherical with a
5–10 µm diameter. The photobiont of Iriomote Islandmaterial could represent a different species from the algal
taxon described in the protologue.
In addition to the identification based on morphology
and chemistry, the BLAST results showed high identity(99%) to B. vanderystii sequences registered in GenBank(Table 1).
Within the genus Byssoloma, B. braulioi Lücking, B.
catarinense L.I. Ferraro & Lücking, B. octomerum
Malcolm & Vězda, B. usambarense Vězda, B. syzygii
Vězda & Vivant, and B. vanderystii consistently have 7septate ascospores. Among the six species, B. vanderystii
is easily distinguished from others by a convex apothecia
with a well-developed byssoid margin spreading laterally
over the thallus surface.
Byssoloma vanderystii has been reported from Asia
(Malaysia [Sipman, 1993], Thailand [Wang et al., 2020],
and Vietnam [Nguyen, 2010]), Oceania (Australia[Sipman, 1991]), North and South America (Argentina[Ferraro, 1997], Brazil, Costa Rica, French Guiana[Lücking, 2008]), and Africa (Democratic Republic of theCongo [type locality] [Sérusiaux, 1979; Van den Broeck
et al., 2014]), showing a pantropical distribution(Lücking, 2008). This species is reported as new to Japan.
At present, Iriomote Island is the northernmost point of
the species.
Specimen examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along trail in the mountains along Urauchi river,Iriomote Island, Yaeyama Islands (N24°19′01″, E123°50′32″), on
Rhynchotechum discolor?, 210 m elev., 19 November 2019, K.
Miyazawa 400 & Y. Ohmura (TNS).

Gallaicolichen pacificus Sérus. & Lücking, Biblioth.
Lichenol. 95: 510, 2007.
Figs. 2 & 3
Thallus crustose, epiphyllous, pale yellowish gray,rounded and usually coalescing with adjacent thalli toform irregular shape, 1.2–2.5 mm in diam., smooth orslightly uneven, up to 20 µm thick, composed of a loose
network of interwoven hyphae and plates of photobiont
cells, containing crystals (up to 50 × 30 µm); prothallus
usually present, membranaceous and whitish.
Peltidiangia, 1–6 per a patch of thallus, almost perfectly
circular, 0.1–0.19 mm in diam. and 80–100 µm tall,formed by a rather thick erect margin with its inner parttypically whitish and its outer part usually covered by thethallus. Peltidia numerous, usually filling up the
peltidiangia cavity, disc-like, 20–30 µm in diam. and 10–
15 µm thick, composed of fungal hyphae and algal cellsof Trentepohliales, all linked to a central structure andcoiled up inwards. Ascomata not seen. Conidiomata notseen. Photobiont Trentepohliales, greenish brown, cells
6–11 × 3–5 µm, regularly and radiately arranged rows inplates.
Chemistry: no secondary substances were detected by
HPTLC.
Gallaicolichen pacificus is characterized by a paleyellowish gray rounded patched thallus with crystals (Fig.
2A, C), peltidiangia filled with peltidia (Figs. 2A, 3A, B),
and photobiont of Trentepohliales (Fig. 2C).The peltidia
size of the Iriomote Island materials is slightly smaller (20–30 µm in diam.) than that mentioned in the protologue
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Fig. 2. Gallaicolichen pacificus collected from Iriomote Island.
A. Thallus with peltidiangia (Y. Ohmura 7952 & G. Tanaka, TNS).
B. Peltidium (G. Tanaka 357 & Y. Ohmura, TNS). C. Plate ofphotobionts and crystals (black arrow indicates) (G. Tanaka 355& Y. Ohmura, TNS). Scales: A = 1 mm; B–C = 20 μm.

(25–55 µm) (Figs. 2B, 3B), but is considered within therange of variation because the ranges partially overlap. As
the ascomata and conidiomata of this genus have not beenfound to date, the taxonomic position under the phylum
Ascomycota is still unclear (Sérusiaux and Lücking, 2007;
Wijayawardene et al., 2018). The mtSSU sequence from theIriomote Island materials could not be obtained in this study.
Gallaicolichen pacificus is from the southeast Pacificregion: Australia (Queensland), Vanuatu, New Caledonia,
and Hawaii (Sérusiaux and Lücking, 2007). The distribution
is now extended to the west side of the Pacific Ocean as thenorthern limit. This species is reported as new to Asia.
Specimens examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along mountain path between Ohtomi-guchi Entrance
and Daiichi Yamagoya-ato, Iriomote Island, Yaeyama Islands(N24°19′49″, E123°51′04″), on Lasianthus attenuatus, 180 m elev., 19
November 2019, K. Miyazawa 492C & Y. Ohmura; along the trail by
Urauchi River from the landing point to Maryudo Waterfall, Iriomote
Island, Yaeyama Islands (N24°21′, E123°47′), on Arenga engleri, 60 m
elev., 9 March 2011, G. Tanaka 355 pr.p. & Y. Ohmura, (in collectionof Porina kamerunensis) (TNS); ditto, G. Tanaka 356 pr.p. & Y.
Ohmura, (in collection of Fellhanera bouteillei) (TNS); ditto, G.
Tanaka 357 pr.p. & Y. Ohmura (in collection of Asterothyrium cf.
decipenes) (TNS); ditto, Y. Ohmura 7950 pr.p. & G. Tanaka (incollection of Coenogonium subluteum) (TNS); ditto, Y. Ohmura 7952
& G. Tanaka (TNS); along the stream at SE foot of Mt. Goza, Iriomote
Island, Yaeyama Islands (N24°17′, E123°49′), on Lasianthus
cyanocarpus, 10 m elev., 11 March 2011, G. Tanaka 432 pr.p. & Y.
Ohmura (in collection of Calopadia sp.) (TNS); ditto, Y. Ohmura 8053
pr.p. & G. Tanaka (in collection of Mazosia bambusae) (TNS); ditto, on
Lasianthus cyanocarpus, Y. Ohmura 8080 pr.p. & G. Tanaka (incollection of Byssoloma leucoblepharum) (TNS); ditto, on Lasianthus
cyanocarpus, Y. Ohmura 8081 pr.p. & G. Tanaka (in collection of
Calopadia puiggarii) (TNS).

160

Fig. 3. SEM images of Gallaicolichen pacificus (Y. Ohmura
7952 & G. Tanaka, TNS). A. Peltidiangium with many peltidia. B.
Margin of peltidiangium and peltidia.

Species new to Iriomote Island
Byssoloma minutissimum Kalb & Vězda, Nova
Hedwigia 51: 445. 1990.
Thor et al. (2000) reported B. minutissimum from
Ishigaki Island in Japan as new to the Paleotropics but
showed that the ascospores size range is broader in widththan usual (10–12 × 2.5–3.5 μm vs. 2–2.5 μm). The
ascospore size range of the Iriomote Island material is evengreater [(10–)10.8–13.2(–16) × (2.5–)2.7–3.9(–5.5) μm (n= 50)]. The wider size range of ascospores was treated as avariation within this species in Thor et al. (2000). Our
study follows their treatment, and B. minutissimum isreported as new to Iriomote Island. However, further
taxonomic study treating on this variation is needed aftersufficient materials have been examined.
Specimen examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along the trail in the mountains along Urauchi river,Iriomote Island, Yaeyama Islands (N24°18′42″, E123°50′24″), on leaf

2022

Miyazawa et al.: Noteworthy foliicolous lichens collected from Iriomote Islandof Cleyera japonica, 220 m elev., 19 November 2019, K. Miyazawa 387& Y. Ohmura (TNS).

Fellhanera rhapidophylli (Rehm) Vězda, Folia Geobot.
Phytotax. 21: 214. 1986.
In Japan, Fellhanera rhapidophylli has been reported
only once from Yakushima Island and the main island of
Okinawa (Thor et al., 2000).
Specimen examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along the trail by Urauchi River from the landing point
to Maryudo Waterfall, Iriomote Island, Yaeyama Islands (N24°21′,
E123°47′), on Cleyera japonica, 60 m elev., 9 March 2011, G. Tanaka
353 pr.p. & Y. Ohmura (in collection of Mazosia bambusae) (TNS).

Additional data on the list of threatened species inJapanThirteen species in the Red List for Japanese species
including DD (Ministry of the Environment, 2020) wereidentified among the materials collected in this study.
Among these species, eight are treated as NT and five as
DD (Appendix 1). The list was formerly evaluated based
mainly on the rarity of the species. However, morequantitative evaluations are needed to clarify the rank ofeach species strictly according to the IUCN criteria. Our
additional data in this study will contribute to future
quantitative evaluation of red-listed species in Japan.
Byssoloma annuum (Vain.) G. Thor, Lücking & Tat.
Matsumoto
In addition to the morphological similarities, theBLAST results showed high identity (99%) to B. annuum
sequences in GenBank (Table 1).
Specimens examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along mountain path between Ohtomi-guchi Entrance
and Daiichi Yamagoya-ato, Iriomote Island, Yaeyama Islands(N24°19′26″, E123°50′49″), on broad-leaf tree, 190 m elev., 19
November 2019, K. Miyazawa 451 pr.p. & Y. Ohmura (in collection of
Sporopodium flavescens) (TNS); along trail in the mountains along
Urauchi river, Iriomote Island, Yaeyama Islands (N24°20′26″,
E123°49′58″), on Arenga engleri, 160 m elev., 19 November 2019, K.
Miyazawa 501 & Y. Ohmura (TNS); ditto (N24°20′52″, E123°49′24″),
on Arenga engleri, 100 m elev., 20 November 2019, K. Miyazawa 558& Y. Ohmura (TNS), K. Miyazawa 583 & Y. Ohmura (TNS).

Eugeniella micrommata (Kremp.) Lücking, Sérus. &Kalb
In addition to the morphological similarities, theBLAST results showed high identity (99%) to an
Eugeniella micrommata sequence in GenBank (Table 1).
Specimens examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along mountain path between Ohtomi-guchi Entrance
and Daiichi Yamagoya-ato, Iriomote Island, Yaeyama Islands(N24°19′01″, E123°50′32″), on broad-leaf tree, 210 m elev., 19
November 2019, K. Miyazawa 403 & Y. Ohmura (TNS); ditto,(N24°19′26″, E123°50′49″), on broad-leaf tree, 190 m elev., 19
November 2019, K. Miyazawa 437 pr.p. & Y. Ohmura (in collection of
Sporopodium flavescens) (TNS), K. Miyazawa 450 pr.p. & Y. Ohmura(in collection of Sporopodium flavescens) (TNS).

Porina kamerunensis F. Schill.
Although Thor et al. (2000) reported Japanese

specimens collected from Ishigaki Island and Iriomote
Island as Porina chrysophora (Stirt.) R. Sant., Lücking(2008) concluded that the specimens are P. kamerunensis(as P. kameruniensis). This study follows the same
treatment.
The ascospore size of the Iriomote Island materials
was (19.0–)19.7–21.8(–22.0) × (2.5–)2.8–3.5(–4.0) μm(n = 15). This is larger than that of the protologue (13–15.6 × 2–2.6 µm, Cameroon) and the description of
Lücking (2008) (14–20 × 2.5–4 µm, the Neotropics) but
is smaller than the description of Vězda (1997) (18–26 ×
3.5–4.5 μm, Vietnam, reported as P. chrysophora). Thoret al. (2000), who examined Japanese materials of Porina
kamerunensis (as P. chrysophora), reported a similar
measurement to that in this study [(19–)20–23(–25) ×(3–)3–4(–4) µm)]. Although the ascospore size differs byregion, we tentatively consider these differences to be
intraspecific variations because they are continuous.
Since the sequence of P. kamerunensis was notregistered in GenBank, the BLAST result showed lessthan 93% identity to the sequences of other Porina
species (Table 1).
Specimens examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along trail of Ohtomi-yuhodo, Iriomote Island,
Yaeyama Islands (N24°18′08″, E123°51′31″), on Asplenium setoi, 30 m
elev., 18 November 2019, K. Miyazawa 376 pr.p. & Y. Ohmura (incollection of Coenogonium subluteum) (TNS); along mountain path
between Ohtomi-guchi Entrance and Daiichi Yamagoya-ato, Iriomote
Island, Yaeyama Islands (N24°19′01'', E123°50′32''), on broad-leaf tree,
210 m elev., 19 November 2019, K. Miyazawa 400 pr.p. & Y. Ohmura(in collection of Byssoloma vanderystii) (TNS); along mountain path
between Daini Yamagoya-ato and Kambire-guchi Entrance, Iriomote
Island, Yaeyama Islands (N24°20′52″, E123°48′59″), on broad-leaf tree;
100 m elev., 20 November 2019, K. Miyazawa 504 & Y. Ohmura (TNS),
K. Miyazawa 517 & Y. Ohmura (TNS); along mountain path between
Itachikigawa River Junction and Daini Yamagoya-ato, Iriomote Island,
Yaeyama Islands (N24°20′52″, E123°49′18″), on broad-leaf tree, 100 m
elev., 20 November 2019, K. Miyazawa 511 pr.p. & Y. Ohmura (incollection of Porina cupreola var. nipponica) (TNS); along the trail by
Urauchi River from the landing point to Maryudo Waterfall, Iriomote
Island, Yaeyama Islands (N24°21′, E123°47′), on Arenga engleri, 60 m
elev., 9 March 2011, G. Tanaka 355 & Y. Ohmura (TNS); Haemida
Beach, Iriomote Island, Yaeyama Islands (N24°16′, E123°49′), on
Ipomoea sp., 10 m elev., 10 March 2011, Y. Ohmura 8029 pr.p. & G.
Tanaka (in collection of Coenogonium subluteum) (TNS).

Semigyalecta paradoxa Vain.
Fig. 4A
The Iriomote Island material exhibited thallicomposed of brown branched hyphae of the symbiotic
fungus and plates of symbiotic algae (Fig. 4A). The platesof symbiotic algae were composed of irregular andusually elongate cells, approximately 3–10 × 2.5–3 µm(Fig. 4A). This observation is more or less similar to thedescription provided by Santesson (1952) (~3–5 × 2–4µm). Although it is debated whether Semigyalecta
paradoxa is lichenized fungus (Santesson, 1952; Kalb
and Vězda, 1994; Thor et al., 2000; Wang and Wei, 2018),
our observation implies that it is lichenized.
Specimen examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along trail in the mountains along Urauchi river,161

Taiwania

Vol. 67, No. 1

Taketomi-cho, Iriomote Island, Yaeyama Islands (N24°19′01″,
E123°50′32″), on broad-leaf tree, 210 m elev., 19 November 2019, K.
Miyazawa 405 & Y. Ohmura (TNS).
Additional specimen examined. JAPAN. RYUKYUISLANDS (Kagoshima Pref.): 7 km SW of Nishinakama village, 100–
200 m N of Yakugachi River, near the road, Suniyo-cho, Oshima-gun,
Amami Islands, (N28°13′, E129°21′), on Quercus sp., 60–80 m elev., 5
February 1995, G. Thor 12987 (holotype, TNS).

ACKNOWLEDGMENTS
We thank Akiyo Naiki of the Tropical Biosphere ResearchCenter, University of the Ryukyus for providing us information
for the permit application for the field survey. We also thank thethree reviewers for carefully reading our manuscript and for theconstructive comments. Permissions for the collections on
Iriomote Island were kindly granted by the Naha Nature
Conservation Office, Kyushu Regional Environment Office, theMinistry of the Environment Government of Japan (No.
1911071), and the Okinawa forest management station, Forestry
Agency, the Ministry of Agriculture, Forestry and Fisheries
Government of Japan (No. 342).

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Fig. 4. Semigyalecta paradoxa and Tricharia kashiwadanii. A.
Thallus of S. paradoxa composed of brown branched hyphae ofthe mycobiont and plates of the photobiont (K. Miyazawa 436 &Y. Ohmura, TNS). B. Thallus with apothecia of T. kashiwadanii(holotype, TNS). C. Black hyphophore with diahyphae of T.
kashiwadanii (holotype, TNS). D. Black hyphophores on thethallus of T. kashiwadanii (holotype, TNS). Scales: A = 20 μm;
B = 0.5 mm; C = 100 μm; D = 0.2 mm.
Iriomote Island, Yaeyama Islands (N24°19′26″, E123°50′49″), on
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Tricharia kashiwadanii G. Thor, Lücking & Tat.
MatsumotoFig. 4B–D
Hyphophores and diahyphae of T. kashiwadanii were
found in the type specimen and in other material collectedin this study, although they were not described in the
protologue. Their features based on these specimens aredescribed as follows. Hyphophores are abundant, black,setiform with an acute apex and up to 0.3 mm tall (Fig.
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The BLAST results show less than 90% identity toGomphilloideae (including Tricharia) sequences in
GenBank (Table 1). This low homology is due to theabsence of registered sequences for this species in
GenBank.
Specimen examined. JAPAN. RYUKYU ISLANDS(Okinawa Pref.): along trail in the mountains along Urauchi river,

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