Bull. Natl. Mus. Nat. Sci., Ser. B, 38(3), pp. 87–92, August 22, 2012

Chapsa leprieurii, Ocellularia cavata and O. pyrenuloides(Graphidaceae, Lichenized Ascomycota) New to Japan
Andreas Frisch* and Yoshihito Ohmura
Department of Botany, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan*E-mail: frisch-a@kahaku.go.jp(Received 16 May 2012; accepted 26 June 2012)

Abstract Chapsa leprieurii (Mont.) Frisch, Ocellularia cavata (Ach.) Müll. Arg. and O. pyrenuloides Zahlbr. are reported from Iriomote and Kita-Iwo Islands and are additions to the lichen biota
of Japan. The species are described and illustrated, and information on their ecology and distribution is provided. The new combination Rhabdodiscus crassus (Müll. Arg.) Frisch is also made.
Key words : chemistry, description, ﬂoristics, Japan, morphology, Thelotremataceae, tropicalislands.

Introduction
The thelotremoid Graphidaceae (＝Thelotremataceae (Nyl.) Stiz.; Stizenberger, 1862) ofJapan are rather well known, particularly the species of Thelotrema s.l., which have been monographed by Matsumoto (2000). Fourty-two species are listed in the checklist of Japanese lichens(Kurokawa and Kashiwadani, 2006) which,according to the present classiﬁcation of the family (e.g., Frisch, 2006; Frisch and Kalb, 2006;
Rivas Plata et al., 2012), belong to the genera
Chapsa, Chroodiscus, Diploschistes, Ingvariella,
Leucodecton, Myriotrema, Ocellularia, Rhabdodiscus, and Thelotrema. Chroodiscus defectus
Papong & Lücking (Papong et al., 2009) andRhabdodiscus crassus (Müll. Arg.) Frisch [asStegobolus crassus (Müll. Arg.) Frisch in Frischand Kalb (2006); see below for the new combination] have recently been added to the Japaneselichen biota, while Thelotrema cinereum Müll.
Arg. [＝Myriotrema cinereum (Müll. Arg.) Hale]and Thelotrema sendaiense Vain. were placed in
the synonymy of Ocellularia microstoma (Müll.
Arg.) Frisch (Mangold et al., 2009; Ohmura,2012), and T. inalbescens Nyl. in the synonymy

of Rhabdodiscus ﬁssus Vain. (Mangold et al.,2009). Additional species can be expected
mainly in the subtropical islands in the south ofthe country. In this contribution we report three
species from Iriomote and Kita-Iwo Islands as
new to Japan, by which the total number of species in this country become six in Chapsa andﬁve in Ocellularia.
Material and Methods
This study is based on the herbarium specimens housed in the National Museum of Natureand Science (TNS) in Tsukuba, in which recent
materials collected by the authors are also
included.
The morphological descriptions of the species
were made using Olympus BX40 and Leica MZ8
microscopes on hand cuttings mounted in water,
1% aqueous iodine solution or 10% aqueous
KOH solution. Measurements were always madeon water preparations. At least 10 mature spores
were measured for each specimen with 1 μm precision. Paraphyses and periphysoids were measured with 0.5 μm precision. Chemical compounds were identiﬁed by means of thin layer

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Andreas Frisch and Yoshihito Ohmura

chromatography (TLC) using solvent C (Orange
et al., 2010).
Results and Discussion
Chapsa leprieurii (Mont.) Frisch, Biblioth.
Lichenol. 92: 105 (2006).
[Fig. 1A]
THALLUS cartilaginous, brown-olive, matt to
weakly shining, continuous to slightly ﬁssured,
uneven to shallowly warty, up to 0.2 mm thick;phenocortex 20–50 μm, moderately dense todense, strongly conglutinated, mainly of periclinal hyphae, often with inclusions of calcium oxalate crystals and decomposed bark cells; photobiont layer 10–100 μm, with calcium oxalate
crystals, largely endophloeodal; medulla not differentiated. PYCNIDIA unknown.
APOTHECIA dispersed or aggregated in loose
groups, level with the thallus, rounded to usuallyangular or shortly elongated, occasionally withshort branches, 0.6–2.5 mm broad and long; margin raised, rim-like, ± jagged to lobed, uprightto strongly recurved, with a prominent, whitefelty to crystalline inner surface; proper exciple
fused to partly free in the apical parts, low andrather thick, white-felty; disc pale brown, covered by thick white pruina. THALLINE MARGIN with phenocortex 15–40 μm thick; photobiont layer incuding the indistinct periderm layer
25–80 μm thick, with numerous calcium oxalate
crystals and decomposed periderm cells, only
weakly conglutinated. PROPER EXCIPLE cupular, 10–15 μm tall and wide, hyaline to pale
brown, of slightly conglutinated prosoplectenchymatic hyphae; periphysoids up to 40 μm
long, 2–3 μm wide, the tips moniliform, ﬁnely
adspersed with greyish to brownish granules.
HYMENIUM 70–90 μm tall, clear; ascospores
8/ascus, 1 seriate, brown, transversely septate,(2–)4 loculate, 10–12×5–6 μm, thick-walled,with subacute ends, I+ purplish-blue; asci narrowly clavate, 70–85×8–9 μm; paraphyses
1.5–2 μm wide, tips moniliform, ﬁnely adspersedwith greyish to brownish granules; epihymenium
greyish to pale brownish, 8–12 μm tall.

CHEMISTRY. No lichen compounds detectedby TLC.
ECOLOGY AND DISTRIBUTION. Chapsa
leprieurii is mainly collected in the canopy andmore open places in rainforests in coastal areas,
coastal forest and open coastal scrub up to 1250
m altitude. On Iriomote Island, C. leprieurii wasfound on the bark of a broad-leaf tree in aslightly shaded rainforest besides the brackish
river.
Chapsa leprieurii is a pantropical species,
being reported from tropical America, Africa,
India, Sri Lanka, The Philippines, Sumatra andAustralia (Awasthi, 1991; Frisch, 2006; Hale,
1981; Mangold et al., 2009). New to Japan.
NOTES. Chapsa leprieurii is easily recognized by the cartilaginous, olive-brown thallus;
the small and typically aggregated chroodiscoid
apothecia with prominently white-felty to crystalline margin; the brown, (2–)4 loculate ascospores 10–16×5–6 μm in size; and the absenceof lichen compounds. The specimens from Iriomote Island show a noticeable proportion of
2–3 loculate spores which are slightly smaller
than given in Frisch (2006: 12–16×5–6 μm) andat the lower end of the range reported by
Mangold et al. (2009) from Australia: 4–7(–8)×1(–2) loculate, 9–25(–28)×6–8(–11) μm.
Chapsa leprieurii has been included in Chapsa
based on the chroodiscoid apothecia with
periphysoids, the thick-walled ascospores andChapsa-type paraphysoids (Frisch, 2006).
Molecular data, however, indicate C. leprieurii tobe more closely related to Ocellularia s.l. than toChapsa (K. Kalb, pers. comm.). Further investigations are needed for a correct placement of thisspecies. For synonymy see Frisch (2006).
Specimens examined: JAPAN. Yaeyama
Islands (Pref. Okinawa). Iriomote Island, along
the stream at SE foot of Mt. Goza, Taketomi-cho,Yaeyama-gun, on bark of broad-leaf tree,
24°17′N, 123°49′E, 10 m a.s.l. 11. iii. 2011, Y.
Ohmura 8045, 8048 (TNS).

Chapsa leprieurii, Ocellularia cavata and O. pyrenuloides New to Japan

Fig. 1. Three Graphidaceae new to Japan. A. Chapsa leprieurii (Y. Ohmura 8045, TNS). B. Ocellularia cavata(Y. Ohmura 8056, TNS). C. O. pyrenuloides (Y. Ohmura 6367, TNS). Scales＝2 mm.

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Andreas Frisch and Yoshihito Ohmura

Ocellularia cavata (Ach.) Müll. Arg., Flora,
Jena 65: 499 (1882).
[Fig. 1B]
THALLUS corticolous, olive-grey, matt to
weakly shining, continuous, uneven to shallowly
warty, up to 0.2 mm thick; medulla of thallus
warts with pale fawn to deep yellow-orange pigment; phenocortex 5–12 μm, continuous, moderately dense, conglutinated, mainly of irregularhyphae, not always clearly separated from thephotobiont layer; photobiont layer 30–70 μm
thick, with numerous photobiont cells and someclusters of calcium oxalate crystals towards thebase; thallus warts ﬁlled with orange granules
and calcium oxalate crystals; medulla thin or
endophloeodal. PYCNIDIA not seen in the Japanese collection.
APOTHECIA dispersed, rounded, moderately
emergent, with ± steep ﬂanks and a broadly
rounded apex, 0.5–1.2 mm in diam.; margin shallowly warty, concolorous with the thallus; pore
0.2–0.3(–0.6) mm wide, rounded to angular, surrounded by an incomplete black ring formed by
the proper exciple; columella black, simple, withbulging tip, ﬁlling the pore; disc pale brown to
wax-coloured, covered by thin white pruina.
THALLINE MARGIN with phenocortex 3–8 μm
thick; photobiont layer 20–45 μm thick, an additional large cavity ﬁlled with orange granules andcalcium oxalate crystals often present below thephotobiont layer; periderm layer 50–60 μm thick,comprising a dense, brown conglutinatedprosoplectenchyme with periderm inclusions.
PROPER EXCIPLE cupular, 10–15 μm tall andhyaline to pale brown at the base, 50–100 μm
wide laterally, strongly carbonised at least in theupper half; columella conical, with a bulging tip,
carbonised throughout. HYMENIUM 100–
120 μm tall, clear; subhymenium 10–15 μm tall;ascospores 8/ascus, 1–2 seriate, hyaline, transversely septate, 6–9 loculate, 21–27×6–7 μm,
with subacute to rounded ends, I+ purplish-blue;
asci narrowly clavate, 90–100×10–14 μm;
paraphyses 1.5–2 μm wide; tips with a single
slightly thickened cell, adspersed with ﬁne greyish to brownish granules; epihymenium unpi-

mented, 5–8 μm tall.
CHEMISTRY. Cinchonarum unknown (seeFrisch 2006; major), unknown 1 (Rf value 5, paleyellow, minor), unknown 2 (Rf value 10, orangebrown, trace), unknown 3 (Rf value 26, grey tobrown, trace), and orange pigment (Rf value 17)
were detected by TLC. The orange pigment is
K+ purple.
ECOLOGY AND DISTRIBUTION. Ocellularia cavata is a common pantropical species,
being collected in a wide range of tropical rain
and cloud forests from sea-level to 2400 m altitude. On Iriomote Island, it was found on thebark of a broad-leaf tree in a slightly shaded rainforest besides the brackish river.
Ocellularia cavata has been reported fromtropical America, Africa, Sri Lanka, Australiaand Hawaii (Frisch, 2006; Hale, 1974, 1978;
Magnusson and Zahlbruckner, 1943; Mangold etal., 2009; Redinger, 1936; Sipman, 1992). New
to Japan.
NOTES. Ocellularia cavata is characterizedby moderately to strongly emergent carbonizedapothecia with a conical columella; pale straw toorange pigment concentrated in warts on thethallus and thalline margin of the apothecia;
hyaline, transversely septate, I+ purplish-blue
ascospores of moderate size (6–16 loculate, 20–
60×6–11 μm); and the cinchonarum unknown
and an unknown orange pigment as lichen compounds. On the world level, O. cavata is a polymorphic species regarding ascospore size, pigmentation and apothecia morphology, andobviously represents a complex of closely related
taxa. With respect to the small ascospores andthe deep orange, K+ purple pigment of the
medulla, the specimen from Iriomote Islandshows afﬁnities to the type of O. punctulata(Leight.) Zahlbr. from Sri Lanka. However, therelationship of this species and the similar andolder O. obturata (Ach.) Spreng. from Sierra
Leone is not clear and needs further study. Until
a thorough revision of O. cavata s.l. on the worldlevel has been done, we prefer to use the broad
circumscription of this species as adopted by
Frisch (2006).

Chapsa leprieurii, Ocellularia cavata and O. pyrenuloides New to Japan

Specimen examined: JAPAN. Yaeyama
Islands (Pref. Okinawa). Iriomote Island, along
the stream at SE foot of Mt. Goza, Taketomi-cho,Yaeyama-gun, on bark of broad-leaf tree,
24°17′N, 123°49′E, 10 m a.s.l. 11. iii. 2011, Y.
Ohmura 8056 (TNS).
Ocellularia pyrenuloides Zahlbr., in Magnusson
and Zahlbruckner, Ark. Bot. 31A(1): 46 (1944).
[Fig. 1C]
THALLUS pale greenish-grey, weakly glossy,smooth to indistinctly verruculose, continuous,
up to 0.07 mm thick, the bark substrate shining
through the thallus surface; phenocortex 3–7 μm
thick or indistinct, loosely organized to moderately dense, mostly not clearly separated fromthe photobiont layer, formed from irregularhyphae; photobiont layer 40–60 μm thick, withlarge complexes of calcium oxalate crystals,
largely endophloeodal; medulla endophloeodal.
PYCNIDIA unknown.
APOTHECIA dispersed, rounded, moderately
emergent, 0.5–0.9 mm in diam.; margin concolorus with the thallus, thin, entire to somewhat ﬁssured, forming a 0.3–0.4 mm wide, rounded toangular pore lined by a brown to black ring of
the exposed proper exciple and periderm layer;columella broad, ﬂat to slightly bulging,±ﬁlling
the pore, black, tipped with white. THALLINEMARGIN with phenocortex 3–5 μm thick toabsent; photobiont layer 20–35 μm thick; periderm layer 15–50 μm thick, moderately dense,comprising a brown conglutinated prosoplectenchyme with periderm inclusions and large
complexes of calcium oxalate crystals. PROPER
EXCIPLE cupular, 7–10 μm tall and hyaline topale brown at the base, 25–60 μm wide laterally,
carbonised, with a thin apical layer of hyaline
hyphae; columella broad trunk-shaped, carbonised, simple, ﬂat, with a thin apical layer ofhyaline hyphae. HYMENIUM 90–110 μm tall,clear, subhymenium 8–10 μm tall; ascospores
8/ascus, 2 seriate, hyaline to pale olivish, transversely 6–8 loculate, 21–25×6–7 μm, with subacute to rounded ends, I+ purplish-blue. Ascinarrowly clavate, 80–105×10–14 μm; paraphy-91

ses 1.5–2 μm wide, tips with a single slightly
thickened cell, adspersed with ﬁne greyish tobrownish granules; epihymenium unpigmented,
8–10 μm tall.
CHEMISTRY. Stictic acid (major), cryptostictic acid (trace) and constictic acid (trace) weredetected by TLC.
ECOLOGY AND DISTRIBUTION. Ocellularia pyrenuloides has been collected in Pandanus forest in the Hawaii Islands and is further
reported from lowland to submontane rainforests
in tropical America, Africa, Australia and Hawaii(Frisch, 2006; Hale, 1974, 1981; Magnusson and
Zahlbruckner, 1943; Mangold et al., 2009). The
specimen from Kita-Iwo Island was collectedfrom Pandanus boninensis in a lowland rainforest. New to Japan.
NOTES. Ocellularia pyrenuloides is an inconspicuous and somewhat nondescript species thatcan be characterized by the thin to endophloeodal, pale greenish-grey to fawn thallus without
clearly deﬁned phenocortex; small immersed tomoderately emergent apothecia of 0.3–0.9 mm
diam., with strong carbonization and a broad,ﬂat, and typically white-tipped columella; hyaline to pale olive, I+ purplish blue ascospores,
6–8 loculate, 18–25×6–8 μm in size; and thestictic acid chemosyndrome. Species of thehabitually similar the genus Clandestinotrema(Rivas Plata et al., 2012) can be distinguished by
the I-negative ascospores of the clandestina-type(Frisch, 2006). The specimen from Kita-Iwo
Island is morphologically close to the type collections from Hawaii.
Specimen examined: JAPAN. Sulphur
Islands. Kita-Iwo Island, on bark of Pandanus
boninensis, 25°26′13.2″N, 141°17′14.6″E, 186 ma.s.l. 17. vi. 2009, Y. Ohmura 6367 (TNS).
Rhabdodiscus crassus (Müll. Arg.) Frisch,comb. nov.
Basionym: Leptotrema crassum Müll. Arg., Flora
65: 332 (1882). — Stegobolus crassus (Müll.
Arg.) Frisch, in Frisch and Kalb, Biblioth.
Lichenol. 92: 448 (2006).
This species had been included in the

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Andreas Frisch and Yoshihito Ohmura

Stegobolus auberianus group by Frisch and Kalb
(2006), which is now accepted as the separate
the genus Rhabdodiscus Vain. (Vainio, 1921) following recent molecular research in the family
Graphidaceae (e.g. Rivas Plata and Lumbsch,
2011; Rivas Plata et al., 2012). Therefore, the
new combination Rhabdodiscus crassus is made.
R. crassus has been reported from Japan basedon the type specimen of Leptotrema oleosum
Zahlbr., collected on Yakushima Island by Abbé
Faurie in 1906 (Frisch and Kalb, 2006).
Acknowledgments
This study is partly funded by the Grant-inAid relating to JSPS Postdoctoral Fellowship for
Foreign Researchers (no. 23-01706). We arethankful to an unknown reviewer for helpful
information on the medulla pigmentation in O.
cavata s.l. and the possible identity of the Japanese collection with O. punctulata.
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