Bull. Natl. Mus. Nat. Sci., Ser. B, 42(1), pp. 1–18, February 22, 2016

Lichens of Mikura-jima Island, Izu Islands, Central Japan
Yoshihito Ohmura1,*, Andreas Frisch2, James C. Lendemer3 and Minoru Nakanishi4
1

Department of Botany, National Museum of Nature and Science,
Amakubo 4–1–1, Tsukuba, Ibaraki 305–0005, Japan2
Department of Ecology, Swedish University of Agricultural Sciences,
P.O. Box 7044, SE–750 07 Uppsala, Sweden
3
Institute of Systematic Botany, New York Botanical Garden, Bronx, NY, 10458–5126, U.S.A.
4
Hiroshima University, Kagamiyama 1–1–1, Higashihiroshima, Hiroshima 739–8524, Japan*E-mail: ohmura-y@kahaku.go.jp(Received 15 November 2015; accepted 22 December 2015)

Abstract Lichens of Mikura-jima Island, Izu Islands, central Japan, were investigated. As theresults of taxonomic study, 78 taxa (77 species and 1 variety) were recognized from the island.
Among them, Graphis mikuraensis Y. Ohmura & M. Nakan. is described as a new species. Lepraria ﬁnkii and L. lobiﬁcans, based on the emended species concept, were conﬁrmed to occur inJapan. A new combination, Rhabdodiscus inalbescens (Nyl.) Frisch is proposed. Phytogeographic
discussion is made for the listed species. The lichen ﬂora of Mikura-jima Island is characterized by
the high percentage of pantemperate element species together with tropical and subtropical species. A remarkable range extension is noted for Physma radians, and the ﬁnd represents the northernmost locality for this paciﬁc element species. Nipponoparmelia laevior, which was common
before 1981, may have become extinct or drastically reduced in this island during the past 30
years.
Key words : inventory, lichenized fungi, new record, new species, phytogeography, taxonomy.

Mikura-jima Island, one of the Izu Seven
Islands, is located ca. 200 km south of central
Tokyo in the Paciﬁc Ocean (N33°52′, E139°36′);
its shape is roughly circular (ca. 5 km in diameter), and the highest peak, Mt. Oyama, reaches
851 m elev. (Fig. 1). This island is a stratovolcano but the last volcanic activity occurred
around 6,000 years ago.
The annual precipitation is more than 5,000 mm(Hamada, 2001). Monthly average temperatures
range from 5.4°C to 24.1°C at 300 m elev., 3.8°C
to 23.0°C at 500 m elev., and 2.1°C to 21.7°C at
720 m elev. (Hoshino et al., 1995).
The natural vegetation is dominated by Castanopsis sieboldii (Makino) T.Yamaz. & Mashibaand Machilus thunbergii Sieb. & Zucc. at lower
elevations, Castanopsis sieboldii at higher elevations, and a mixed forest with Trochodendron

aralioides Sieb. & Zucc., Daphniphyllum macropodum Miq., Cerasus speciosa (Koidz.) H.
Ohba, Dendropanax triﬁdus (Thunb.) Makino,
etc. in the plateau at the top of the island (Ohba,
1971).
From a phytogeographic point of view, it
should be noted that subalpine species of vascular plants [e.g. Maianthemum dilatatum (A.W.
Wood) A.Nelson & J.F.Macbr., Viburnum furcatum Blume ex Maxim., and Rhus ambigua
Lavall. ex Dipp. (Ohba, 1971)] and bryophytes[e.g. Pleurozium schreberi (Brid.) Mitt., Campylopus fragilis (Bridel) Bruch & Schimper, Dicranum viride var. hakkodense (Cardot) Takaki, andGollania varians (Mitt.) Broth. (Higuchi andNishimura, 2001)] have been reported from thisisland, although they are usually found at elevations higher than 1,000 m in central Honshu and/

Yoshihito Ohmura et al.

2

Fig. 1.

Map showing investigated sites in Mikura-jima Island.

or on Hokkaido, northern Japan.
The lichen ﬂora of Mikura-jima Island has
been poorly known and the ﬁrst report of 11 species was made by Asahina (1955) based on thespecimens collected by Dr. Hisashi Tsuyama in
August 1952. Subsequently two additional species and one variety were reported in the area
(Asahina, 1969; Kashiwadani and Kurokawa,
1981; Miyawaki, 1988). As part of an environmental educational project organized by the
Mikurajima-mura Administration and the Nonproﬁt Organization Nature center risen (Mikura
Museum, 2012), the ﬁrst author had opportunities to collect lichens in 2010 and 2011. The purpose of this study was to inventory the lichens of
Mikura-jima Island and discuss the species fromthe phytogeographic point of view.
Materials and Methods
Field surveys were carried out by the ﬁrst
author on 20–22 December 2010 and 10–13
November 2011. The locality data of the collection sites are shown in Fig. 1. In addition to theﬁrst authorʼs collections, herbarium specimensfrom Mikura-jima Island housed in the National
Museum of Nature and Science (TNS) were also
used for the present study. A total of 154 specimens were examined. All collections examined
are housed in TNS unless otherwise indicated.
Morphological observations were made using
Olympus SZ61 and BX51 microscopes on handLichens of Mikura-jima Island, Japan

cuttings mounted in GAW solution (glycerin : ethanol : water＝1 : 1 : 1). Chemical substances were detected by means of thin layer
chromatography (TLC) with solvents A, B′ and/or C (Orange et al., 2010).
Collection sites
The data for collection sites (A–P) investigated by the ﬁrst author and the collection dates(d/m/y) are provided as below. The symbols(A–P) correspond to those in Fig. 1. For the
specimens of other collectors, the locality information on the specimen labels is shown in thelist of species.
A: Tokurine, Mikura-jima Island, 20 m elev.
10/11/2011
B: Maenohama, Mikura-jima Island, 10 m
elev. 21/12/2010
C: Inane Shrine, Sato, Mikura-jima Island,
100–120 m elev. 22/12/2010 or 12,
13/11/2011
D: Sato, Mikura-jima Island, 165 m elev.
13/11/2011
E: ca. 1.5 km E of Sato, Mikura-jima Island,
300 m elev. 13/11/2011
F: along the trail, NW slope of Mt. Oyama,
Mikura-jimaIsland,550 m
elev.
21/12/2010 or 10/11/2011
G: along the trail, NW slope of Mt. Oyama,
Mikura-jima Island, ca. 600 m elev.
21/12/2010
H: Suzuhara Shitsugen, N slope of Mt.
Oyama, Mikura-jima Island, ca. 700 m
elev. 21/12/2010
I: N slope of Mt. Oyama, Mikura-jimaIsland, ca. 750 m elev. 21/12/2010
J: along the Prefectural Road 223, NE slopeof Mt. Nagataki, Mikura-jima Island,
680 m elev. 21/12/2010
K: NE slope of Mt. Nagataki, Mikura-jimaIsland, ca. 700 m elev. 21/12/2010 and
10/11/2011
L: around the top of Mt. Nagataki, Mikurajima Island, ca. 800 m elev. 21/12/2010
M: along the Prefectural Road 223, W slope of3

Mt. Oyama, Mikura-jima Island, 510 m
elev. 20/12/2010
N: along the trail of Mt. Kurosaki-Takao,
Mikura-jima Island, ca. 500 m elev.
20/12/2010
O: Mt. Kurosaki-Takao, Mikura-jima Island,
530 m elev. 20/12/2010
P: Inane Shrine, Mt. Kurosaki-Takao,
Mikura-jimaIsland,370 m
elev.
20/12/2010
Results and Discussion
Based on the results of taxonomic study andthe reference information, 78 taxa (77 species
and 1 variety) are reported from the present area.
They are arranged alphabetically in the list
below. Some specimens related to Adelolecia,
Enterographa, Lecanora, Lecidella, Ocellularia,
Parmentaria, Porina, Strigula, and unidentiﬁed
specimens even at the genus level are notincluded in the list as further taxonomic revisions
are required for their complete species-level
identiﬁcation.
Abbreviations for collectors are as follows.
HM: H. Miyakawa, JH: J. Hasegawa, MH: M.
Higuchi, NN: N. Nishimura, NS: N. Satomi, YD:
Y. Doi, and YO: Y. Ohmura. Numeral following
collector abbreviation indicates the collection
number of MH, NN or YO.
List of species1). Agonimia paciﬁca (H.Harada) Diedrich
Specimen examined. K: YO&HM-7815(on bark of Buxus microphylla var. japonica).
2). Caloplaca spodoplaca (Nyl.) Zahlbr.
Specimen examined. B: YO&HM-7844(on rock).
3). Cladonia coniocraea (Flörke) Spreng.
This species was already reported fromthe present area by Asahina (1955), and it
was collected again during the current
investigations.
Specimen examined. K: YO&HM-7825

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Yoshihito Ohmura et al.

(on trunk of Buxus microphylla var. japonica).
4). Cladonia fruticulosa Kremp.
Specimens examined. G: YO&HM-7838(on humus). N: YO&HM-7781 (on soil).
5). Cladonia furcata (Huds.) Schrad.
Specimens examined. J: YO&HM7809(over humus on sloping road-bank). K:
YO-8447 (over humus on sloping roadbank).
6). Cladonia kurokawae Ahti & S.Stenroos
This species was reported as C. conistea(Delise) Asahina (nom. illegit.) by Asahina(1955), and it was collected again during thecurrent investigations.
Specimens examined. C: YO-8542, 8543
(over mosses on stone wall; 12/11/2011).
7). Cladonia macilenta Hoffm.
Specimen examined. N: YO&HM-7803(on decayed stump).
8). Cladonia merochlorophaea Asahina
Specimen examined. L: YO&HM-7829(on decayed wood).
9). Cladonia ochrochlora Flörke
Specimens examined. I: YO&HM-7830(on decayed wood). N: YO&HM-7798 (ontrunk base).
10). Cladonia ramulosa (With.) J.R.Laundon
Specimens examined. A: YO-8534, 8537
(on soil). C: YO-8541 (on soil on stonewall; 12/11/2011). J: YO&HM-7808, 7813
(on soil on sloping road-bank).
11). Cladonia scabriuscula (Delise) Leight.
Specimens examined. F: YO&HM-7839(on soil; 21/12/2010). J: YO&HM-7807 (onsoil on sloping road-bank).
12). Cladonia submultiformis Asahina
Specimen examined. J: YO&HM-7810(on soil on sloping road-bank).
13). Coccocarpia erythroxyli (Spreng.) Swinscow & Krog
Specimens examined. C: YO&HM-7855(on rock; 22/12/2010). L: YO&HM-7828(on bark of Pittosporum tobira). N:
YO&HM-7786 (on bark of Ternstroemia
gymnanthera).

14). Coccotrema porinopsis (Nyl.) Imshaug ex
Yoshim.
Specimens examined. H: YO&HM-7836(on bark of Prunus sp.). I: YO&HM-7833(on decayed wood).
15). Collema japonicum (Müll.Arg.) Hue
Specimen examined. C: YO&HM-7851(on rock; 22/12/2010).
16). Collema subﬂaccidum Degel.
Specimens examined. C: YO-8548 (onstone wall; 12/11/2011), YO&HM-7847 (onbark of Cinnamomum camphora; 22/12/
2010).
17). Cresponea proximata (Nyl.) Egea & Torrente
Specimens examined. C: YO&HM-7848(on bark of Castanopsis sieboldii;
22/12/2010). D: YO-8561 (on bark of Castanea crenata).
18). Dibaeis sorediata Kalb & Gierl
Specimen examined. F: YO-8532 (onsoil; 10/11/2011).
19). Diorygma soozanum (Zahlbr.) M.Nakan. &Kashiw.
Specimens examined. C: YO&HM-7849(on bark of Castanopsis sieboldii;
22/12/2010). D: YO-8564 (on bark of Castanea crenata).
20). Enterographa leucolyta (Nyl.) Redinger
Specimens examined. A: YO-8538 (onrock). B: YO&HM-7845 (on rock).
21). Fissurina inabensis (Vain.) M.Nakan. &Kashiw.
Specimens examined. P: YO&HM-7796(on bark of Camellia japonica). Without
detailed locality, YD-s.n. (on bark ofCamellia japonica; 29/9/1978).
22). Graphis assimilis Nyl.
This species has been confused with G.
intricata Fée by Japanese lichenologists(Kurokawa and Kashiwadani, 2006). However, G. intricata has elongate lirellae andshorter spores (15–30 μm), while G. assimilis has short lirellae and longer spores (30–45 μm) (Lücking et al., 2009).
Specimens examined. D: YO-8568 pr.

Lichens of Mikura-jima Island, Japan

maj.p. (on bark of Ginkgo biloba). N:
YO&HM-7791 (on bark of decayed wood).
Along the route between Sato and Mt.
Oyama, 250 m elev., MH-10252 (on trunk;
14/7/1981).
23). Graphis cleistoblephara Nyl.
Specimens examined. D: YO-8565 (onbark of Castanea crenata), YO8568 pr.p.,
YO8569 (on bark of Ginkgo biloba).
24). Graphis hossei Vain.
This species was known as G. connectens
Zahlbr. for Japanese lichenologists (Kurokawa and Kashiwadani, 2006), which wassynonymized into G. hossei by Lücking etal. (2009).
Specimens examined. K: YO&HM-7816(on bark of Prunus sp.). Along the route
between Sato and Mt. Oyama, 440 m elev.,
MH-10294A (on tree trunk; 14/7/1981).
25). Graphis japonica (Müll.Arg.) A.W.Archer& LückingThis species was reported in Nakanishi(1966), but later, it was treated as a synonym of G. ﬁssofurcata Leight. by Nakanishi (1974). According to Lücking et al.
(2009), G. japonica contains 2–4 spores per
ascus while G. streblocarpa (Bél.) Nyl.
(syn. G. ﬁssofurcata) contains one spore per
ascus. These features are considered to bestable and G. japonica should be treated asa distinct species.
Specimens examined. N: YO&HM-7789(on bark of Ternstroemia gymnanthera),
YO&HM-7782 (on bark of Daphniphyllum
teijsmannii). O: YO&HM-7799 pr.maj.p.
(on bark of Castanopsis sieboldii).
26). Graphis longiramea Müll.Arg.
This species has been known as Graphis
awaensis Vain. by Japanese lichenologists(Kurokawa and Kashiwadani, 2006). It wasplaced in synonymy with G. longiramea(Lücking et al., 2009). The specimens collected in Mikura-jima Island have sporeswith 62–90×11–12 μm and 15–20-locules,
which agree with the feature of G. longiramea.

5Specimens examined. I: YO&HM-7831(on decayed wood). Along the route
between Sato and Mt. Oyama, 350 m elev.,
MH-10281 (on decaying wood; 14/7/1981).
27). Graphis mikuraensis Y.Ohmura & M.
Nakan., sp. nov.
Mycobank No.: MB 815136
Type: Japan, Izu Islands (Tokyo Metropolis): along the route between Sato and Ienosawa, Mikura-jima Island, on tree branch,elevation 500 m, 15 July 1981, M. Higuchi
10474 (Holotype, TNS).
This species is similar to Graphis gonimica Zahlbr. and G. sapii Zahlbr. but differsfrom G. gonimica in the larger spore andfrom G. sapii in lirella morphology.
[Fig. 2]
Thallus corticolous, off-white to palefawn, smooth, dull. Ascomata conspicuous,
numerous, crowded, sessile, erumpent, simple or sparsely branched, straight or curved,
terminally rounded, with lateral thalline
margin, up to 4 mm long, 0.3–0.4 mm wide;
labia black, entire, disc concealed, non-pruinose. Proper exciple completely carbonized, 40–90 μm thick at the base. Hymenium
92–142 μm thick, inspersed with oil droplets. Ascospores, 8 per ascus, colorless,transversely 10–14-locular, (35–)43.7±
5.7(–54)×(6.7–)8.1±0.8(–10) μm (n＝18).
Chemistry: norstictic acid and an unidentiﬁed substance (Rf class 2, same position
with stictic acid, in solvent Bʼ and Rf class
1, lower than salazinic acid, in solvent C;
brownish yellow after heating, 110°C,
10 min. with 10% H2SO4) as the major substances.
Etymology: the epithet refers to the type
locality, Mikura-jima Island.
This species is characterized by 1) the
erumpent lirellae with black, entire and concealed labia and lateral thalline margin,
which are up to 4 mm and simple to sparsely
branched with rounded terminals [“lineolamorph” sense Lücking (2009) and Lücking
et al. (2009)] (Fig. 2A), 2) the completely

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Yoshihito Ohmura et al.

Fig. 2. Graphis mikuraensis Y.Ohmura & M.Nakan. (Holotype, TNS). A. Lirellae. B. Cross section of apothecium. C. Spores. Scales: A＝1 mm, B＝100 μm, C＝10 μm.

carbonized proper exciple (Fig. 2B), 3) the
inspersed hymenium, 4) the transversely
septate spores, 10–14-locular and 35–54×
6.7–10 μm in length (Fig. 2C), and 5) the
presence of norstictic acid.
Graphis mikuraensis resembles G. gonimica Zahlbr. (W-lectotype!) and G. sapii
Zahlbr. (W-lectotype!), which were both

described from China. However, G. mikuraensis is distinguished by lager spores thanin G. gonimica [22–32×5–9 μm in length, 8
locules in Zahlbruckner (1932: 431)]; andby the lineola-morph lirellae with rounded
terminals from G. sapii which has elongate
and irregularly branched lirellae with acuteterminals [“deserpense-morph” sense Lück-

Lichens of Mikura-jima Island, Japaning (2009) and Lücking et al. (2009)]
although the range of spore size and septation in G. mikuraensis entirely overlaps
with that of G. sapii [40–44×6–7 μm in
length, 12 locules in Zahlbruckner (1930:40)]. It should be noted that Lücking et al.
(2009) treated G. sapii as a synonym of G.
gonimica, but we consider them as a distinct
species according to the morphological differences mentioned above. Graphis mikuraensis is also similar to G. desquamescens(Fée) Zahlbr. which occurs in tropical andadjacent temperate regions including southern Japan. However, G. mikuraensis has alateral thalline margin on the lirellae, while
G. desquamescens lacks a lateral thalline
margin [“hossei-morph” sense Lücking(2009)].
Graphis mikuraensis has been found only
on Mikura-jima Island, and is possibly an
endemic species to Japan. It was collected
on tree bark at elevations between 440 and
500 m along a stream valley.
Specimen examined (except type). En
route from Sato to Mt. Oyama, 440 m elev.,
MH-10294B (on tree trunk; 14/7/1981).
28). Graphis scripta (L.) Ach.
Specimens examined. D: YO-8563 (onbark of Castanea crenata). En route from
Sato to Mt. Oyama, 140 m elev., MH-10227(on trunk of Camellia japonica; 14/7/1981);
the same locality, 250 m elev., MH-10252(on trunk; 14/7/1981).
29). Graphis tsunodae Zahlbr.
Graphis tsunodae was described by Zahlbruckner (1916: 47) based on a specimen
collected from Mt. Akagi in Japan. Nakanishi (1966) considered it as a synonym of G.
proserpens Vain. However, Lücking et al.
(2009) distinguished G. tsunodae from G.
proserpens by having prominent sessile lirellae which lack thalline margins or havepoorly developed thalline margins only atthe base, while G. proserpens has erumpent
lirellae with distinct lateral thalline margins.
Moon et al. (2012) also reported G.

7tsunodae from Korea followed by Lücking
et al. (2009). The features for the collections
from Mikura-jima Island agree with G.
tsunodae, although the occurrence of G.
proserpens in Japan should be carefully
revised by further research.
Specimens examined. L: YO&HM-7827(on bark of Pittosporum tobira). O:
YO&HM-7799 pr.p. (on bark of Castanopsis sieboldii). En route from Ohshimawake
River to Nangou, 450 m elev., MH-10554(on trunk of Buxus microphylla var. japonica; 16/7/1981).
30). Hemithecium oshioi (M.Nakan.) M.Nakan.
& Kashiw.
Specimen examined. P: YO&HM-7794(on bark of Camellia japonica).
31). Herpothallon japonicum (Zahlbr.) G.Thor
Specimen examined. E: YO-8560 (onbark of Cryptomeria japonica).
32). Heterodermia diademata (Taylor) D.D.
Awasthi
Specimen examined. K: YO&HM-7814(on bark of Buxus microphylla var. japonica).
33). Heterodermia pandurata (Kurok.) J.C.Wei
Specimens examined. Mt. KurosakiTakao-san, NS-s.n. (on twigs of Buxus
microphylla var. japonica; 3/10/1963).
Between Ienosawa River to Inane Shrine,
500 m elev., NN-8350 (on trunk; 17/7/1981).
34). Hypotrachyna exsecta (Taylor) HaleThis species was reported in Asahina(1955), and it was collected again during thecurrent investigations.
Specimen examined. K: YO&HM-7821(on bark of Buxus microphylla var. japonica).
35). Hypotrachyna koyaensis (Asahina) Hale
Specimen examined. K: YO&HM-7818,
7822 (on bark of Buxus microphylla var.
japonica).
36). Lecanora leprosa Fée
D: YO-8567 (on bark of Castanea crenata).
37). Lecanora mikuraensis H.Miyaw.

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Yoshihito Ohmura et al.

This species was reported in Miyawaki(1988). It was collected at 380 m elev. without detailed locality (MH-10441, HIRO, notseen).
38). Lecanora subimmergens Vain.
Specimen examined. C: YO-8546 (onstone wall; 12/11/2011).
39). Lepraria cupressicola (Hue) J.R.Laundon
Specimen examined. C: YO-8554 (onbark of Cryptomeria japonica; 12/11/2011).
40). Lepraria ﬁnkii (de Lesd.) R.C.Harris,
Evansia 2: 45. 1985.
Basionym: Crocynia ﬁnkii de Lesd., Bull.
Soc. Bot. France 71: 334. 1924. Type:
U.S.A., Iowa, Fayette Co., Fayette, 7.
vii.1894, on moss and limestone, B. Fink
s.n. (NY!, lectotype; COLO!, DUKE!, NY!,
isolectotypes).
See description and illustrations in Lendemer (2013).
This species is characterized by the thickgranular crustose thallus without crisped
margins, and the presence of atranorin, zeorin, norstictic acid, stictic acid aggregate,
and ±fatty acid.
The specimen from Mikura-jima Island
agrees with the type specimens and description of Lendemer (2013). New to Japan.
Lepraria ﬁnkii has historically been confused with L. lobiﬁcans Nyl. which is a distinct and chemically variable species (Lendemer, 2013). As L. ﬁnkii was found together
with L. lobiﬁcans Nyl. on Mikura-jimaIsland, the Japanese materials which wereformerly identiﬁed as L. lobiﬁcans should
be carefully revised in further researches.
Specimen examined. N: YO&HM-7800(on bark of Daphniphyllum teijsmannii).
41). Lepraria lobiﬁcans Nyl., Flora 56: 196.
1873.
Type: France, Collioure, Pyrénées-Orientales, 4.vii.1872, on soil, W. Nylander s.n.
(H-NYL-10864!, lectotype; H-NYL-10865!,
isolectotype).
See synonyms in Lendemer (2013).
See descriptions and illustrations in Cre-

spo et al. (2006), Saag et al. (2007), Tretiach et al. (2009) and Lendemer (2013).
This species is characterized by its thickgranular thallus (Leproloma-like thallus)
with crisped margins and the presence of
atranorin and a chemosyndrome of stictic
acid.
Seven chemotypes have been reported forthe species (as L. santosii; Tretiach et al.,2009): (1) atranorin, roccellic acid, zeorin,
stictic and constictic acids; (2) atranorin,
UV– vivid pink unknown, stictic and constictic acids; (3) atranorin, roccellic acid,stictic and constictic acids; (4) atranorin,roccellic acid, stictic and constictic acids(± or ＋), fumarprotocetraric or protocetraric acid; (5) atranorin, roccellic acid,protocetraric acid; (6) atranorin, roccellicacid, protocetraric acid, fumarprotocetraricacid; (7) atranorin, roccellic acid, fumarprotocetraric acid. The specimens collected from
Mikura-jima Island are related to the “chemotype (1)”, but they contain ±different
fatty acid (not roccellic acid nor rangiformicacid).
Lepraria lobiﬁcans Nyl. in the sense of
the type (Lendemer, 2013) is conﬁrmed tooccur in Japan. Previous occurrences of thisspecies were in France and Canary Islands(Crespo et al., 2006; Lendemer, 2013). The
distribution is now extended to Japan but
further careful examinations would expand
the distribution. Given the difﬁculty
involved in delimitation within L. lobiﬁcans-L. isidiata the identity of the Japanese
populations should also be conﬁrmed withmolecular data in the future (Tretiach et al.,2009).
Specimens examined. A: YO-8535 (onrock). C: YO-8557 (on stone wall;
12/11/2011).
42). Leptogium azureum (Sw.) Mont.
Specimens examined. C: YO&HM-7850,
7852 (on rock; 22/12/2010).
43). Leptogium cyanescens (Pers.) Körb.
Specimens examined. N: YO&HM-7788

Lichens of Mikura-jima Island, Japan(on bark of Ternstroemia gymnanthera),
YO&HM-7801 (on bark of Daphniphyllum
teijsmannii).
44). Leptogium pichneum (Ach.) Nyl.
This species was reported by Asahina(1955), but the voucher specimen was notfound in TNS. In Japan, Leptogium pichneum is a common species at low elevations, and is usually found on stonewalls
and tree trunks in humid habitats. It mightoccur in Mikura-jima Island, but was notcollected during the current investigations.
45). Lobaria discolor (Bory) Hue
Specimens examined. H: YO&HM-7835(on bark of Trochodendron aralioides). N:
YO&HM-7787 (on bark of Ternstroemia
gymnanthera).
46). Lobaria retigera (Bory) Trevis.
This species was reported in Asahina(1955), but the voucher specimen could not
be found in TNS. Lobaria retigera isbroadly distributed from Hokkaido to
Ryukyu in Japan, and might occur in
Mikura-jima Island. However, it was notfound during the present investigations.
47). Myelochroa entotheiochroa (Hue) Elix &HaleSpecimens examined. C: YO&HM-7854(on rock; 22/12/2010), YO-8547 (on stonewall; 12/11/2011).
48). Nipponoparmelia laevior (Nyl.) A.Crespo& al.
Although four voucher specimens arepreserved in TNS, this species could not befound even at the same locality during thisinvestigation.
Specimens examined. Between Ienosawa
Riv. and Inane Shrine, ca. 500 m elev.,
NN-8349 (on trunk; 17/7/1981). Mt. Kurosaki-Takao-san, NS-s.n. (3/10/1963). Mt.
Oyama (label as “Mt. Misen”), NS-s.n.
(4/10/1963). Without detailed locality, NSs.n. (7/10/1963).
49). Normandina pulchella (Borrer) Nyl.
This species was reported in Asahina(1955), and it was collected again during the9current investigations.
Specimens examined. C: YO&HM-7853(on mosses over rock; 22/12/2010). K:
YO&HM-7823 (on bark of Buxus microphylla var. japonica).
50). Ocellularia microstoma (Müll.Arg.) Frisch
Specimen examined. I: YO&HM-7832
pr.maj.p. (on decayed wood).
51). Ochrolechia parellula (Müll.Arg.) Zahlbr.
Specimen examined. C: YO-8572 (onrock; 13/11/2011).
52). Parmelinopsis minarum (Vain.) Elix &HaleSpecimen examined. I: YO&HM-7834(on decayed wood).
53). Parmelinopsis spumosa (Asahina) Elix &HaleSpecimens examined. C: YO&HM-7846(on bark of Cinnamomum camphora;
22/12/2010), YO-8556 (on bark of Cryptomeria japonica; 12/11/2011). K: YO&HM7817 (on bark of Prunus sp.), YO&HM7826 (on trunk of Trochodendron
aralioides).
54). Parmotrema tinctorum (Despr. ex Nyl.)HaleSpecimen examined. C: YO-8553 (onbark of Cryptomeria japonica; 12/11/2011).
55). Peltigera dolichorrhiza (Nyl.) Nyl.
This species was reported in Asahina(1955), but the voucher specimen could not
be found in TNS. Peltigera dolichorrhiza is
a common terricolous species in the temperate region and might occur in Mikura-jimaIsland. However, it could not be found during the current investigations.
56). Peltigera pruinosa (Gyeln.) Inumaru
Specimens examined. F: YO&HM-7840(on humus with moss; 21/12/2010). J:
YO&HM-7806, 7811 (over humus on sloping road-bank), YO&HM-7812 (over soilon sloping road-bank).
57). Pertusaria multipuncta (Turner) Nyl.
Specimen examined. C: YO-8539 (onstone wall; 12/11/2011).
58). Phaeophyscia limbata (Poelt) Kashiw.

10

Yoshihito Ohmura et al.

This species is known as one of the commonest species of Phaeophyscia from lowland to montane regions in Japan, where it
grows on tree bark with mosses or on rockwith mosses (Kashiwadani, 1984). However, the taxonomic position of this taxon
has been controversial and different treatments have been applied e.g. P. hispidula
subsp. limbata by Poelt (1974) andEsslinger (1978), P. limbata by Kashiwadani (1984), Aptroot et al. (2002),
Harada et al. (2004) and Kurokawa andKashiwadani (2006), or as a synonym of P.
hispidula var. exornatula by Moberg (1995).
Although Kashiwadani (1984) excluded P.
hispidula from the Japanese lichen ﬂora,
several of his subsequent publications
reported P. hispidula, occasionally with P.
limbata, from Japan (Kashiwadani et al.,2000, 2002; Kurokawa and Kashiwadani,
2006; Shiba et al., 2008; Ohmura et al.,2012, 2014). Further taxonomic researchbased on molecular data in addition to the
known morphological and chemical features
are needed to solve the taxonomic problems
among P. hispidula and related species.
However, preliminary molecular data examined by Y. Ohmura seems to show that theJapanese populations of P. hispidula (s. lat.)
have low genetic variation and form a different clade from those collected in othercountries (unpublished data), which might
support the observation of Kashiwadani(1984). Hence, this study tentatively applies
the name of P. limbata to the species in thesense of Kashiwadani (1984).
Specimen examined. C: YO&HM-7858(on concrete; 22/12/2010).
59). Physciella melanchra (Hue) Essl.
Specimen examined. C: YO&HM-7857(on concrete; 22/12/2010).
60). Physma radians Vain.
This species was reported in Asahina(1955), and it was collected again during thecurrent investigations.
Physma radians is widely distributed

along the Paciﬁc side of Honshu, Shikoku,
Kyushu, and Ryukyu in Japan (Yoshimura,1974), Philippines (Vainio, 1921), Taiwan(Wang-Yang and Lai, 1973) and Thailand(Magain and Sérusiaux, 2014). However, it
seems to be relatively rare in Japan (Kashiwadani and Kurokawa, 1995). Mikura-jimaIsland is the northernmost locality for thisspecies.
Specimen examined. C: YO-8540 (overmosses on stone wall; 12/11/2011).
61). Porpidia albocaerulescens (Wulfen) Hertel& KnophSpecimen examined. C: YO-8551 (onstone wall; 12/11/2011).
62). Pyrenula fetivica (Kremp.) Müll.Arg.
Specimens examined. Without detailed
locality, ca. 150 m elev., MH-32 (on trunksof Shiia sieboldii; 13/7/1981). Along theroute between Ienosawa River and Inane
Shrine, 500 m elev., NN-8348 (on trunk ofCamellia japonica; 17/7/1981).
63). Pyrenula leucostoma Ach.
Specimen examined. Sato, ca. 150 m
elev., MH-96 (on bark of Camelia japonica;
13/7/1981).
64). Pyrenula platystoma (Müll.Arg.) Aptoot
It was reported as Anthracothecium oculatum Müll.Arg. in Kashiwadani and Kurokawa (1981).
Specimen examined. Without detailed
locality, YD-s.n. (on bark of Camellia
japonica; 29/9/1978).
65). Pyxine endochrysina Nyl.
Specimen examined. C: YO-8549 (onstone wall; 12/11/2011).
66). Ramalina litoralis Asahina
Specimens examined. A: YO-8536 (onrock). B: YO-7843 (on rock).
67). Rhabdodiscus crassus (Müll.Arg.) Frisch
Specimens examined. H: YO&HM7837B (on bark of Prunus sp.). N:
YO&HM-7785 (on bark of Camellia japonica).
68). Rhabdodiscus inalbescens (Nyl.) Frisch,comb. nov.

Lichens of Mikura-jima Island, Japan

Mycobank, no. 815287.
Basionym: Thelotrema inalbescens Nyl.,
Lich. Japon.: 57. 1890.
Type: Japan, Joumoto-Hakoné, corticola,
1879, E. Almquist s.n. (H-NYL-22453!,
holotype).
[Fig. 3]
Rhabdodiscus inalbescens is characterized by the pustulate thallus, the pustules
often bursting at the top to expose theﬁbrose to indistinctly granular-sorediate

11

medulla, or developing a granular-sorediate
surface; strongly emergent to adnate apothecia with reticulate columella; 1 seriate,brown, submuriform, 15–21×8–10 μm
large, 4–6×1–2 loculate ascospores withrounded to subacute ends, I＋ purplish-blue;
and psoromic acid.
This species has been frequently collected from Hokkaido to Kyushu up to
1,100 m elevation. It seems to be conﬁned toconiferous trees including Abies ﬁrma,

Fig. 3. Rhabdodiscus inalbescens (Nyl.) Frisch. A. Thallus (Y. Ohmura 8555, TNS). B. Granular-sorediate surface (Y. Ohmura 8555, TNS). C. Pustule bursting at the top with soredia (H. Kashiwadani 49141, TNS). Scale:
A＝1 cm, B & C＝0.5 mm.

12Yoshihito Ohmura et al.

Cryptomeria japonica (Thunb. ex L.f.) D.
Don, Picea glehnii, Pinus densiﬂora andTsuga sieboldii Carrière, and shows a preference for older trees with ﬁssured or ﬂaky
bark.
Although Mangold et al. (2009) considered Thelotrema inalbescens Nyl. as a synonym of R. ﬁssus Vain., the investigation of
the holotype and of numerous collections in
TNS conﬁrmed it as a separate species that,to present knowledge, is restricted to Japan.
From R. ﬁssus it is readily distinguishable
by the pustulate thallus which is most distinct in the numerous sterile specimens, andby its distinct ecology.
Specimen examined. C: YO-8555 (onbark of Cryptomeria japonica; 12/11/2011).
Additional specimens examined. JAPAN.
Hokkaido. Prov. Kushiro: along the trail
from Taro-ko to the top of Mt. O-akan,
Akan-cho, Akan-gun (43°26′N, 144°08′E),
on bark of Picea glehnii, 430‒700 m elev.,
5/9/1995, Y. Ohmura 1261 & H. Kashiwadani. Honshu. Prov. Musashi (Tokyo
Metropolis): Mt. Takao, Hachiohji-city, onbark of Cryptomeria japonica, 390 m elev.,
13/12/1993, H. Kashiwadani 44711. Prov.
Shinano (Pref. Nagano): Shimonakane,
Minamishinano-mura, Shimoina-gun, on barkof Abies ﬁrma, ca. 550 m elev., 5/10/2000,
H. Kashiwadani 42593; north western slopeof Oyama mountain, Chichibu-Tama-Kai N.
P.,
Kawakami-mura,
Minamisaku-gun(N35°56′41.0″, E138°43′21.9″), on large
Tsuga diversifolia, 1,658 m elev., 27/3/2013,
A. Frisch 13/Jp281 & Y. Ohmura. Prov.
Mikawa (Pref. Aichi): Mt. Iwagoya, Waichi,
Shitara-cho, Kitashitara-gun (35°01′N,
137°39′E), on bark of Pinus densiﬂora, 780 m
elev., 20/10/2003, H. Kashiwadani 46140.
Prov. Ohmi (Pref. Shiga): Samegai Gorge,
Maibara-city (N35°17′58.0″, E136°20′20.0″),
on Cryptomeria japonica, 180 m elev.,
13/7/2012, A. Frisch 12/Jp340, Y. Ohmura& G. Thor; ca. 2 km WNW of Mt. Amagoi,
along Shibukawa River, Eigenji-cho, Kan-zaki-gun, on bark of Pinus densiﬂora, ca.
400 m elev., 12/11/2009, H. Kashiwadani
50010. Prov. Kii (Pref. Wakayama): Nyu
Shrine, Higashi, Ryujin-mura, Tanabe-city,
on bark of Cryptomeria japonica, ca. 300 m
elev., 29/3/2011, H. Kashiwadani 49141.
Shikoku. Prov. Tosa (Pref. Kochi): None,
Toyo-cho, Aki-gun, on bark of Cryptomeria
japonica, ca. 5 m elev., 7/3/2011, H. Kashiwadani 49059, 49066. Kyushu. Prov. Buzen(Pref. Ohita): Mt. Hikosan, Soeda-cho,
Tagawa-gun, on bark of Cryptomeria japonica, 1,106 m elev., 26/10/1996, Y. Umezu 283.
Prov. Bungo (Pref. Ohita): en route from
Notohge Pass to Mt. Ichino-dake, Yamakuni-cho, Shimoge-gun, on bark of Tsuga
sieboldii, 720‒910 m elev., 10/10/1996, H.
Kashiwadani 39994 & Y. Umezu.
69). Sarcographa tricosa (Ach.) Müll.Arg.
Specimen examined. D: YO-8562 (onbark of Castanea crenata).
70). Stereocaulon exutum Nyl.
Specimen examined. M: YO&HM-7805(on rock).
71). Stereocaulon japonicum Th.Fr.
This species was reported in Asahina(1955) with the recognition of S. japonicum
var. aogasimense (Asahina) I.M.Lamb.
occurring in another island, AogashimaIsland. Stereocaulon japonicum is a common saxicolous species at lower to middle
elevations of Japan, and might occur in
Mikura-jima Island. However, it could not
be found in the ﬁeld investigations, and thevoucher specimen was also not found in
TNS.
72). Stereocaulon japonicum var. aogasimense(Asahina) I.M.Lamb.
Asahina (1969) reported the occurrenceof this variety in this island. This taxon ischaracterized by 1) the minute size of thephyllocladia, which are cylindric-coralloid,
not over 0.5 mm long and less than 0.1 mm
in thickness; 2) the persistent primary thallus; and 3) the presence of atranorin, norstictic and stictic acids. It could not beLichens of Mikura-jima Island, Japan

found during this investigation and novoucher specimen was found in TNS.
73). Stereocaulon sorediiferum Hue
This species was reported in Asahina(1955), and it was also collected by the ﬁrst
author during the current investigations andother collectors.
Specimens examined. K: YO&HM-7820(on rock). M: YO&HM-7804 (on rock).
Without detailed locality, JH-s.n. (17/7/
1966). Mt. Kurosaki-Takao-san, NS-s.n. (onrocks; 3/10/1963). Mt. Oyama (label as “Mt.
Misen”), NS-s.n. (on rocks; 4/10/1963).
74). Stereocaulon verruculigerum Hue
Specimens examined. F: YO&HM-7842(on rock; 21/12/2010), YO-8446 (on rock;
10/11/2011). Along the route between Satoand Ohshimawake River, ca. 180 m elev.,
MH-10181 (on rock at cliff; 13/7/1981).
75). Sticta gracilis (Müll.Arg.) Zahlbr.
This species was reported in Asahina(1955). Sticta gracilis is usually found in
humid forests of temperate to subtropicalregions, and might occur in Mikura-jimaIsland. However, it could not be found during the current investigations, and thevoucher specimen examined by Asahina(1955) was also not found in TNS.
76). Thecaria montagnei (Bosch) Staiger
In Japan, this species is rare but has been
widely collected from various localities: i.e.
Kohzu-shima Island that is one of Izu
Islands (Kashiwadani, 1979), and warmtemperate to subtropical regions in southern
Kyushu and Ryukyu Islands (Nakanishi,
1966, 1977). Additional examination of herbarium specimens in TNS revealed that thisspecies is also distributed in Bonin Islands
and Sulphur Islands in Japan.
Specimens examined. Along the route
between Sato and Ohshimawake River,
140 m elev., MH-10169 (on trunk of Camellia japonica; 13/7/1981); the same locality,
180 m elev., MH-10187 (on trunk; 13/7/
1981).
Additional specimens examined. JAPAN.

13

Bonin Islands: Mt. Mikazuki, Chichijimaisland, ca. 210 m elev., 10/6/1977, H. Kashiwadani 13861; en route from Kohshinzuka
to Kitamura, Hahajima-island, 30/11/1977,
N. Amano s.n. Sulphur Islands: Kita-iwo
Island (N25°26′20″, E141°17′02″), on barkof Eurya japonica 430 m elev., 18/6/2009, Y.
Ohmura 6432.
77). Thelotrema similans Nyl.
Specimen examined. I: YO&HM-7832
pr.p. (on decayed wood).
78). Trapelia coarctata (Turner ex Sm.) M.Choisy
Specimen examined. C: YO-8544 (onstone wall; 12/11/2011).
Phytogeographical notes
According to Kurokawa (2006), sixteen phytogeographical elements are recognized for thelichen ﬂora of Japan on basis of their world distribution patterns, namely 1) holarctic, 2) bipolar,
3) alpine, 4) boreal, 5) Beringian, 6) eastern
Asiatic-North American disjunctive, 7) northeast
Asiatic, 8) pantemperate, 9) Sino-Japanese, 10)
endemic, 11) pantropical, 12) paleotropical, 13)
Indo-Malayan, 14) Paciﬁc, 15) circum-Paciﬁc,and 16) Australian. Among these elements, ten
were recognized for the species occurring in
Mikura-jima Island.
1. Eastern Asiatic-North American disjunctive (1 species, 1.3%): Lecanora subimmergens(see Kurokawa, 2006).
2. Northeast Asiatic element (3 species,3.8%): Cladonia kurokawae, Peltigera dolichorrhiza (see Shiba et al., 2008); Nipponoparmelia
laevior (see Kurokawa, 2006).
3. Pantemperate element (25 species, 32.1%):
Cladonia furcata, C. scabriuscula, Porpidia
albocaerulescens (see Kurokawa, 2006); C.
merochlorophaea,
Leptogium
cyanescens,
Parmelinopsis minarum, P. spumosa, Trapelia
coarctata [they were classiﬁed as cosmopolitan in
Galloway (2007)]; C. coniocraea, C. macilenta,
C. ochrochlora, C. ramulosa, Coccocarpia
erythroxyli, Collema japonicum, C. subﬂaccidum,
Graphis scripta, Heterodermia diademata,14

Yoshihito Ohmura et al.

Leptogium azureum, Myelochroa entotheiochroa,
Normandina pulchella, Physciella melanchra(see Shiba et al., 2008); Lepraria ﬁnkii (seeLendemer, 2013). Although Agonimia paciﬁca
was suggested as Indo-Malayan (＝Western
Paciﬁc) element by Galloway (2007) or boreal
element by Shiba et al. (2008), it can be classiﬁed into the pantemperate element because of its
wide distribution in Asia, Oceania and Brazil(Harada, 1993; Aptroot et al., 1997, 2002; Moon,
1999; Aptroot, 2002, 2003; Galloway, 2007).
Lepraria lobiﬁcans might be classiﬁed into thiselement because it is reported from Japan, France
and Canary Islands (Crespo et al., 2006). Pertusaria multipuncta, which is commonly found in
the beech forests in Japan and the Northern
Hemisphere (Oshio, 1968; Urbanavichus, 2010),
can be classiﬁed into this element although Shibaet al. (2008) considered it as boreal element.
4. Sino-Japanese element (10 species,
12.8%): Stereocaulon sorediiferum (see Kurokawa, 2006); Phaeophyscia limbata (see Shiba etal., 2008). The following species can be classiﬁed into this element: Caloplaca spodoplaca,
which was only rarely collected in Japan (Nylander, 1890) and Korea (Kondratyuk et al.,2012); Diorygma soozanum, distributed in warm
temperate to subtropical regions of Japan, Taiwan, and China (Nakanishi, 1966; Meng andWei, 2008); Fissurina inabensis, which isreported from Japan, Taiwan and Thailand(Nakanishi, 1966; Nakanishi et al., 2001);
Hemithecium oshioi, which is reported fromJapan and China (Nakanishi, 1966; Jia et al.,2011); Ochrolechia parellula, which is reported
from Japan (Yoshimura, 1974) and Korea (Moon,
1999); Ramalina litoralis, which is reported fromJapan, Korea and Taiwan (Kashiwadani et al.,2006); Stereocaulon exutum, which is reported
from Japan (Yoshimura, 1974) and Korea (Moon,2013); Thelotrema similans, is distributed in temperate regions of Japan (Matsumoto, 2000) andJeju Island in Korea (Joshi et al., 2012).
5. Endemic to Japan (5 species, 6.4%):
Graphis mikuraensis, only reported from
Mikura-jima island (this study); Herpothallon

japonicum, reported from the nemoral to subtropical region (Aptroot et al., 2009); Lecanora
mikuraensis, reported from Izu Islands, Shikoku
and Kyushu (Miyawaki, 1988); Rhabdodiscus
inalbescens, distributed in Hokkaido to Kyushu(this study); Stereocaulon japonicum var. aogasimense, reported from Izu Islands, Ryukyu Islands
and Hirado Island (Asahina, 1969).
6. Pantropical element (11 species, 14.1%):
Enterographa leucolyta (see Sparrius, 2004);
Graphis cleistoblephara, G. tsunodae (seeLücking et al., 2009); Lecanora leprosa (seeRyan et al., 2004); Pyrenula fetivica, P. leucostoma, P. platystoma (see Aptroot, 2012). Cresponea proximata was suggested for this element by
Egea et al. (1996). Although Galloway (1996)
considered C. proximata as both Neotropical andIndo-Malayan elements, it is widely distributed
in warm temperate and tropical regions of Asia
and South America (Egea and Torrente, 1993;
Breuss and Brunnbauer, 1997; Messuti and Ferraro, 2002). Dibaeis sorediata can also be classiﬁed into this element because it occurs in warm
temperate to subtropical regions of Japan (Haradaet al., 2007), Nepal, Australia, and Réunion(Gierl and Kalb, 1993). Leptogium pichneum canbe classiﬁed into this element because it occurs
in warm temperate to subtropical regions ofJapan, Taiwan, Southeast Asia, Australia, andSouth America (Zahlbruckner, 1925; Yoshimura,1974). Parmotrema tinctorum belongs to this element with adjacent temperate regions (Nash andElix, 2002) although Shiba et al. (2008) considered it as the pantemperate element.
7. Paleotropical element (10 species,
12.8%): Cladonia fruticulosa (see Galloway,2007); Graphis assimilis, G. japonica, G. longiramea (see Lücking et al., 2009); Lobaria discolor (see Shiba et al., 2008); L. retigera, Pyxine
endochrysina (see Kurokawa, 2006); Rhabdodiscus crassus (see Frisch and Kalb, 2006; Frischand Ohmura, 2012). The following species can
be classiﬁed into this element: Graphis hossei,
which occurs in warm temperate regions ofJapan, China (Nakanishi, 1966, 1974), Indonesia,
Philippines, Thailand (Lücking et al., 2009), andLichens of Mikura-jima Island, Japan

Bolivia (Schiefelbein et al., 2014); Thecariamontagnei, which occurs in tropical to subtropical regions of East and South-East Asia, Oceania,
and West Africa (Staiger, 2002; Archer, 2006,2009).
8. Indo-Malayanelement(＝Western
Paciﬁc element) (6 species, 7.7%): Cladonia
submultiformis, Stereocaulon verruculigerum(see Kurokwa, 2006); Hypotrachyna exsecta (seeGalloway, 2007). The following species can beclassiﬁed into this element: Heterodermia pandurata, which occurs in warm temperate to subtropical regions of Japan, Taiwan, Thailand (Kurokawa, 1962), and India (Singh and Sinha, 2010);
Lepraria cupressicola, which is reported fromJapan, Australia, Norfolk Island, and Sri Lanka(Elix, 2009); Sarcographa tricosa, which isreported from southern Japan, Ceylon, Southeast
Asia, and Australia (Nakanishi, 1966; Archer,2009).
9. Paciﬁc element (6 species, 7.7%): Hypotrachyna koyaensis (see Kurokawa, 1996),
Physma radians (see Kurokwa, 2006). The following species can be classiﬁed into this element: Ocellularia microstoma, which is reported
from Japan, Australia and the Philippines(Mangold et al., 2009; Kraichak et al., 2014);
Peltigera pruinosa, which is reported fromJapan, China, Taiwan, Australia (Yoshimura,
1974; Louwhoff, 2009), and Korea (Moon,2013); Stereocaulon japonicum, which isreported from Japan, Taiwan, China, Java
(Yoshimura, 1974), and Korea (Moon, 2013);
Sticta gracilis, which is recorded from Japan,
China and New Caledonia (Yoshimura, 1974).
10. Circum-Paciﬁc element (1 species,1.3%): Coccotrema porinopsis (see Messuti,
1996; Kurokawa, 2006).
Judging from the ﬂoristic composition outlined above, the lichen ﬂora of Mikura-jimaIsland is characterized by the high percentage of
pantemperate species in combination with tropical to subtropical species. Despite of the previous
reports for vascular plants and bryophytes (Ohba,
1971; Higuchi and Nishimura, 2001), subalpine
species of lichens could not be found in the15

investigated area. It should be also noted that
Nipponoparmelia laevior, a conspicuous macrolichen which belongs to the northeast Asiatic element, could not be found during this study. The
voucher specimens of N. laevior collected fromvarious localities in this island before 1981 suggest that this species may have been common informer times but may have become extinct ordrastically reduced in this island during the past
30 years.
Acknowledgments
We wish to express our sincere gratitude to M.
Iwama, H. Miyakawa and Y. Miyakawa of Nonproﬁt Organization Nature center risen, S. Hirose
and M. Hino of Mikura-jima Island for their kind
help during the ﬁeld surveys. We are alsoindebted to the village ofﬁce for the coordinationof this investigation and the residents of Mikurajima Island for their warm encouragement
regarding our exploration. Thanks are extended
to H. Ebihara, Y. Mouri and K. Matsukura for
their technical assistances in the laboratory
experiments, M. Higuch of National Museum of
Nature and Science for providing locality information of his collections, and an anonymous
reviewer for the critical reading of this manuscript. This study is partly funded by the JSPS
KAKENHI no. 24300314 to Y. Ohmura and
JSPS Postdoctoral Fellowship for Foreign
Researchers (no. 2301706) to A. Frisch.
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