The Lichenologist 43(2): 141–146 (2011)
doi:10.1017/S0024282910000812
© British Lichen Society, 2011
Caloplaca aequata is a synonym of C. cinnabarina
(Teloschistaceae)
Yogesh JOSHI, Holger THÜS and Jae-Seoun HUR
Abstract: In 1915 Hue described Lecanora (sect. Placodium) aequata growing over quartzite rocks
from Busan and the Kangwon Provinces of South Korea. Later Zahlbruckner (1931) transferred it to
the genus Caloplaca. Careful examination of syntype material of L. aequata lodged at The Natural
History Museum, UK (BM) demonstrates that the species is conspecific with C. cinnabarina (Ach.)
Zahlbr. and we henceforth synonymize it with that species.
Key words: Asia, Hue, Lecanora, lichen-forming fungi, South Korea, Zahlbruckner
Introduction
At the beginning of the twentieth century, the
French lichenologist Auguste-Marie Hue
(1840–1917) described a saxicolous lichen
under the name Lecanora (sect. Placodium)
aequata, based on material collected in Kangouen-to, Corea (=Kangwon-do, South
Korea, where do means province) and Fusan
(=Busan) (Hue 1915). The taxon was
later transferred to the genus Caloplaca by
Zahlbruckner (1931).
Caloplaca aequata (Hue) Zahlbr., was later
included in the checklists of Japan, Korea
and Thailand by Harada et al. (2004), Hur
et al. (2005) and Wolseley et al. (2002), respectively. Unfortunately, the exact locality
from which the type collections were made is
unclear. Hue (1915) in his protologue cited
“In Corea quartzicolam legit R. P. Faurie in
Kang-oue-to, n. 4289 et 4319, ac in Fusan,
n. 4395, julio et octobri 1901”, while
Zahlbruckner (1931) cites “Corea, quartzicola”. In addition, there is no material to be
found under either Lecanora or Caloplaca
aequata in the major herbaria of the Asian
countries.
Y. Joshi and J.-S. Hur: Korean Lichen Research Institute, Sunchon National University, Sunchon 540-742,
Republic of Korea. Email: jshur1@sunchon.ac.kr
H. Thüs: Botany Department, The Natural History
Museum, Cromwell Road, London SW7 5BD, UK.
Judging from the species epithet, we first
suspected C. aequata to be an aquatic species,
growing either near fresh water or in maritime regions. However, after consulting the
original publication, we came to the conclusion that these collections had all the
characteristic features in common with C.
cinnabarina and to clarify this situation a
thorough taxonomic revision of the relevant
type material of C. aequata (Fig. 1C–E) was
necessary. After studying syntype material, it
became clear that C. aequata and C. cinnabarina are same taxon, and that C. aequata
should be synomymized under the earlier
name.
An updated list of various synonyms belonging to this species, provided by earlier
workers (Poelt & Hinteregger 1993;
Wetmore & Kärnefelt 1999), is also included
in the text.
Materials and Methods
We examined type specimens of C. aequata (deposited in
BM) and C. cinnabarina (deposited in H), as well as
additional material [C. almorensis (Räsänen) D.D.
Awasthi and C. cinnabarina] from the herbaria of LWGLWU, LWG-AWAS, Hector Osorio and KoLRI for
comparative purposes. Several standard Asian and
American flora and a few specific papers were used for
identification (e.g. Kärnefelt 1988; Poelt & Hinteregger
1993; Wetmore & Kärnefelt 1999).
Morphology and anatomy were examined by standard microscopic techniques. Thallus and apothecial
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THE LICHENOLOGIST
characters were studied on hand-cut sections mounted
in water. For characters such as size of thallus, apothecia
and thickness of the hymenium, hypothecium, exciple,
five measurements were recorded for each specimen; 10
measurements per specimen were recorded for ascospore dimensions. Ascospore measurements are given
as: (min.–) {x̄−SD}–{x̄+SD} (–max.), where min. and
max. are the extreme values, x̄ the arithmetic mean, and
SD the corresponding standard deviation. Chemistry
was analyzed using standardized TLC (Orange et al.
2010).
Results and Discussion
Caloplaca cinnabarina (Ach.) Zahlbr.
In Engler & Prantl., Nat. Pflanzenfam. 1(1): 228 (1908).
Basionym: Lecanora cinnabarina Ach., Lichenogr.
Univers.: 402 (1810); type: Lesser Antilles, St. Bartholomie
[St. Barthelemy], Forström (H-Ach. 1242!).
Synonyms nova: Lecanora (sect. Placodium) aequata
Hue, Annals. mycol. 13(2): 78 (1915); type: Corea
(=South Korea), Kan.ouen.to (=Kangwon-do), julio
1901, leg. Faurie, det. Hue 4319 (BM!, lectotype
selected here).—Caloplaca aequata (Hue) Zahlbr., Cat.
Lich. Univ. 7: 211 (1931).
Additional synonyms (after Poelt & Hinteregger
1993; Wetmore & Kärnefelt 1999): Parmelia cinnabarina
(Ach.) Spreng., Syst. Veget. 4(1): 301 (1827).—
Placodium cinnabarinum (Ach.) Nyl., in Mémoir. Soc.
Imp. Scienc. Natur. Cherbourg. 3: 178 (1856).—
Callopisma cinnabarinum (Ach.) Müll. Arg., Flora 64:
513 (1881).—Blastenia cinnabarina (Ach.) Lindau, Die
Flechten: 207 (1913).
Caloplaca holochracea (Nyl.) Zahlbr., Cat. Lich. Univ.
7: 144 (1930).—Lecanora holochracea Nyl. J. Linn. Soc.
London, Bot. 15: 171 (1876); type: India, Mt. Khasya,
J. D. Hooker (H-Nyl. 29527).
?Caloplaca intermedia (B. de Lesd.) Zahlbr., Cat. Lich.
Univ. 10: 624 (1940).
Callopisma almorense Räsänen, Suom. Elain-ja Kasvit.
Seuran Van. Tiedon. 6: 82 (1952); type: India, Kumaun,
Almora district (now Pithoragarh district), Berinag, near
Pinus forest, alt. 5000 ft., on stones, 26 September 1950,
D. D. Awasthi 804 (Isotype-LWG-AWAS!).—Caloplaca
almorensis (Räsänen) D. D. Awasthi, Beih. Nova
Hedwigia 17: 33 (1965).
(Fig. 1)
Thallus saxicolous, continuous, smooth,
cracked-areolate in centre, marginal areoles
elongated with a distinct and abrupt thallus
edge that does not taper off to a prothallus;
reddish orange to orange (Fig. 1B & F).
Cortex thin, paraplectenchymatous.
Apothecia immersed in central areoles to
finally raised, 1–3 per areole, 0·1–0·4 mm
Vol. 43
diam., disc little darker orange than the thallus. Epihymenium golden brown, 10–20 m
high. Hymenium hyaline, 40–55 m high.
Hypothecium hyaline, of indistinct cells, with
oil globules. Parathecium poorly developed.
Amphithecium with algae. Paraphyses simple
or sparsely branched, with no swollen tips
or 1–2 slightly swollen cells. Asci 8-spored,
ascospores
polarilocular,
(8–)8·3–10·7
(–11·2) × (3·9–)4–4·5(–5·6) m, isthmus
(1·8–)2–2·8(–4·2) m.
For further morphological and anatomical
descriptions see Wetmore & Kärnefelt
(1999) who placed this species within its own
group.
Chemistry. Spot tests: thallus K+ red, C−,
P−. TLC: parietin (major), emodin, fallacinal. Wetmore & Kärnefelt (1999) mentioned in their description of that species a
C+ red reaction of the thallus. The species is
never C+ red. It was a typographic error as
suggested by Dr. Wetmore (personal communication) and should be corrected to
C−. Parietin along with traces of emodin
and fallacinal are reported in the specimens
examined.
For additional secondary metabolites reported in this species, see Santesson (1970)
who found teloschistin, xanthorin and parietinic acid as compounds as well as those
reported in our study.
Ecology and distribution. It is a common
rock dwelling lichen generally found growing
over siliceous rocks (granite, quartzite), but
sometimes on calcareous rocks in rather
exposed situations, throughout tropical and
subtemperate regions of the world. The
taxon has previously been reported from
America (Wetmore & Kärnefelt 1999), Asia
(Poelt & Hinteregger 1993; Aptroot &
Seaward 1999; Aptroot & Sipman 2001;
Joshi 2008; Joshi et al. 2010), West and
South Africa (Kärnefelt 1988), New Zealand
(Galloway 2007), Australia (Wetmore &
Kärnefelt 1999) and Oceania (Elix &
McCarthy 1998).
One of the authors (YJ) noticed that the
species has a wide distribution in India, extending from arid tropical regions to cold
2011
Caloplaca aequata, a synonym of C. cinnabarina—Joshi et al.
143
F. 1. A, lectotype collection of Caloplaca cinnabarina (Ach.) Zahlbr.(H, Ach = 1242); B. magnified image of
Lecanora cinnabarina; C, syntype collection of Lecanora aequata Hue (Abbé Faurie: Lichens de Coreé, no. 4319 =
BM000733815) [ = Caloplaca cinnabarina (Ach.) Zahlbr.]; D, specimens of Caloplaca aequata (Hue) Ach. (anon.,
Kaw Tao, L41 = BM000733814); E, magnified image of Lecanora aequata; F, isotype collection of Callopisma
almorense Räsänen lodged at LWG-AWAS showing subeffigurate marginal areoles and immersed apothecia. Scales:
B = 1 mm; E = 2 mm; F = 3 mm.
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THE LICHENOLOGIST
subalpine regions. In temperate to subalpine
regions, the species is found between 1440
and 2700 metres growing with Caloplaca
flavovirescens (Wulfen) Dalla Torre and
Sarnth. and Ioplaca pindarensis (Räsänen)
Poelt & Hinter., while in tropical regions
(500–1170 m), it is found growing with C.
awasthii Y. Joshi & Upreti, C. poliotera (Nyl.)
J. Steiner, C. subpoliotera Y. Joshi & Upreti,
Endocarpon spp. and Peltula placodizans
(Zahlbr.) Wetmore (Joshi 2008). However,
South Korean specimens show a greater
affinity to maritime regions, having a tropical
distribution (10–260 m), and are found
growing on cliffs and boulders along with C.
bogilana Y. Joshi & J. S. Hur and C. decipiens
(Arnold) Blomb. & Forssell (Joshi et al.
2010).
Nomenclatural notes. Two well preserved
and fertile samples of C. aequata lodged at
BM, 4319 (BM000733815) and L41
(BM000733814) were investigated. Sample
4319 contains one piece of rock glued onto a
card, which itself is mounted on a big sheet
(Fig. 1C). The text on the card reads
“Lecanora aequata Hue, Kan-ouen-to, julio
1901, leg. Faurie, det. Hue, Abbé Faurie
Lichens de Coreé, no. 4319 (syntype)”, in
black ink, except the word “syntype”, which
is written in red ink. It corresponds with the
locality and collection in the protologue, and
consists of a large piece of lichen with an
orange-red thallus having immersed to sessile dark red apothecia. On the other hand,
the sample L41 contains three pieces of rock
glued onto a card (Fig. 1D). There was no
text on the card, but the annotation slip and
envelope bears the information “Caloplaca
aequata Ach., Kaw Tao – 1929, det. R.
Paulson, no. L41, Herb. Hort. Bot. Reg.
Kew” in black ink. Both samples of C.
aequata at BM, including the syntype are in
good condition, and their macroscopic and
microscopic characters fit well with those of
C. cinnabarina. In the syntype, the marginal
areoles are 0·5–0·8 mm long and 0·3–0·4
wide. The thallus is 72–95 m thick, without
a necral layer, the cortex is 14–20 m thick,
the algal layer continuous and the medulla
dense and without crystals. The apothecia
Vol. 43
are immersed to raised, 1–4 per areole, round
to irregular and 0·1–0·35 mm diam. The
golden brown epihymenium is 10·5–12·5 m
high and the hyaline hymenium, 37·5–50 m
high. The hyaline hypothecium is made
up of indistinct cells without oil droplets.
The parathecium is poorly developed and the
amphithecium with alga extending below
the hypothecium. The paraphyses are simple
to ±furcate at the tips, with or without 1–2
swollen cells at the tips. Ascospores are
(8–)10–11 × 4–4·5 m and the isthmus 2·3–
2·8 m. Hence all the characters fall within
the range of C. cinnabarina as outlined above
and described in detail by Wetmore &
Kärnefelt (1999).
The date of the combination of C. aequata
is given as 1931 (Zahlbruckner 1931), while
that of C. cinnabarina is 1908 (Zahlbruckner
1931), which means that C. cinnabarina has
nomenclatural precedence over C. aequata.
Henceforth, C. aequata is herewith synonymized with C. cinnabarina. In addition, C.
aequata should be cited as C. aequata (Hue)
Zahlbr., not C. aequata Ach. (as mentioned
by R. Paulson on the packet L41).
Despite the fact that Hue did not give an
exact locality (type locality), select a holotype
specimen or mention any herbarium, the
names [L. (sect. Placodium) aequata and C.
aequata] are valid. The publication antedates
the effective dates of the respective articles in
the International Code of Botanical Nomenclature [ICBN Art. 37.1 (type specimen): 1
January 1958; ICBN Art. 37.6 (inclusion of
“typus” or “holotypus”): 1 January 1990;
ICBN Art. 37.7 (herbarium name): 1 January
1990] (McNeill et al. 2006).
Hue (1915) in his original description of
Lecanora (sect. Placodium) aequata, mentioned three specimens from Korea: two
from Kang-ouen-to (4289 and 4319) and
one from Fusan (4395). We studied the
specimen 4319 along with the specimen
of Caloplaca aequata (L41) determined by
Paulson. Both of them were identical to
C. cinnabarina. As sample 4319 is in good
condition and bears numerous apothecia and
well developed thallus, we have selected it
here as the lectotype according to Art. 9.2 of
ICBN Vienna Code (McNeill et al. 2006).
2011
Caloplaca aequata, a synonym of C. cinnabarina—Joshi et al.
Remarks. Caloplaca cinnabarina is macroscopically a distinctive species because of its
areolate to subeffigurate, cinnabarine coloured thallus with small, immersed to ± sessile apothecia mostly obscuring the thallus.
The only three species in the Asian mycobiota with which it might be confused are
Ionaspis lacustris (With.) Lutzoni, Ioplaca
pindarensis and Caloplaca rubelliana (Ach.)
Lojka. Ionapsis lacustris differs in lacking
anthraquinones and having halonate, simple
spores. Ioplaca pindarensis is similar in external morphology, but differs in having a dark
under surface (black hypothallus), egg-yolk
coloured thallus and is endemic to the
Himalayan region. The widespread Caloplaca rubelliana, another member of the
Cinnabarina group differs in having thallus
edges tapering off gradually either to nothing
or to a pale prothallus, while thallus edges in
C. cinnabarina have distinct abrupt edges
without thinning or prothalli. Besides that,
C. rubelliana has a scarlet red disc and a
greyish to brown-orange thallus. Caloplaca
subunicolor (Nyl.) Zahlbr., a species with
scattered records across the Southern
Hemisphere, has gradually tapering thallus
margins without prothallus and apothecia
soon becoming sessile, while in C. cinnabarina the thallus edge is always abrupt and the
apothecia remain more or less immersed or
finally become raised.
Specimens examined as C. aequata. Korea: Kaw Tao:
on rock, 1929, det. R. Paulson 41 (BM). Kan-ouen-to: on
rock, 1901, coll. A. Faurie, det. Hue 4319 (BM).
Specimens examined as C. cinnabarina. India:
Himachal Pradesh: alt. 2700 m, 1949, Harish C.
Raghubir 430 (LWG-AWAS); alt. 1440 m, 2002, S.
Nayaka & R. Srivastava 02-001208 (LWG). Karnataka:
alt. c. 980 m, 1979, D. D. Awasthi, D. K. Upreti & U. C.
Misra 79-212, 79-213, 79-231, 79-242, 79-261, 79-269
(LWG-LWU); alt. 900 m, 1997, S. Nayaka 413
(LWG); 1956, D. D. Awasthi 3595 (LWG-AWAS).
Madhya Pradesh: 18 iv 1973, P. S. H. Khan & party s. n.
(LWG); 12 vii 2004, D. K. Upreti s. n. (LWG); alt.
500 m, 2004, Y. Joshi 04-004542 (LWG). Orissa: 17 ix
1993, Vivek Pande s. n. (LWG). Rajasthan: alt. 1680 m,
1973, A. Singh 101472 (LWG); alt. 1170 m, 1973,
A. Singh 101421 (LWG). Tamil Nadu: alt. 950 m, 1996,
G. N. Hariharan CS4 (MSSRF); alt. 1000 m, 1999, S.
Nayaka 99-75925 (LWG); alt. 700 m, 1999, S. Nayaka
99-75952 (LWG); alt. 1490 m, 1999, S. Nayaka 307/B
(LWG); alt. 2100 m, 1959, D. D. Awasthi 4544 (LWGAWAS); alt. c. 2100 m, 1971, K. P. Singh 71.776
(LWG-LWU); alt. c. 2250 m, 1971, D. D. Awasthi & K.
145
P. Singh 71.270 (LWG-LWU); alt. c. 2400 m, 1971, K.
P. Singh 71.721 (LWG-LWU); alt. 1400–1500 m,
1990, D. K. Upreti & Hariharan 202170 (LWG).
Uttarakhand: alt. 1800 m, 1956, D. D. Awasthi 3464
(LWG-AWAS); alt. 1700 m, 2005, D. K. Upreti 05005058 (LWG); alt. 2400 m, 1950, D. D. Awasthi & A.
M. Awasthi 620 (LWG-AWAS); alt. 1760 m, 2007, Y.
Joshi & S. Joshi 07-010304, 07-010305, 07-010084,
07-010085 (LWG); alt. 1350 m, 1967, A. Singh & party
91539 (LWG); alt. 1950 m, 1954, D. D. Awasthi 2673
(LWG-AWAS); alt. 1500 m, 1999, H. C. Pande 9965596 (LWG); alt. 2300 m, 2000, V. Pant 20-67548
(LWG); alt. 1600 m, 1989, D. K. Upreti 201883
(LWG); alt. 1500 m, 1950, D. D. Awasthi 803 (LWGAWAS); 2001, V. Pant 02-223452 (LWG); 1951, D. D.
Awasthi 855 (LWG).—South Korea: Gyeongnam
Prov.: 34°46#08·3$N, 127°59#24$E, alt. 171 m, 2004,
Jae-Seoun Hur 040022 (KoLRI). Jeonbuk Prov.:
35°37#02·7$N, 126°35#01·7$E, alt. 260 m, 2004,
Jae-Seoun Hur 040167 (KoLRI). Jeonnam Prov.:
34°25#20·8$N, 127°08#43·1$E, alt. 10 m, 2005,
Jae-Seoun Hur 050208/1 (KoLRI); 34°09#64$N,
126°35#11·5$E, alt. 15 m, 2010, Y. Joshi, H. S. Jeon &
M. H. Jeong 100166, 100194 (KoLRI); 34°10#90$N,
126°31#83·6$E, alt. 11 m, 2010, Y. Joshi, H. S. Jeon &
M. H. Jeong 100256, 100257 (KoLRI); 34°38#23·9$N,
127°25#19·5$E, alt. 85 m, 2010, Y. Joshi, H. S. Jeon &
G. S. Han 100270 (KoLRI).—Uruguay: MaldonadoCerro Pan de Azucar, 100 m.s.m., 1951, H. S. Osorio
4652 (hb. Hector Osorio).—W Nepal: Seti zone, alt.
1500 m, 1977, L. R. Sharma 77.4 (LWG-LWU);
Karnali-zone, alt. 2250 m, 1977, L. R. Sharma
77.207A, 77.207B (LWG-LWU); alt. 2250 m, 1977,
L. R. Sharma 77.202 (LWG-LWU).
We are grateful to the directors of the herbaria cited
for placing material at our disposal, Drs S. Kondratyuk
and L. Lőkös for critically reviewing the manuscript
and providing Hue’s reprints, and to Sunchon National
University for providing the laboratory facilities. This
work was supported by grants from the Korea National
Research Resource Center Program (2010-0000660)
through the National Research Foundation of Korea
(NRF) and the Korean Forest Service Program (KNA
2010) through Korea National Arboretum. The first
author also thanks Dr Leena Myllys for providing the
type specimen of Lecanora cinnabarina Ach. on a loan
basis and to Mr Xin Yu Wang, Miss Thi Thuy Nguyen,
Jung Ae Ryu and Hae Sook Jeon for their help and
kindness.
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