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 Kang-ouen-to, Corea (=Kangwon-do, South Korea, where do means province) and Fusan (=Busan) (Hue Reference Hue1915). The taxon was later transferred to the genus Caloplaca by Zahlbruckner (Reference Zahlbruckner1931).
Caloplaca aequata (Hue) Zahlbr., was later included in the checklists of Japan, Korea and Thailand by Harada et al. (Reference Harada, Okamoto and Yoshimura2004), Hur et al. (Reference Hur, Koh and Harada2005) and Wolseley et al. (Reference Wolseley, Aguirre-Hudson and McCarthy2002), respectively. Unfortunately, the exact locality from which the type collections were made is unclear. Hue (Reference Hue1915) 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 (Reference Zahlbruckner1931) 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.
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 Reference Poelt and Hinteregger1993; Wetmore & Kärnefelt Reference Wetmore and Kärnefelt1999), 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 LWG-LWU, 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 Reference Kärnefelt1988; Poelt & Hinteregger Reference Poelt and Hinteregger1993; Wetmore & Kärnefelt Reference Wetmore and Kärnefelt1999).
Morphology and anatomy were examined by standard microscopic techniques. Thallus and apothecial
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.–) {
−SD}–{
+SD} (–max.), where min. and max. are the extreme values,
the arithmetic mean, and SD the corresponding standard deviation. Chemistry was analyzed using standardized TLC (Orange et al. Reference Orange, James and White2010).
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 Reference Poelt and Hinteregger1993; Wetmore & Kärnefelt Reference Wetmore and Kärnefelt1999): 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)
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.
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 diam., disc little darker orange than the thallus. Epihymenium golden brown, 10–20 µm high. Hymenium hyaline, 40–55 µm 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) µm, isthmus (1·8–)2–2·8(–4·2) µm.
For further morphological and anatomical descriptions see Wetmore & Kärnefelt (Reference Wetmore and Kärnefelt1999) who placed this species within its own group.
Chemistry. Spot tests: thallus K+ red, C−, P−. TLC: parietin (major), emodin, fallacinal. Wetmore & Kärnefelt (Reference Wetmore and Kärnefelt1999) 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 (Reference Santesson1970) 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 Reference Wetmore and Kärnefelt1999), Asia (Poelt & Hinteregger Reference Poelt and Hinteregger1993; Aptroot & Seaward Reference Aptroot and Seaward1999; Aptroot & Sipman Reference Aptroot and Sipman2001; Joshi Reference Joshi2008; Joshi et al. Reference Joshi, Wang, Yamamoto, Koh and Hur2010), West and South Africa (Kärnefelt Reference Kärnefelt1988), New Zealand (Galloway Reference Galloway2007), Australia (Wetmore & Kärnefelt Reference Wetmore and Kärnefelt1999) and Oceania (Elix & McCarthy Reference Elix and McCarthy1998).
One of the authors (YJ) noticed that the species has a wide distribution in India, extending from arid tropical regions to cold 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 Reference Joshi2008). 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. Reference Joshi, Wang, Yamamoto, Koh and Hur2010).
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 µm thick, without a necral layer, the cortex is 14–20 µm thick, the algal layer continuous and the medulla dense and without crystals. The apothecia 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 µm high and the hyaline hymenium, 37·5–50 µm 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 µm and the isthmus 2·3–2·8 µm. Hence all the characters fall within the range of C. cinnabarina as outlined above and described in detail by Wetmore & Kärnefelt (Reference Wetmore and Kärnefelt1999).
The date of the combination of C. aequata is given as 1931 (Zahlbruckner Reference Zahlbruckner1931), while that of C. cinnabarina is 1908 (Zahlbruckner Reference Zahlbruckner1931), 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. Reference McNeill, Barrie, Burdet, Demoulin, Hawksworth, Marhold, Nicolson, Prado, Silva and Skog2006).
Hue (Reference Hue1915) 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. Reference McNeill, Barrie, Burdet, Demoulin, Hawksworth, Marhold, Nicolson, Prado, Silva and Skog2006).
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 (LWG-AWAS); alt. c. 2100 m, 1971, K. P. Singh 71.776 (LWG-LWU); alt. c. 2250 m, 1971, D. D. Awasthi & K. 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 05-005058 (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 99-65596 (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 (LWG-AWAS); 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: Maldonado-Cerro 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.
