The Lichenologist 41(3): 249–255 (2009) © 2009 British Lichen Society
doi:10.1017/S0024282909008214 Printed in the United Kingdom
Caloplaca himalayana, a new epiphytic lichen from the
Indian subcontinent
Yogesh JOSHI, Dalip K. UPRETI and Suresh C. SATI
Abstract: Twenty-three species of epiphytic Caloplaca are recognized in India, of which Caloplaca
himalayana Joshi, Y. & Upreti, characterized by thin, continuous, yellowish thallus, numerous
ferruginous-red to brownish red, flat to subconvex apothecia with a persistent proper margin and
lignicolous habitat, is described here as a new species. Because of the ferruginous coloured apothecial
disc, the new species is provisionally placed in the ferruginea group until a taxonomic treatment of the
whole group has been carried out. The ferruginea group is characterized by a whitish to grey (rarely
yellowish) crustose thallus, a ferruginous to rarely blackish apothecial disc, biatorine to lecanorine
exciple, thick-walled spore septa and the chemosyndrome C. The taxonomic affinities of the new taxa
with several other taxa, including species of ferruginea group, are discussed. Two new records for India
are also recorded: Caloplaca alnetorum Giralt, Nimis & Poelt and C. crocea (Kremp.) Hafellner & Poelt.
A key to all the corticolous Caloplaca species occurring in the Indian subcontinent is also provided.
Key words: Calplaca ferruginea group, India, Teloschistaceae, new species, new records
Introduction
Corticolous and/or lignicolous species of the
lichen genus Caloplaca have been taxonomically well-treated by various workers across
the world (e.g. Arup 2006; Giralt et al. 1992;
Hafellner & Poelt 1979; Hansen et al. 1987;
Khodosovtsev et al. 2004; Laundon 1992;
Magnusson 1944; Poelt & Hinteregger 1993;
Søchting 1989; Wade 1965; Wetmore 1994,
2001, 2004 a, b, 2007). For the Indian
subcontinent, Awasthi (1991), Poelt &
Hinteregger (1993) and Joshi & Upreti
(2007, 2008 a, b) reported 11, 24 and 11
corticolous species of Caloplaca, respectively.
While revising Teloschistacean taxa from
India, one of the authors (YJ) reported 23
corticolous species of this genus from India
(Joshi 2008c), one of which was collected 3
to 6 years ago in temperate regions of the
Western Himalayas growing over dead
wood. It is described here as new to science
Y. Joshi and S. C. Sati: Department of Botany, Kumaun
University, D.S.B. Campus, Nainital-263002, India.
Email: yogesh36953@rediffmail.com
D. K. Upreti: Lichenology Laboratory, National
Botanical Research Institute (C.S.I.R.), Rana Pratap
Marg, P. B. No. 436, Lucknow-226001, India.
along with two new records to the Indian
lichen flora. A revised and updated key of all
the corticolous Indian epiphytic Caloplaca
species is also provided.
Materials and Methods
The present study is based on collections in LWG (including LWU–AWAS). The morphological characters
were examined on dry material under a dissecting
microscope (×40). The thallus and ascomata were examined with a compound microscope (×100, oil immersion). The sections cut for studying anatomical details
were mounted in water. For characters such as size of
thallus, apothecium and thickness of the hymenium, 5
measurements were recorded from each specimen; 10
measurements per specimen were recorded for ascospore dimensions. The dimensions of epihymenium,
hymenium and ascospores are generally presented as
(minimum value recorded–) lowest specimen arithmetic
mean observed—highest specimen arithmetic mean observed (–maximum value recorded). All measurements
were made on material mounted in water, but the paraphyses were studied after replacing water with 25%
KOH (Wetmore 1994). Chemicals used in identification were 10% KOH (K), calcium hypochlorite (C),
and para-phenylenediamine (PD). Secondary metabolites were identified by TLC as described by Walker &
James (1980). The chromatograms were developed in
solvent systems A (toluene: 1, 4-dioxane: acetic acid)
and C (toluene: acetic acid). Terminology for tissues
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THE LICHENOLOGIST
Vol. 41
F. 1. Caloplaca himalayana, habitus (holotype in LWG). Scale = 3 mm.
generally follows that of Nash & Gries (2002) and
Wetmore (1994).
The New Species
Caloplaca himalayana Y. Joshi &
Upreti sp. nov.
Caloplaca flavorubescens similis sed margine thallino deficienti, amphithecio sine algis, sporis parvis differt.
Typus: India, Himachal Pradesh, Shimla district,
Rohru, Sungri, alt. 2600 m, on dead wood, 22 May
2002, S. Nayaka & R. Srivastava 02-219451 (LWG—
holotypus; CAL—isotypus).
(Fig. 1)
Thallus crustose, lignicolous, thin, continuous, determinate, irregular, 0·4–2(–3)
cm diam., 25–30 µm thick, yellowish. Cortex
thin, 8–13 µm thick, paraplectenchymatous,
of thin- walled cells, algal layer even and
continuous. Medulla macroscopically white,
of loose hyaline hyphae, prosoplectenchymatous. Prothallus absent.
Apothecia biatorine, numerous, scattered
to ± aggregated (1–4), sessile, round to ±
angular owing to pressure, 0·2–1(–1·2) mm
diam.; disc ferruginous red to reddish brown,
plane to subconvex, glossy; proper margin
smooth, entire, persistent, thin, 0·2–0·5 mm,
flush, concolorous or paler than disc, glossy;
thalline margin absent. Epihymenium golden
brown, 15–30 µm thick; hymenium hyaline,
50–80 µm high; hypothecium hyaline, of isodiametric cells, with numerous oil-droplets;
parathecium cupular, of thin walled elongated
hyphal cells; amphithecium of thin-walled
elongated hyphal cells, without algae or algae
± restricted to the base of parathecium.
Paraphyses thin, simple, ± furcated at the
tips, sometimes with ± inflated apices. Ascus
8-spored, ascospores polaribilocular, ellipsoid
to broadly ellipsoid to ovoid, 9–11 × (4–)5–
6(–7) µm, isthmus 2–4 µm.
Pycnidia not seen.
Chemistry. Thallus, apothecial disc and
epihymenium K+ purple, C−, Pd−. Medulla
K−, C−, Pd−. TLC: 7-chloroemodin
(major) and parietinic acid (traces).
Ecology and distribution. Currently C. himalayana is known from two localities in
Himachal Pradesh and one locality in Jammu
2009
Caloplaca himalayana—Joshi et al.
& Kashmir. The species was found growing
over dead wood between elevations of 2150–
2800 m in temperate regions of the Western
Himalayas. The three known communities in
which the new taxon has been encountered,
have a similar lichen composition (Lecanora
and Rinodina spp.), along with a dense mat of
cyanophycean species.
Remarks. The new taxon can be recognized
by a thin, continuous yellowish thallus with
ferruginous apothecia, absence of both a
thalline margin and algae in the amphithecium, and lignicolous habitat. It is often
confused with the citrine or orange-grey coloured species of C. herbidella (= C. herbidella
f. citrinescens H. Magn.) which differs from
the new taxa in being isidiate. Sometimes
C. flavorubescens (Huds.) J. R. Laundon with
deep reddish brown to brown apothecia is
difficult to separate from the new taxon, but
the former differs in the presence of a thalline
margin, algae in the amphithecium and larger
spores [12–20(–25) × 5–10(–13·6) µm].
Caloplaca ammiospila (Wahlenb.) H. Olivier,
C. caesiorufella (Nyl.) Zahlbr., C. ferruginea
(Huds.) Th. Fr., C. ferrugineofusca (Vain.) H.
Magn., C. jenisejensis H. Magn., C. spitsbergensis H. Magn. and C. subathallina H.
Magn., are other lignicolous or corticolous
species having ferruginous or reddish brown
coloured apothecial discs, but differ from the
new taxon in having endoxylic, poorly to
well-developed grey thalli and larger spores
(12–17µm). On the other hand, C. asserigena
(J. Lahm) Della Torre & Sarnth., having a
ferruginous coloured apothecial disc and
similar sized spores to that of the new taxon
(9–12 × 5–6·5 µm), differs in its grey thallus
and lecanorine to zeorine small apothecia
(0·2–0·4 mm broad). Caloplaca juniperi Poelt
& Hinter., another related species having a
yellowish thallus and ferruginous coloured
apothecial disc, differs from the new taxon in
having squamulose thalli and pale yellow
margined concave apothecia.
Additional specimens examined. India: Himachal
Pradesh: Shimla district, Narkanda, 3–4 km towards
Hatu Peak, alt. 2800 m, on dead wood, 2002, S. Nayaka
& R. Srivastava 02-81522 (LWG). Jammu & Kashmir:
Udhampur district, Patnitop, Tourist area, alt. 2150 m,
on dead wood, 9 xii 2005, M. Sheikh s.n. (LWG).
251
New Records
Caloplaca alnetorum Giralt, Nimis &
Poelt
Cryptog. Bryol. Lichénol. 13: 269 (1992).
This species is characterized by: irregular
to ± orbicular, very thin (almost confused
with the substratum) continuous to ± areolate, golden yellow thallus; orange apothecial
discs; persistent proper margin; parathecium
of ± elongated cells or paraplectenchymatous; amphithecium with algae; and spores
with thickened end walls, 11–12 × 5–7 µm,
with isthmus 4–6 µm. In external appearance
the species is close to Caloplaca cerinelloides
(Erichs.) Poelt, but differs in having a yellowish thallus and spores of variable shapes with
thickened end-walls. Caloplaca cerinelloides
has a greyish K− thallus. Caloplaca flavorubescens another related species, differs in having
a greenish yellow thallus with a prominent
thalline margin and larger broadly ellipsoid
spores [12–20(–25) × 5–10(–13·6) µm].
Poelt (1969, as “C. alnetorum ad int.”)
and Giralt, Nimis & Poelt (1992) reported
this species from Europe and Nordic
countries. Alon & Galun (1971) reported this
species growing over bark of Quercus calliprinos and Pistacia palestina from Israel. It is a
new record for India and is found growing on
bark of Cedrus and Abies between elevations
of 2000–2800 m in the Central Himalayas.
Additional specimens examined. India: Himachal
Pradesh: Chamba district, Chamba, around Joth, alt.
2000–2350 m, on Cedrus bark, 2001, D. K. Upreti & S.
Nayaka 01-75556 (LWG); Shimla district, Narkanda,
3–4 km towards Hatu Peak, alt. 2800 m, on Abies bark,
2002, S. Nayaka & R. Srivastava 02-67177 (LWG).
Caloplaca crocea (Kremp.) Hafellner &
Poelt
J. Hattori Bot. Lab. 46: 19–24 (1979). – Lecidea crocea
Kremp., Flora 59: 317 (1876).
The species is characterized by: a thin continuous to subareolate, smooth, whitishgrey, UV+ cream-yellow, K− thallus; yellow
to yellowish orange apothecial disc; proper
margin pale orange-brown; thalline margin
persistent, smooth, thick, raised above the
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THE LICHENOLOGIST
level of disc, concolorous with thallus; hypothecium paraplectenchymatous, with oil
droplets; parathecium of radiating elongated
hyphal cells, amphithecium with algae, outer
surface with hyaline crystals, paraphyses simple to ± branched at the tips, spores trilocular, 16–18 × 8–10 µm, middle locule of
similar size to the polar locules. In external
morphology, the species is similar to C. aff.
cerina (Ehrh. ex Hedw.) Th. Fr., a corticolous species, growing in temperate regions
of India. However, the trilocular spores of C.
crocea separate it from species of C. cerina
group.
Vol. 41
Hafellner & Poelt (1979) reported C.
crocea growing on bark and wood in Africa,
Madagascar, Argentina, Brazil, Paraguay
and Uruguay, while Wetmore (2007) reported it from North and Central America. It
is a new record for India and is reported from
the tropical regions of southern India, where
it was found growing on trees at an altitude of
1100 m with C. bassiae (Willd. ex Ach.)
Zahlbr.
Specimen examined. India: Tamil Nadu: Chennai,
Pulmedu, Siruvani Hills, alt. 1100 m, on bark, 1996,
G. N. Hariharan & Balaji CC9 (MSSRF).
Key to the epiphytic species of Caloplaca from the Indian subcontinent
A total of 32 species of epiphytic Caloplaca from the Indian subcontinent are keyed out, of
which 23 occur in India (bold in key) and the remaining 9 are reported from other parts of
the Indian subcontinent. Figures for spore measurements include (minimum value
recorded–) lowest specimen arithmetic mean observed—highest specimen arithmetic mean
observed (–maximum value recorded); dimensions given for apothecia all refer to their
diameter. Vegetative propagules play an important role in demarcating the species, hence
their presence or absence is an important distinguishing character. The number of locules
within the spores varies between the species, but one type, polaribilocular is more common
than than any other; the type of spore is only given when deviating from the normal type or
where it is important for identification. The key in this paper will certainly not solve all
problems, but should be consulted together with previous publications (cited above).
1
Thallus sorediate, isidiate or blastidiate . . . . . . . . . . . . . . . . . . . . . . 2
Thallus without soredia, isidia or blastidia . . . . . . . . . . . . . . . . . . . . 12
2(1)
Thallus isidiate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Thallus sorediate or blastidiate . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3(2)
Thallus grey; apothecial disc ferruginous red . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . C. herbidella (Nyl. ex Hue) H. Magn.
Thallus yellow; apothecial disc orange-brown to brownish red . . . . . . . . . . 4
4(3)
Apothecia biatorine; cosmopolitan . . . . C. bassiae (Willd. ex Ach.) Zahlbr.
Apothecia lecanorine; restricted to temperate regions . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . C. kashmirensis Joshi, Y. & Upreti
5(2)
Thallus blastidiate . . . . . . . . . . . . . . . . . . . . C. farinosa Poelt & Hinter.
Thallus sorediate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6(5)
Thallus leprose, completely sorediate or nearly so . . . . . . . . . . . . . . . . . 7
Thallus only partly dissolved into soredia . . . . . . . . . . . . . . . . . . . . . . 9
7(6)
Thallus completely leprose, golden to brownish yellow, consorediate . . . . . . . .
. . . . . . . . . . . . . . . . . . C. chrysodeta (Vain. ex Räsänen) Dombr.
Thallus usually completely dissolved into soredia but not leprose . . . . . . . . . 8
8(7)
Soredia arising marginally . . . . . . . . . . . . . . . C. phlogina (Ach.) Flagey
Soredia of medullary origin . . . . . . . . . . . C. phloginopsis Poelt & Hinter.
2009
Caloplaca himalayana—Joshi et al.
253
9(6)
Thallus grey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Thallus yellow-orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10(9)
Thallus distinctly grey, rarely sorediate, with numerous brown to blackish apothecia;
tropical distribution . . . . . . . . C. atrosanguinea (G. Merr.) I. M. Lamb
Thallus grey, sometimes yellow tinged, distinctly sorediate, rarely apotheciate;
temperate distribution . . . . . . . . . . . C. granularis (Müll. Arg.) Zahlbr.
11(9)
Soralia mainly marginal, disc yellowish orange . C. flavocitrina (Nyl.) H. Olivier
Soralia of medullary origin, almost crater-like, disc ferruginous red . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . C. chrysophthalma Degel.
12(1)
Spores trilocular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Spores polaribilocular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
13(12) Apothecial disc different shades of brown, K−; differs entirely from C. cerina. . 14
Apothecial disc yellow to orange to rust brown, K+ purple; often confused with
C. cerina . . . . . . . . . . . . . . . . C. crocea (Kremp.) Hafellner & Poelt
14(13) Middle locule rhomboidal in shape . . . . . . . . . . C. brebissonii (Fée) Zahlbr.
Middle locule not rhomboidal . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15(14) Anthraquinone and/or hyaline crystals absent in amphithecium and parathecium
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. homologa Nyl.
Anthraquinone and/or hyaline crystals present in amphithecium and parathecium
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16(15) Anthraquinone crystals present in epihymenium; hypothecium with oil droplets;
middle locule of spores not elongated; thallus UV+ cream-yellow . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . C. jatolensis Joshi, Y. & Upreti
Anthraquinone crystals absent in epihymenium; hypothecium without oil droplets;
middle locule of spores elongated; thallus UV+ orange . . C. triloculans Zahlbr.
17(12) Thallus different shades of yellow to orange . . . . . . . . . . . . . . . . . . . 18
Thallus different shades of grey . . . . . . . . . . . . . . . . . . . . . . . . . . 22
18(17) Apothecia ± stipitate; ascus 12–16 spored; thallus Xanthoriella-type (i.e. lobate
species of Caloplaca resembling Xanthoria but devoid of lower cortex and rhizines)
. . . . . . . . . . . . . . . . . . . . . . . C. persica (Stein.) M. Stein. & Poelt
Apothecia sessile; ascus 8 spored; thallus not of Xanthoriella type . . . . . . . . 19
19(18) Thallus obligate on Juniperus bark or on dead wood; apothecial disc ferruginous red
but never with orange tinge . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Thallus not obligate over Juniperus bark or on dead wood; apothecial disc orangeyellow to brownish orange, rarely ferruginous red . . . . . . . . . . . . . . . 21
20(19) Mature thalli squamulose; apothecia deeply concave in the beginning; spores 10–14
× 4·5–8 µm; obligate on Juniperus. . . . . . . . . . . C. juniperi Poelt & Hinter.
Mature thalli continuous; apothecia plane from the beginning; spores 9–11 × 5–6
µm; obligate on dead wood . . . . . . . . C. himalayana Joshi, Y. & Upreti
21(19) Thallus greenish yellow to dull greyish, continuous to cracked areolate, prothallus
black; hypothecium with oil droplets; spores ellipsoid with thin end walls . . . .
. . . . . . . . . . . . . . . . . . . C. flavorubescens (Huds.) J. R. Laundon
Thallus golden yellow, continuous to areolate, prothallus absent; hypothecium
without oil droplets; spores ellipsoid with thickened end walls . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . C. alnetorum Giralt, Nimis & Poelt
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Vol. 41
22(17) Apothecial disc different shades of brown to blackish . . . . . . . . . . . . . . 23
Apothecial disc yellow to orange or red . . . . . . . . . . . . . . . . . . . . . . 27
23(22) Spores > 20 µm long . . . . . . . . . . . . C. yuennana (Zahlbr.) Poelt & Hinter.
Spores shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24(23) Disc dark brown to black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Disc pale brown with some reddish or orange tinge . . . . . . . . . . . . . . . 26
25(24) Epihymenium first K+ strong reddish purple then turning blue, thallus ± sorediate
. . . . . . . . . . . . . . . . . . . C. atrosanguinea (G. Merr.) I. M. Lamb
Epihymenium K+ purple not turning blue, thallus esorediate . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . C. rinodinopsis Poelt & Hinter.
26(24) Thallus granular to verruculose, prothallus present (bluish grey); spores with thin
end-walls. . . . . . . . . . . . . . . . . . . . . C. haematites (Chaub.) Zwack
Thallus continuous to areolate, prothallus absent; spores with thick end-walls . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . .C. pollinii (A. Massal.) Jatta
27(22) Asci usually 12–16 spored; thallus indistinct . . . . . C. cerinella (Nyl.) Flagey
Asci usually with 8 or fewer spores; thallus distinct. . . . . . . . . . . . . . . . 28
28(27) Margins whitish to dark bluish-grey, never yellow . . C. cerina (Hedw.) Th. Fr.
Margins usually yellow to orange at least in the upper part and around the disc, the
flanks often discolouring, whitish or greyish . . . . . . . . . . . . . . . . . . 29
29(28) Spores with thick end-walls (hour glass type); parathecium distinctly paraplectenchymatous and cupulate . . . . . . . . . . . . . . C. cupulata Poelt & Hinter.
Spores not with thick end-walls; parathecium neither paraplectenchymatous nor
cupulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30(29) Thallus granulose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Thallus smooth or crustose-membranaceous . . . . . . . . . . . . . . . . . . . 32
31(30) Prothallus absent, cortex of the amphithecium very thick; from Karakorum Range in
Pakistan . . . . . . . . . . . . . . . . . . . . . . C. baltistanica Poelt & Hinter.
Prothallus distinct, pale grey with bluish tinge; from Europe & Israel . . . . . . . .
. . . . . . . . . . . . . . . . . . C. haematites (Chaub. ex St.-Amans) Zwackh
32(30) Apothecia uniformly ferrugineous . . . . . . . . C. ferruginea (Huds.) Th. Fr.
Apothecia yellow to orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33(32) Apothecia uniformly ceraceous yellow, but the thin, sharp margin often discoloured,
becoming olivaceous or greyish black. . . . . . . . . . C. borealis (Vain.) Poelt
Apothecia yellow to orange, but margins usually not discoloured . . . . . . . . 34
34(33) Apothecia up to 1·5 mm in diam., distinctly zeorine, spores 14–22 × 7–11 µm . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. satparae Poelt & Hinter.
Apothecia mostly 0·2–0·7(–1) mm in diam., ± indistinctly zeorine, spores smaller
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35(34) Apothecia yellow to orange-yellow, margins somewhat brighter than discs; spores
9–13 × 5–7 µm . . . . . . . . . . . . . . . C. cerinelloides (Erichsen) Poelt
Apothecia yellowish orange to orange or brownish orange, but flanks of the margins
often becoming greyish; spores usually 11–14 × 5–7 µm. . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pyracea (Ach.) Th. Fr.
2009
Caloplaca himalayana—Joshi et al.
We thank Dr R. K. Tuli, Director, National Botanical
Research Institute, Lucknow, for providing laboratory
facilities and to Dr S. Nayaka, Dr R. Srivastava and Mr
Mukhtar Sheikh for collecting the specimens.
R
Alon, G. & Galun, M. (1971) The genus Caloplaca in
Israel. Israel Journal of Botany 20: 273–292.
Arup, U. (2006) A new taxonomy of the Caloplaca citrina
group in the Nordic countries, except Iceland.
Lichenologist 38: 1–20.
Awasthi, D. D. (1991) A key to the microlichens of
India, Nepal and Sri Lanka. Bibliotheca Lichenologica 40: 1–337.
Giralt, M., Nimis, P. L. & Poelt, J. (1992) Studien über
den formenkreis von Caloplaca flavorubescens in
Europa. Cryptogamie, Bryologie et Lichénologie 13:
261–273.
Hafellner, J. & Poelt, J. (1979) Die Arten der Gattung
Caloplaca mit pluriloculären Sporen (Meroplacis,
Triophthalmidium, Xanthocarpia). Journal of the
Hattori Botanical Laboratory 46: 1–41.
Hansen, E. S., Poelt, J. & Søchting, U. (1987) Die
Flechtengattung Caloplaca in Grönland. Meddelelser
om Grønland, Bioscience 25: 1–52.
Joshi, Y. & Upreti, D. K. (2007) New species and new
records of the lichen genus Caloplaca from India.
Lichenologist 39: 505–508.
Joshi, Y. & Upreti, D. K. (2008a) Sorediate and isidiate
species of the lichen genus Caloplaca (Ascomycetes,
Teloschistaceae) from India. Nova Hedwigia 86:
259–272.
Joshi, Y., Upreti, D. K. & Sati, S. C. (2008b) Three new
species of Caloplaca from India. Lichenologist 40:
535–541.
Joshi, Y. (2008c) Morphotaxonomic studies on lichen family
Teloschistaceae from India. Ph.D. Thesis. Kumaun
University.
Khodosovtsev, A. Y., Kondratyuk, S., Makarova, I. &
Oxner, A. (2004) Handbook of the Lichens of Russia.
9. Fuscideaceae, Teloschistaceae. St Petersburg:
Nauka Publishing.
255
Laundon, J. R. (1992) Caloplaca Th. Fr. (1860). In The
Lichen Flora of Great Britain and Ireland (O. W.
Purvis, B. J. Coppins, D. L. Hawksworth, P. W.
James & D. M. Moore, eds): 141–159. London:
Natural History Museum Publications.
Magnusson, A. H. (1944) Studies in the ferruginea –
group of the genus Caloplaca. Göteb. Kungl. Vetensk.
Vitt. Samh. Handl. Sjätte Följd. Ser. B, 3: 1–71.
Nash III, T. H. & Gries, C. (2002) Introduction. In
Lichen Flora of the Greater Sonoran Desert Region.
Vol. I. (T. H. Nash III, B. D. Ryan, C. Gries &
F. Bungartz, eds): 1–53. Tempe: Lichens Unlimited, Arizona State University.
Poelt, J. (1969) Bestimmungsschlüssel Europäischer Flechten. Lehre, Cramer.
Poelt, J. & Hinteregger, E. (1993) Beiträge zur Kenntnis
der Flechtenflora des Himalaya. Bibliotheca Lichenologica 30: 1–247.
Søchting, U. (1989) Lignicolous species of the lichen
genus Caloplaca from Svalbard. Opera Botanica 100:
241–257.
Wade, A. E. (1965) The genus Caloplaca Th. Fr. in the
British Isles. Lichenologist 3: 1–28.
Walker, F. J. & James, P. W. (1980) A revised guide to
microchemical techniques for the identification of
lichen products. Bulletin of British Lichen Society 46:
13–29.
Wetmore, C. M. (1994) The lichen genus Caloplaca in
North and Central America with brown or black
apothecia. Mycologia 86: 813–838.
Wetmore, C. M. (2001) The Caloplaca citrina group in
North America. Bryologist 104: 1–11.
Wetmore, C. M. (2004a) The isidiate corticolous
Caloplaca species in North and Central America.
Bryologist 107: 284–292.
Wetmore, C. M. (2004b) The sorediate corticolous
Caloplaca species in North and Central America.
Bryologist 107: 505–520.
Wetmore, C. M. (2007) Notes on Caloplaca cerina
(Teloschistaceae) in North and Central America.
Bryologist 110: 798–807.
Accepted for publication 2 March 2009