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The lichen family Physciaceae in Thailand—II. Contributions to the genus Heterodermia sensu lato

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A revision of 245 Heterodermia s.lat. collections from Thailand preserved in RAMK, herb. F. Schumm and herb. K. Kalb is presented. The 39 species found in this material are assigned to the genera Heterodermia Trevis. s.str. with a lower cortex and Pachysporaria-type ascospores (without sporoblastidia), Leucodermia Kalb, gen. nov., with foliose to subfruticose, linear-elongate, ribbon-like, dichotomously branched lobes, loosely attached to the substrate, without a lower cortex, apothecia often with pruinose discs and Polyblastidium-type ascospores (with sporoblastidia) and Polyblastidium Kalb, gen. nov. distinguished by a foliose thallus attached to the substrate, no lower cortex and mostly Polyblastidium-type ascospores. The remaining species are assigned to groups without a formal generic name, awaiting further results from molecular phylogenies. The Heterodermia comosa group contains species which are similar to Leucodermia, but differ in having a subfruticose or rosulate thallus with ascending, spathulate or paddle-shaped lobes and the Heterodermia obscurata group which contains species with a fluffy cottony woolly lower surface which is totally or at least partly impregnated by anthraquinones. Standardized descriptions are provided for all the better known Heterodermia species s.str. worldwide and all other species found in the material studied or reported for Thailand in the literature, as well as species which might be expected to occur in the country. The new species, Leucodermia borphyllidiata Kalb & Meesim, similar to L. boryi, but differing in having a phyllidiate thallus is described, and the new combinations, Leucodermia appalachensis (Kurok.) Kalb, L. arsenii (Kurok.) Kalb, L. boryi (Fée) Kalb, L. ciliatomarginata (Linder) Kalb, L. circinalis (Zahlbr.) Kalb, L. fertilis (Moberg) Kalb, L. leucomelos (L.) Kalb, L. lutescens (Kurok.) Kalb, L. vulgaris (Vain.) Kalb, Polyblastidium appendiculatum (Kurok.) Kalb, P. casarettianum (A. Massal.) Kalb, P. corallophorum (Taylor) Kalb, P. dendriticum (Pers.) Kalb, P. fragilissimum (Kurok.) Kalb, P. hypocaesium (Yasuda) Kalb, P. hypoleucum (Ach.) Kalb, P. japonicum (M. Satô) Kalb, P. magellanicum (Zahlbr.) Kalb, P. microphyllum (Kurok.) Kalb, P. neglectum (Lendemer, R.C. Harris & Tripp) Kalb, comb. nov. [Mycobank MB 813853; Basionym: Heterodermia neglecta Lendemer, R.C. Harris & Tripp, The Bryologist 110(3): 490 (2007)], P. propaguliferum (Vain.) Kalb, P. queenslandicum (Elix) Kalb, P. subneglectum (Elix) Kalb and P. violostriatum (Elix) Kalb, are made. Chaudhuria Zahlbr. is a synonym of Heterodermia s.str. and Chaudhuria indica Zahlbr. is an older synonym for Heterodermia verdonii Elix, but as the name Heterodermia indica (H. Magn.) D.D. Awasthi already exists for another species, the correct name for this taxon reverts to H. verdonii. Chemical analyses revealed that Heterodermia reagens (Kurok.) Elix must be placed into synonymy with Polyblastidium propaguliferum (Vain.) Kalb. Dichotomous keys for the identification of all species are provided. Characteristic TLC profiles for selected species are presented and Rf values for the most important terpenes and pigments are given for the first time in the standard solvents A, B’ and C. Photographs showing the species as well as characteristic structures in the genera and groups are also provided. A new epitype for H. comosa from Réunion is designated.
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Accepted by Christian Printzen: 22 Jul. 2015; published: 20 November 2015
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PHYTOTAXA
ISSN 1179-3155 (print edition)
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(online edition)
Copyright © 2015 Magnolia Press
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http://dx.doi.org/10.11646/phytotaxa.235.1.1
PHYTOTAXA
The lichen family Physciaceae in Thailand—II. Contributions
to the genus Heterodermia sensu lato
PACHARA MONGKOLSUK
1*
, SANYA MEESIM
1
,
VASUN POENGSUNGNOEN
1
, KAWINNAT
BUARUANG
1
,
FELIX SCHUMM
2
& KLAUS KALB
3,4
1
Lichen Research Unit, Biology Department, Faculty of Science, Ramkhamhaeng University, Ramkhamhaeng 24 road, Bangkok,
10240 Thailand.
2
Mozartstraße 9, D-73117 Wangen, Germany.
3
Lichenologisches Institut Neumarkt, Im Tal 12, D-92318 Neumarkt, Germany. Email: klaus.kalb@arcor.de
4
University of Regensburg, Institute of Plant Sciences, Universitätsstraβe 31, D-39040 Regensburg, Germany
*
Corresponding author—email: pm-tamrapap@ru.ac.th
Magnolia Press
Auckland, New Zealand
235
MONGKOLSUK ET AL.
2
Phytotaxa 235 (1) © 2015 Magnolia Press
PACHARA MONGKOLSUK, SANYA MEESIM,
VASUN POENGSUNGNOEN, KAWINNAT
BUARUANG,
FELIX SCHUMM
& KLAUS KALB
The lichen family Physciaceae in Thailand—II. Contributions to the genus Heterodermia sensu lato
(Phytotaxa 235)
66 pp.; 30 cm.
20 November 2015
ISBN 978-1-77557-839-0 (paperback)
ISBN 978-1-77557-840-6 (Online edition)
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UBLISHED I
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ISSN 1179-3155 (Print edition)
ISSN 1179-3163 (Online edition)
Phytotaxa 235 (1) © 2015 Magnolia Press
3
THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Abstract
A revision of 245 Heterodermia s.lat. collections from Thailand preserved in RAMK, herb. F. Schumm and herb. K.
Kalb is presented. The 39 species found in this material are assigned to the genera Heterodermia Trevis. s.str. with a
lower cortex and Pachysporaria-type ascospores (without sporoblastidia), Leucodermia Kalb, gen. nov., with foliose to
subfruticose, linear-elongate, ribbon-like, dichotomously branched lobes, loosely attached to the substrate, without a
lower cortex, apothecia often with pruinose discs and Polyblastidium-type ascospores (with sporoblastidia) and
Polyblastidium Kalb, gen. nov. distinguished by a foliose thallus attached to the substrate, no lower cortex and mostly
Polyblastidium-type ascospores. The remaining species are assigned to groups without a formal generic name, awaiting
further results from molecular phylogenies. The Heterodermia comosa group contains species which are similar to
Leucodermia, but differ in having a subfruticose or rosulate thallus with ascending, spathulate or paddle-shaped lobes
and the Heterodermia obscurata group which contains species with a fluffy cottony woolly lower surface which is totally
or at least partly impregnated by anthraquinones. Standardized descriptions are provided for all the better known
Heterodermia species s.str. worldwide and all other species found in the material studied or reported for Thailand in the
literature, as well as species which might be expected to occur in the country. The new species, Leucodermia
borphyllidiata Kalb & Meesim, similar to L. boryi, but differing in having a phyllidiate thallus is described, and the new
combinations, Leucodermia appalachensis (Kurok.) Kalb, L. arsenii (Kurok.) Kalb, L. boryi (Fée) Kalb, L.
ciliatomarginata (Linder) Kalb, L. circinalis (Zahlbr.) Kalb, L. fertilis (Moberg) Kalb, L. leucomelos (L.) Kalb, L.
lutescens (Kurok.) Kalb, L. vulgaris (Vain.) Kalb, Polyblastidium appendiculatum (Kurok.) Kalb, P. casarettianum (A.
Massal.) Kalb, P. corallophorum (Taylor) Kalb, P. dendriticum (Pers.) Kalb, P. fragilissimum (Kurok.) Kalb, P.
hypocaesium (Yasuda) Kalb, P. hypoleucum (Ach.) Kalb, P. japonicum (M. Satô) Kalb, P. magellanicum (Zahlbr.) Kalb,
P. microphyllum (Kurok.) Kalb, P. neglectum (Lendemer, R.C. Harris & Tripp) Kalb, comb. nov. [Mycobank MB
813853; Basionym: Heterodermia neglecta Lendemer, R.C. Harris & Tripp, The Bryologist 110(3): 490 (2007)], P.
propaguliferum (Vain.) Kalb, P. queenslandicum (Elix) Kalb, P. subneglectum (Elix) Kalb and P. violostriatum (Elix)
Kalb, are made. Chaudhuria Zahlbr. is a synonym of Heterodermia s.str. and Chaudhuria indica Zahlbr. is an older
synonym for Heterodermia verdonii Elix, but as the name Heterodermia indica (H. Magn.) D.D. Awasthi already exists
for another species, the correct name for this taxon reverts to H. verdonii. Chemical analyses revealed that Heterodermia
reagens (Kurok.) Elix must be placed into synonymy with Polyblastidium propaguliferum (Vain.) Kalb. Dichotomous
keys for the identification of all species are provided. Characteristic TLC profiles for selected species are presented and
R
f
values for the most important terpenes and pigments are given for the first time in the standard solvents A, B' and C.
Photographs showing the species as well as characteristic structures in the genera and groups are also provided. A new
epitype for H. comosa from Réunion is designated.
Key words: Appalachin, Asia, lichen chemistry, lichen pigments, lichenized Ascomycota, new species, taxonomy
Introduction
In tropical rainforests, members of Graphidaceae play a particularly important role in the composition of lichen
communities. Members of Physciaceae, however, dominate in open situations, such as on road side trees, in
thornbush forests, dry dipterocarp forests, coastal vegetation or dry evergreen forests (Cáceres et al. 2007, 2008),
where they grow on various substrates such as bark, wood, rocks or over mosses, especially if more or less basic
and/or nutrient rich environments. The genus Heterodermia Trevis. s.lat. is quite large and consists of ca. 115
mainly pantropical to subtropical species (www.mycobank.org, accessed: 11 February 2015) with a few species
extending into temperate or oceanic regions, e.g. H. speciosa and H. obscurata (Elix 2011c, Wirth et al. 2013).
The first species of Heterodermia recorded from Thailand were Heterodermia albicans (Vainio 1909)
published as Physcia crispa var. mollescens (Nyl.) Vain. from Trat Province, Koh Chang Island in the Gulf of
Thailand, H. leucomelos (L.) Poelt, H. hypoleuca (Ach.) Trevis. and H. speciosa (Wulfen) Trevis. from Chiang Mai
Province, Doi Suthep (Vainio 1921). Later, Kurokawa (1959, 1973) added H. diademata (Taylor) D.D. Awasthi [as
H. esorediata (Vain.) Du Rietz & Lynge] and H. hypocaesia (Yasuda) D.D. Awasthi to the lichen biota of Thailand.
Wolseley et al. (2002) added H. fragilissima (Kurok.) J.C. Wei & Y.M. Jiang and H. microphylla (Kurok.) Skorepa.
Subsequently, Aptroot et al. (2007) reported Heterodermia antillarum (Vain.) Swinscow & Krog, H. comosa
(Eschw.) Follmann & Redón, H. flabellata (Fée) D.D. Awasthi, H. galactophylla (Tuck.) Trevis., H. isidiophora
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(Vain.) D.D. Awasthi, H. japonica (K. Satô) Swinscow & Krog, H. obscurata (Nyl.) Trevis. and H. podocarpa
from different regions of Thailand. From the RAMK website (http://joomlas.ru.ac.th/lichen/index.php/herbarium/
checklist, accessed: 30 May 2013) the following species were added: H. appendiculata, H. hypochraea, H.
magellanica and H. pseudospeciosa. So before the present work, 20 species were known from the country.
Material and methods
All collections of Heterodermia s.lat. from Thailand housed in RAMK, hb. K. Kalb and hb. F. Schumm were
studied, but specimens from other herbaria including B, BM, CANB, F, G, GZU, H, M, NY, S, TUR, UPS and W
were also investigated for comparison. Type collections of many species mentioned in this paper were studied and
their terpene chemistry was investigated for the first time. The lichens were examined with a Wild M3Z Plan
stereomicroscope and an Olympus BH-A research microscope. Sections were prepared using a freezing microtome
Leitz Kryomat 1321, mounted in tap water and lactophenol cotton blue. Photographs were taken with a Canon EOS
600D digital camera and an Olympus BX51 light microscope. Identifications were done with the help of published
floras and keys, such as Awasthi (1960, 1973), Kurokawa (1962), Culberson (1967), Elix (2011c), Martins (2007),
Moberg (2011), Moberg & Nash (1999), Swinscow & Krog (1976), Schumm (2000, 2001a, 2001b), Schumm &
Aptroot (2012), Schumm & Elix (2014) as well as papers with descriptions of individual species. In addition to
these papers, specimens in the herbarium K. Kalb and reports in Aptroot (1988a, 1988b), Aptroot et al. (2007),
Awasthi (1960, 1973), Kashiwadani et al. (1990), Kurokawa (1973), Lendemer (2009), Lopez Figueiras (1986),
Marcano et al. (1996), Moberg (2004a, 2004b), Moberg & Purvis (1997), Thomson (1963), Trass (1992),
Weerakon & Aptroot (2013) and Wei et al. (2014) were consulted in order to establish the distribution of the
species. Natural compounds were characterized by thin-layer chromatography (TLC) according to the methods
standardized for lichen products by Elix (2014).
Results and discussion
In their checklist of Thai lichen species, Aptroot and Spier (2013) (http://www.tropicallichens.net/checklists,
accessed: 30. May 2014) mentioned a total of 27 Heterodermia species, including several literature based,
misidentified species, e.g. Heterodermia lepidota, Polyblastidium magellanicum and Leucodermia lutescens.
However, Heterodermia albicans, H. antillarum, H. galactophylla, H. palpebrata, Polyblastidium chilense, P.
dendriticum, P. fragilissimum, and P. microphyllum were not among the collections studied, but may well occur in
Thailand. They are all described below, except for P. dendriticum. The only report of this species from Thailand
(Kurokawa 1962) is probably from Laos (Wolseley et al. 2002), and furthermore, the description in Kurokawa
(1962) is based on at least two different species, one with and another without norstictic acid. The true nature of P.
dendriticum is still unclear (Swinscow & Krog 1976) and this problem will be addressed in a forthcoming paper
(Kalb & Schumm, in prep.).
In the present paper, we report 39 species for Thailand, including one new to science. Like the genera Physcia
and Pyxine, where terpenes play an important role in species delimitation (Elix et al. 2009; Kalb 1987, 1994;
Mongkolsuk et al. 2012), we have demonstrated the importance of triterpenes for the delimitation of species and
species groups in Heterodermia. This required running TLCs of many of the species in three solvent systems A, B’
and C, in order to record previously missing information (Table 1).
Three other well-studied tropical regions with modern regional monographs of Heterodermia [South America
(Moberg 2011), Africa (Moberg 2004; Swinscow & Krog 1988) and Australia (Elix 2011c)] are compared with
Heterodermia biota of Thailand and the study revealed that out of 39 species so far known from Thailand, 14 are
common with South America (33 species mentioned by Moberg 2011). However this number seems to be far too
low, as Martins (2007), in an unpublished thesis which covered only the state São Paulo of Brazil, recorded 46
species (sixteen of them new to science), 20 with Africa (26 species known) and 28 with Australia (42 species
known). six of the Thai species are not known from South America, Africa or Australia, namely Heterodermia
firmula, H. incana, H. pandurata, H. paradoxa, H. rubescens and Polyblastidium togashii, while 12 of the Thai
species occur in all of the four regions, namely Heterodermia albicans, H. comosa, H. diademata, H. flabellata, H.
Phytotaxa 235 (1) © 2015 Magnolia Press
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
isidiophora, H. obscurata, H. podocarpa, H. pseudospeciosa, H. speciosa, Leucodermia leucomelos,
Polyblastidium chilense and P. japonicum.
FIGURE 1. A. Heterodermia pseudospeciosa, longitudinal section through thallus showing the periclinal hyphae of the upper cortex
forming a prosoplectenchyma characterizing Heterodermia s.lat. (scale 20 µm). B. Physcia erumpens, longitudinal section through
thallus showing the anticlinal hyphae of the upper cortex forming a paraplectenchyma, characteristic for Physcia s.lat. C. Leucodermia
leucomelos f. albociliata, lobes dichotomously branched, ribbon-like with conspicuous long marginal rhizines. D. Polyblastidium
subneglectum (Elix) Kalb, comb. nov., Mycobank MB 813851, Basionym: Heterodermia subneglecta Elix, Australas. Lichenol. 68: 17
(2011), lobes irregularly to sympodially branched, appressed to the substrate not or only slightly ascending towards the tips. E.
Heterodermia galactophylla, tufts of apically ascending lobes with white marginal rhizines. F. Heterodermia comosa, thallus
subfruticose, lobes affixed mainly near the base, with paddle-shaped tips and white cilia.
MONGKOLSUK ET AL.
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Morphology and anatomy
The Thallus
Thallus lobate, irregular or forming rosettes 2–10 cm wide, these occasionally combining to form extensive
radiating mats or tangled clumps. Lobes discrete or contiguous, closely adnate and appressed to ascending and
loosely attached, linear or linear-cuneate to spathulate, ± dichotomously to irregularly branched, often with simple
or densely branched cilia-like marginal rhizines, seldom with cilia. Upper surface whitish, gray to yellow-gray,
plane to convex, or concave towards the periphery, dull or glossy, with or without isidia, soredia, phyllidia and
lobules; pseudocyphellae sometimes present along margins of the lobes and/or along the margins of the apothecia;
cortex composed of periclinal hyphae forming a prosoplectenchyma (Fig. 1A) [contrary to the upper cortex of
Physcia s.lat. which is formed from anticlinal hyphae, pattering a paraplectenchyma (Fig. 1B)]. Hypothallus
absent. Photobiont forming a continuous or seldom, e.g. in H. hypoleuca an interrupted and discontinuous layer.
Medulla well defined, white or partly yellow or shades of orange or brown.
Growth form:—Three different types of the growth form were found among Thai Heterodermia s.lat. species:
1. Leucomelos-growth form: Thallus loosely adnate or, in part, not attached to the substrate; dichotomously branched, lobes
mostly long, narrow, ecorticate below and with marginal rhizines. For the species with this growth form the new genus
Leucodermia is introduced (Fig. 1C).
2. Speciosa-growth form: Thallus entirely attached to the substrate; lobes sympodially or irregularly branched, mostly short
or sublinear-elongate, corticate or ecorticate below (Fig. 1D), with or without marginal rhizines. The species with a corti-
cate lower surface are retained in the genus Heterodermia s.str., while those with an ecorticate lower surface and without a
fluffy cotton-woolly lower surface, which is impregnated by anthraquinones, are placed in the new genus Polyblastidium.
3. Galactophylla-growth form: Thallus attached to the substrate in central parts or forming small rosettes or tufts of ascend-
ing lobes, ecorticate below, subfruticose to fruticose, always with white marginal rhizines (at least when young) (Figs. 1E,
1F).
Pseudocyphellae:—Pseudocyphellae have not been mentioned in Heterodermia by other workers [Elix (2011c),
Kurokawa (1962), Moberg (2004a, 2004b, 2011), Moberg & Nash (1999, 2002)] and Elix (2011c) explicitly writes
in the generic description—pseudocyphellae absent. But in some species, e.g. Heterodermia angustiloba and H.
verdonii, they are very distinct and clearly visible as white spots along the borders of the lobes as well as on the
crenate margins of the apothecia (Figs. 2A, 2B). Initially they look like soralia, but the above species are always
esorediate. Hence, it can be argued whether these structures are really pseudocyphellae, and if not, then a new term
must be created for them. At present we are treating them as pseudocyphellae.
Phyllidia (Squamules):—These are usually marginal, terete or flattened outgrowths of the thallus and in
contrast to isidia, they are never corticate throughout (Figs. 2C, 9F). Often they disintegrate and having the
appearance of coarse soredia or isidia. A total of 8 species with phyllidia are known from Thailand, namely, H.
coralloidea, H. dissecta, H. koyana, Leucodermia borphyllidiata, Polyblastidium appendiculatum, P.
fragilissimum, P. microphyllum and H. violostriatum.
Isidia:—These are ± terete outgrowths and completely corticate. Only two isidiate species occur in Thailand,
namely H. isidiophora and H. antillarum (Figs. 2D and 2E).
Soralia:—These may be marginal (e.g. in H. albicans; Fig. 2F), labriform and spathulate (e.g. in H.
galactophylla; Fig. 3A) or capitate (e.g. in Polyblastidium hypocaesium). Only a few species, especially in
Leucodermia and the comosa-group produce soralia on the arachnoid lower surface (e.g. in L. leucomelos or L.
vulgaris; Fig. 3B) or on the underside of the spoon-like lobe tips (e.g. in H. galactophylla and H. comosa). It
should be noted that the type specimens of Leucodermia boryi, L. leucomelos and L. vulgaris are fertile and lack
soralia.
Lower surface:—In general, the lower surface may be either corticate or ecorticate (e.g. Kurokawa 1962;
Moberg 2011; Swinscow & Krog 1976). When corticate a distinct, usually shiny layer is formed from periclinally
orientated strands of thick-walled hyphae that are clearly separated from the medulla, and the rhizines originate
from the entire lower surface. All species of Heterodermia s.str. have such a corticate lower surface (Fig. 3C).
However, when ecorticate, the lower surface is fluffy, cotton-woolly or arachnoid (Fig. 3D), and is formed from
medullary hyphae and may contain various pigments (see below). Meanwhile, there is a third, intermediate type of
lower surface, which is found predominantly in East Asian species producing dissectic acid and has been treated as
corticate in the literature, e.g. Elix (2011c). In this type it is quite difficult to discern whether there is a real cortex
Phytotaxa 235 (1) © 2015 Magnolia Press
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
like structure as in the species of Heterodermia s.str. or whether there is only a compact layer of medullary hyphae
which is not clearly separated from the medulla. For this type, the term pseudocortex is used in this paper (Figs. 3E
and 3F). It is characterized by being mostly ± rough, dull and always without lower surface rhizines. Unfortunately,
there are no DNA sequences available from species with this type of lower surface. In the near future, this species
group which is further characterized by the presence of dissectic acid and an East Asian distribution, may turn out
to be monophyletic and to deserve the rank of an independent genus.
FIGURE 2. A. Heterodermia verdonii, thallus lobes with pseudocyphellae along margins (see arrows), (scale 1 mm). B. Heterodermia
verdonii, pseudocyphellae along margin of apothecia (scale 1 mm). C. Polyblastidium fragilissimum, flattened marginal lobuli,
ecorticate below (scale 1 mm). D. Heterodermia isidiophora, ± cylindrical isidia, completely corticate. E. Heterodermia antillarum,
radial symmetric isidia, partly flattened, completely corticate. F. Heterodermia albicans, marginal soralia with granular soredia (scale
1 mm).
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FIGURE 3. A. Heterodermia galactophylla, soralia on lower surface of spathulate lobes (scale 1 cm). B. Leucodermia vulgaris,
soralia on lower surface; note also the superimposed layer of cotton-woolly hyphae impregnated by anthraquinones. C. Heterodermia
pseudospeciosa, corticate ± glossy lower surface with laminal and marginal rhizines (scale 1 mm). D. Polyblastidium japonicum,
ecorticate arachnoide lower surface only with marginal rhizines. E. Heterodermia koyanoides, pseudocortex with compact lower
surface, ± dull, not glossy, without laminal rhizines (scale 1 mm). F. Heterodermia koyana, longitudinal section through thallus;
pseudocortex without border between medulla and lower surface (scale 100 µm).
Cilia and rhizines:—These two terms are variably and arbitrarily used in the literature. For example, Elix
(2011a, 2011b, 2011c) used the term cilia for the hairy outgrowths in Leucodermia boryi but rhizines for similar
structures in H. flabellata, while all hairy outgrowths from the upper (or marginal) cortex are called cilia by
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Moberg (2011). Since the presence or absence of cilia and rhizines are important key characters, a consistent
terminology is required for them, and in this paper, we use the following definitions for these structures.
Cilia: are hairy outgrowths originating from the upper cortex of the thallus or from the receptacle of the
apothecia (Figs. 4A and 4B) and may be ramified or not. Cilia occur mainly in species with the Galactophylla-
growth form. They are mostly white (very seldomly short and black), but may become brownish at the tips with
age. Elix (2011c) and Moberg (2011) also treated black or especially long, lash-like excrescences from the upper
cortex as cilia, but these were called marginal rhizines by Kurokawa (1962).
FIGURE 4. A. Heterodermia comosa, cilia orginating from the upper surface (scale 2 mm). B. Heterodermia comosa, cilia orginating
from the exciple of the apothecia (scale 1 mm). C. Heterodermia echinata, white marginal rhizines, some becoming dark at the tips
(scale 5 mm). D. Leucodermia leucomelos f. albociliata, ± white, simple marginal rhizines (scale 5 mm). E. Leucodermia boryi, black
branched marginal rhizines (scale 0.5 cm). F. Polyblastidium propaguliferum, black, branched, marginal rhizines (scale 1 mm).
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FIGURE 5. A. Heterodermia pseudospeciosa, lower surface rhizines originating from the corticate lower surface (scale 1 mm). B.
Leucodermia boryi, apothecia with pruinose disks and a star-like margin with triangular reflexed lobules (scale 1 mm). C.
Heterodermia angustiloba, margin of apothecia crenulate or with small lobules (scale 2 mm). D. Polyblastidium corallophorum
(Taylor) Kalb, comb. nov., Mycobank MB 813852, Basionym: Parmelia corallophora Taylor, London J. Bot. 6: 164 (1847), exciple of
apothecia with isidia (scale 2 mm). E. Polyblastidium squamulosum, exciple of apothecia with lobules (scale 2 mm). F. Leucodermia
ciliatomarginata (Linder) Kalb, comb. nov., Mycobank MB 813834, Basionym: Anaptychia ciliatomarginata Linder, Proc. Calif.
Acad. Sci. IV, 21: 217 (1934), exciple of apothecia with cilia, becoming black at the tips (scale 2 mm).
Marginal rhizines: these are hairy outgrowths somewhat similar to cilia, but originate from the corticate
margin of the thallus (i.e. marginal cortex) and may be ramified or not. Marginal rhizines occur mainly in species
with Speciosa- and Leucomelos-growth form. They may be white in color [e.g. H. echinata or Leucodermia
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
leucomelos f. albociliata (Figs. 4C and 4D)] or black [e.g. L. boryi (Fig. 4E) and Polyblastidium propaguliferum
(Fig. 4F)].
Lower surface rhizines: these are hairy outgrowths originating from the lower surface of the thallus and may
be ramified or not. Lower surface rhizines occur exclusively in the species of Heterodermia s.str. with a corticate
lower surface (Fig. 5A).
Remarks:—There are some exceptions to the terminology proposed above. In very few species, mainly with
the Galactophylla-growth form (e.g. H. trichophora and H. comosa), the same structures are called marginal
rhizines when growing along the margins of the lobes but cilia when growing on the upper cortex of the lobes or
from the exciple of the apothecia. Consequently, the terminology proposed here does have some shortcomings, but
still can be useful for discriminating species.
Ascomata:—Apothecia laminal or terminal, rounded, sessile to stipitate, lecanorine; disc pale to dark brown or
black, concave to ± plane, pruinose or not; thalline exciple prominent or reflexed (Fig. 5B), margin entire, crenulate
(Fig. 5C), phyllidiate, sorediate, isidiate (Fig. 5D) or lobulate (Fig. 5E), sometimes with whitish or, very rarely,
blackish cilia (Fig. 5F). Epihymenium pale brown to brown-black; hymenium colorless, I+ blue; hypothecium
usually colorless, rarely pale yellowish. Paraphyses unbranched or with few branches; apical cells brown and
expanded. Asci cylindrical to subclavate or clavate, Physcia-type, with 8 ascospores; apex wall layers thickened;
apex amyloid, with a distinct axial mass (Hafellner et al. 1979). Ascospores Pachysporaria- (without
sporoblastidia; Fig. 6A) or Polyblastidia-type (with 1 or more sporoblastidia; Fig. 6B), gray-brown to brown or
dark brown, ellipsoidal to oblong or fusiform, 1-septate, ± constricted at the septum, very thick-walled; internal
apical wall thickenings becoming distinct only after the septum is formed (type-A ontogeny); torus thin or absent;
spore surface smooth.
Conidiomata:—Pycnidia, immersed in the thallus or becoming emergent (Fig. 6C); conidiogenous cells in
branched chains, short, cylindrical, enteroblastic. Conidia bacilliform to short-cylindrical.
Chemistry
Terpenes, depsidones, depsides:—As in the other foliose genera of Physciaceae, terpenes play an important role
in species delimitation in Heterodermia (for Physcia see Elix et al. 2009; for Pyxine see Kalb 1987, 1994 and
Mongkolsuk et al. 2012). However, before the pivotal papers by Elix (2010, 2011a, 2011b, 2014), only few
terpenes such as zeorin, spathulene (= spathulin) and japonene (= japonin) were known . As zeorin is present in all
Heterodermia species (and many Physcia species), it is not useful for species or generic delimitation. The major
triterpene present in H. spathulifera Moberg & Purvis was called spathulin by Moberg & Purvis (1997), but this
name had been used previously for a pseudoguaianolide present in the higher plant genus Gaillardia (Asteraceae)
by Herz & Srinivasan (1974). We could have used the systematic name, 16β-acetoxyhopane-22-ol, for it but
propose it be termed spathulene instead. The same applies for a ± common triterpene present in Heterodermia,
initially called japonin by Elix (2011a). However, the name was introduced earlier for an alkaloid present in the
higher plant Orixa japonica (Rutaceae) by Hā-Huy-Kê & Luckner (1970). Therefore, we propose that this
triterpene be termed japonene. In addition to zeorin, the depside atranorin is present in all species of Heterodermia
s.lat., and is not a useful chemical character for delimiting species or genera.
However, by following Elix (2011a, 2011b), we found the presence or absence of the depsidone norstictic acid
and the β-orcinol meta-depside dissectic acid (in some Asian species, except H. neglecta from N-America) as a
valuable character for separating similar species. Until the marvellous work by Elix (2014), the R
f
values for most
of the terpenes in the standard solvents A and B’ could not be determined from the literature, so we have
summarized them together with other relevant substances for Heterodermia s.lat. (Table 1).
Pigments:—Many species, especially those without a lower cortex, produce characteristic yellow, orange,
brownish, red or even blue-green colored pigments. The pigments as well as their distribution on the lower surface
or in the medulla are distinctive and in many cases can be used to separate similar species (Table 3).
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TABLE 1. R
f
values for taxonomically important substances occurring in Heterodermia s.lat.
The following mixtures of the solvents were used:
A: toluene 180 ml, dioxane 45 ml, glacial acetic acid 5 ml
: hexane 140 ml, methyl tert-butyl ether 72 ml, formic acid 18 ml
C: toluen 195 ml, glacial acetic acid 35 ml
Usually, solvent C provides a very good discrimination of lichen substances and it is particularly stable and
reliable, and Elix (2014) also recomments its use for separating hopane triterpenoids. But the four most common
and major terpenes in Heterodermia are best separated in solvent A (Table 2). Unfortunately, in solvent B',
japonene and 6α-acetoxyhopane-16β,22-diol have identical R
f
values and leucotylin is obscured because of having
nearly the same R
f
value as 16β-acetoxyhopane-6α,22-diol, while in solvent C, the R
f
values of japonene and 16β-
acetoxyhopane-6α,22-diol are identical. Even if the R
f
values are not always constant, their sequence stays the
same, but co-chromatography with known species can be very helpful.
Solvent A B' C
Atranorin 75 73 79 yellow
6α-Acetoxyhopane-22-ol 62 56 53
Anaptychin-5 61 54 53
Dissectic acid 58 51 46 brown
Emodin 54 58 41 yellow
7-Chloroemodin 52 55 47 yellow
Spathulene =16β-Acetoxyhopane-22-ol 52 51 58
Zeorin 50 42 43
5,7-Dichloroemodin 49 52 63 orange
6α,16β-Diacetoxyhopane-22-ol 48 41 42 (in H. tremulans)
Emodic acid 47 39 33 yellow
Testacein 42 25 20 pink
16β-Acetoxyhopane-6α,22-diol 41 33 32
Norstictic acid 40 32 30 yellow
6α-Acetoxyhopane-16β,22-diol 38 36 34
Japonene 35 36 32
α-Acetylhypoconstictic acid. 34 14 16 red
Leucotylin 21 31 20 gray
Salazinolide 18 27 8 gray
Appalachin 17 13 9 bluish gray
Hybocarpone (depending on concentration) 15–23 46 19 red
Hypoconstictic acid 11 5 4 red
Salazinic acid 10 7 4 orange
Connorstictic acid 8 11 3 yellow
Desmethylhybocarpone = Norhybocarpone 5 37 5 red
3-O-Methylsalazinic acid 3 7 4 red
Consalazinolide 1 14 4 gray
3-O-Methylconsalazinic acid 1 1 1 orange-red
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
TABLE 2. R
f
values of 4 major terpenes occurring in Heterodermia s.lat.
TABLE 3. R
f
values of pigments reported from Heterodermia s.lat.
Taxonomy
Key to the genera and groups of Heterodermia sensu lato
1. Lower cortex or pseudocortex present, lower cortex shiny and often with rhizines.................................Heterodermia s.str.
- Lower cortex absent, lower surface mostly dull and never with rhizines.............................................................................. 2
2. Thallus lobes linear elongate, ± dichotomously branched with conspicuous long marginal rhizines, Leucomelos-growth
form .....................................................................................................................................................................Leucodermia
- Thallus lobes not distinctly elongate, ± subdichotomously or irregularly branched, obviously foliose, color of marginal
rhizines variable, Speciosa-growth form, or thallus forming small rosettes or tufts of ascending lobes, or subfruticose to
fruticose, mostly with white marginal rhizines, Galactophylla-growthform......................................................................... 3
3. Thallus, at least when young, with white marginal rhizines, somewhat darkened at the tips... Heterodermia comosa-group
- Thallus with marginal rhizines, which soon become jet-black.............................................................................................. 4
4. Lower surface with superimposed layer of cotton-woolly hyphae, impregnated by yellow to orange-brown pigments ........
................................................................................................................................................Heterodermia obscurata-group
- Lower surface formed by ± arachnoid hyphae, without pigments or pigments only patchy ...........................Polyblastidium
I. The lichen genus Heterodermia s.str.
Heterodermia Trevis., Atti Soc. Ital. Sci. Nat. 11: 613 (1868)
Pseudophyscia Müll. Arg., Bull. Herb. Boissier 2, app. 1: 10 (1894a). Type species (selected here) Pseudophyscia speciosa (Wulfen)
Müll. Arg.—Chaudhuria Zahlbr., Ann. Mycol. 30 (5–6): 433 (1932). Type species: Chaudhuria indica Zahlbr., Ann. Mycol.
30(5–6): 434 (1932).
The genus as defined here is characterized by a thallus with lobes appressed to the substrate, not or only slightly
ascending towards the tips, a prosoplectenchymatous upper and lower cortex, the lower surface with rhizines
(rarely only marginal); apothecia laminal; ascospores thick walled, brown 1-septate, without sporoblastidia
(Pachysporaria-type). It conforms to the species with a lower cortex of Anaptychia sect. Anaptychia ser. Speciosae
(Kurokawa 1962: 23) and Anaptychia Körb. subgen. Heterodermia (Trev.) Kurok. ser. Speciosae (Jatta) Kurok.,
Kurokawa (1973: 585).
Type species (selected by Poelt 1965: 31): Heterodermia speciosa (Wulfen) Trevis., Atti Soc. Ital. Sci. Nat. 11:
Solvent A B' C
Zeorin 50 42 43
16β-Acetoxyhopane-6α,22-diol 41 32 32
6α-Acetoxyhopane-16β,22-diol 38 36 34
Japonene 35 36 32
Solvent A B' C
Obscurata yellow (= 7-Chloroemodin) 52 55 47 yellow
Obscurata orange (= 5,7-Dichloroemodin) 49 52 63 orange
Lutescens 381327yellow
Vulgaris high 41 23 29 violet
Vulgaris low 41 14 23 pink
Boryquinone 9 44 8 red
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614 (1869 [1968]); Lichen speciosus Wulfen, in Jacquin, Coll. Bot. 3: 119 (1789); Anaptychia speciosa (Wulfen)
A. Massal.; Mem. Lich.: 36 (1853); Physcia speciosa (Wulfen) Nyl., Actes Soc. Linn. Bordeaux 21: 307 (1856);
Pseudophyscia speciosa (Wulfen) Müll. Arg., Bull. Herb. Boissier 2, app. 1: 40 (1894b); Pseudophyscia speciosa
(Wulfen) Müll. Arg., Bull. Soc. Roy. Bot. Belgique 32: 129 (1893b), invalidly published (Art. 35.1., Art. 38.1., Art.
38.5.; no description provided for the generic name and the genus was not monotypic). Chaudhuria Zahlbr., Ann.
Mycol. 30(5–6): 433 (1932). Type species: Chaudhuria indica Zahlbr., Ann. Mycol. 30(5–6): 434 (1932).
Key to some common worldwide species of Heterodermia s.str.
(TLC, at least in solvent A is necessary for definitive identification)
1. Thallus without vegetative propagules .................................................................................................................................. 2
- Thallus with isidia, phyllidia or soralia.................................................................................................................................. 9
2. Medulla at least partly yellowish, orange or brownish yellow, K+ purple ............................................................................ 3
- Medulla completely white...................................................................................................................................................... 5
3. Medulla with dissectic acid......................................................................................... H. albidiflava (Kurok.) D.D. Awasthi
- Medulla without dissectic acid............................................................................................................................................... 4
4. Lobes 0.3–1.2 mm wide, apothecia 0.5–2.2 mm wide, India, Nepal, Thailand.............................. H. firmula (Nyl.) Trevis.
- Lobes 0.7–2 mm wide, apothecia 1.5–6 mm wide, North America: Texas, Mexico ............ H. rugulosa (Kurok.) Wetmore
5. Medulla without depsidones, only terpenes present ..................................................... H. diademata (Taylor) D.D. Awasthi
- Medulla with both terpenes and depsidones .......................................................................................................................... 6
6. Medulla with salazinic and hypoconstictic acids........................................................................ H. tropica (Kurok.) Sipman
- Medulla without salazinic and hypoconstictic acids.............................................................................................................. 7
7. Medulla without dissectic acid, with norstictic acid, with small lateral lobules and pseudocyphellae ...................................
.................................................................................................................................... H. rubescens (Räsänen) D.D. Awasthi
- Medulla with dissectic acid, norstictic acid present or absent............................................................................................... 8
8. Medulla K+ dark yellow; norstictic acid absent ........................................................................................... H. verdonii Elix
- Medulla K+ yellow then red; norstictic acid present ........................................... H. angustiloba (Müll. Arg.) D.D. Awasthi
9. Thallus sorediate .................................................................................................................................................................. 10
- Thallus with isidia or phyllidia ............................................................................................................................................ 13
10. Medulla without depsidones, only terpenes present .............................................................................................................11
- Medulla with depsidones...................................................................................................................................................... 12
11. Lobes linear-elongate, spores 30–37 × 14–18 μm, 16β-acetoxyhopane-6α,22-diol absent....... H. speciosa (Wulfen) Trevis
- Lobes short, often flexuose-bent, spores 23–32 × 12–14 μm, 16β-acetoxyhopane-6α,22-diol present as major terpene.......
.................................................................................................................................... H. tremulans (Müll. Arg.) W.L. Culb.
12. Medulla with hypoconstictic and salazinic acids, soralia ± linear and continuous along margins of the lobes, not apical.....
..................................................................................................................................... H. albicans (Pers) Swinscow & Krog
- Medulla without hypoconstictic and salazinic acids, but with norstictic acid (major), soralia apical on main lobes and on
short lateral lobes, in delimited areas, ± labriform................................................... H. pseudospeciosa (Kurok.) W.L. Culb.
13. Thallus with distinct isidia ................................................................................................................................................... 14
- Thallus with phyllidia, sometimes becoming sorediate-isidiate-granular............................................................................ 16
14. Medulla with both salazinic and hypoconstictic acids ........................................... H. antillarum (Vain.) Swinscow & Krog
- Medulla with or without salazinic acid but lacking hypoconstictic acid ............................................................................. 15
15. Medulla with salazinic acid, isidia mostly laminal ............................................................ H. granulifera (Ach.) W.L. Culb.
- Medulla without salazinic acid, with terpenes only, isidia mostly marginal......... H. isidiophora (Vain.) Swinscow & Krog
16. Medulla with dissectic acid, with or without norstictic acid................................................................................................ 17
- Medulla without dissectic acid............................................................................................................................................. 18
17. Norstictic acid present...................................................................................................... H. dissecta (Kurok.) D.D. Awasthi
- Norstictic acid absent ....................................................................................................................... H. koyana (Kurok.) Elix
18. Norstictic acid present.............................................................................................................................. H. coralloidea Elix
- Norstictic acid absent, with terpenes only............................................................................................................................ 19
19. Thallus 2–5 cm wide, lobes 0.1–0.5 mm wide, phyllidia becoming sorediate-isidiate-granular; medulla with 6α-acetoxy-
hopane-16β,22-diol present as major terpene....................................................................................... H. isidiophorella Elix
- Thallus 5–20 cm wide, lobes 0.7–2.5 mm wide, phyllidia not becoming sorediate-isidiate-granular; medulla with 16β-ace-
toxyhopane-6α,22-diol present as major terpene .................................................................... H. lepidota Swinscow & Krog
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
1. Heterodermia albicans (Pers.) Swinscow & Krog, Lichenologist 8: 113 (1976) (Fig. 2F)
Parmelia albicans Pers., Ann. Wetterauischen Ges. gesammte Naturk. 2(1): 17 (1810). Type:—ST. DOMINGO [Haiti]. Poiteau s.n.
(holotype L, fide Swinscow & Krog 1988); Physcia albicans (Pers.) J.W. Thomson, Beih. Nova Hedwigia 7: 88
(1963).—Parmelia domingensis Ach. (nom. nov. illegit.), Syn. Lich.: 212 (1814); Hagenia domingensis (Ach.) De Not., Giorn.
Bot. Ital., 2 (1), 1: 186 (1846); Anaptychia domingensis (Ach.) A. Massal., Mem. lich.: 39 (1853); Squamaria domingensis (Ach.)
A. Massal., Atti Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 250 (1860); Physcia stellaris (L.) Nyl. var. domingensis (Ach.) Tuck.,
Proc. Amer. Acad. Arts 4: 396 (1860).—For further synonymy see Swinscow & Krog (1976).
Thallus foliose, orbicular to irregularly spreading, comparatively small, adnate, to 4 cm wide. Lobes 0.5–1.0 mm
wide, to ca. 1.5–2.5 mm wide at the tips, ± plane to weakly convex, sublinear-elongate, usually richly
dichotomously branched; lobe apices not ascending, eciliate, sorediate. Upper surface whitish gray to brownish
gray, darker at the apices, occasionally sparsely pruinose; soralia white to bluish gray, forming small ± continous
marginal soralia towards the thallus centre. Medulla white. Lower surface corticate, whitish to pale brown, rarely
dark gray, smooth to corrugate. Lower surface rhizines frequent, evenly distributed on the lower surface, simple or
irregularly branched, usually short (ca. 1 mm long), pale to dark brown or black. Apothecia very rare, laminal,
substipitate, 0.5–3.0 mm wide; margin becoming sorediate; disc concave, dark brown, epruinose. Ascospores
Pachysporaria-type, narrowly ellipsoidal, 21–32 × 7–14 µm. Pycnidia ± common, immersed, with black weakly
protruding apices; conidia bacilliform, 4–6 × 1 µm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (minor), leucotylin (minor), 6α-
acetoxyhopane-16β,22-diol (trace), salazinic acid (major), hypoconstictic acid (submajor), 3-O-methylconsalazinic
acid (minor), consalazinic acid (minor) and -acetylhypoconstictic acid (minor).
Distribution and habitat:—This rare species occurs on bark in Trat Province, also rarely on rocks in
Australia, North, Central and South America, Macaronesia, South Africa and China.
Remarks:—Heterodermia albicans is characterized by lobes with marginal soralia towards the thallus centre
and the presence of salazinic acid and hypoconstictic acids in the medulla.
Material reported from Thailand (Wolseley et al. 2002: 27):—Trat Province: Koh Chang Island, Gulf of
Thailand, ad corticem arboris prope Lem Ngob, J. Schmidt 9, 32.
Specimens examined: GUATEMALA. Suchitepéquetz: National Highway 11, ca. 20 km N of Patulul, on
deciduous trees alongside the street, 400 m, 14°40' N, 91°07' W, 16 January 1979, K. Kalb & G. Plöbst (herb. Kalb
39583) )—BRAZIL. Paraná: Guaíra, in a rainforest along Rio Paraná, 200 m, 24°05’ S, 54°15’ W, 9 August 1980,
K. Kalb & m, Marcelli (herb. Kalb 39569)—BRAZIL. São Paulo: Serra do Mar; ca. 20 km E of Cruzeiro, in a light
and relatively dry secondary rainforest, 1600 m, 22°33' S, 44°45' W, 3 November 1978, K. Kalb & G. Plöbst (herb.
Kalb 40303).
Heterodermia albidiflava (Kurok.) D.D. Awasthi, Geophytology 3(1): 113 (1973)
Anaptychia albidiflava Kurok., Beih. Nova Hedwigia 6: 42 (1962); Type:—INDIA. Darjeeling: above Kurseong, on tree trunk, 1680
m, Awas th i 3912 (holotype LWG-Awas., fide Awasthi 1962).
Remarks:—Species not recorded for Thailand are not numbered, but selected specimens studied for
morphological, anatomical and chemical characters are cited. This species is almost identical to H. firmula (see
below), but differs chemically in containing dissectic acid. We have not seen any authentic material. All specimens
identified as H. albidiflava studied by us turned out to be H. firmula and lacked dissectic acid.
2. Heterodermia angustiloba (Müll. Arg.) D.D. Awasthi, Geophytology 3(1): 113 (1973) (Fig. 6D)
Physcia speciosa (Wulfen) Müll. Arg. var. angustiloba Müll. Arg., Flora 66: 78 (1883). Type:—AUSTRALIA. Queensland: prope
Toowoom ba, Hartmann (holotype G, fide Elix 2011c); Pseudophyscia speciosa (Wulfen) Müll. Arg. var. angustiloba (Müll. Arg.)
Müll. Arg., Bot. Jahrb. 20: 259 (1894c); Anaptychia speciosa (Wulfen) A. Massal. var. angustiloba (Müll. Arg.) Zahlbr., Cat. lich.
univ. 7: 741 (1931); Anaptychia angustiloba (Müll. Arg.) Kurok., Beih. Nova Hedwigia 6: 39 (1962).
MONGKOLSUK ET AL.
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FIGURE 6. A. Polyblastidium hypoleucum, ascospores (mostly) without sporoblastidia (scale 10 µm). B. Polyblastidium
magellanicum, Polyblastidia-type ascospores with one or more sporoblastidia (scale 10 µm). C. Heterodermia tropica, black
conidiomata, immersed in the thallus or slightly emergent (scale 1 mm). D. Heterodermia angustiloba (scale 5mm). E. Heterodermia
antillarum (scale 1 mm). F. Heterodermia coralloidea, holotype (scale 5 mm).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 4–6 cm wide, but sometimes thalli coalescing
and forming larger colonies. Lobes short and narrow, 0.5–1.2 mm wide, rather flexuose and discrete at the
periphery, ± plane to weakly convex, sublinear-elongate, repeatedly di- or trichotomously branched, eciliate,
lacking isidia, phyllidia and soralia, with distinct pseudocyphellae along the margins of the lobes. Upper surface
grayish white to gray, epruinose. Medulla white. Lower surface with pseudocortex, whitish to pale brown, dark
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
gray-brown near the centre, smooth. Lower surface rhizines absent or rare, marginal rhizines frequent, concolorous
with the thallus or darkening near the apices, not or irregularly branched, 1–3 mm long. Apothecia laminal,
substipitate to subsessile, 0.5–2.5 mm wide; margin ± crenate with pseudocyphellae; disc concave, brown to
blackish brown, weakly gray-pruinose. Ascospores Pachysporaria-type, ellipsoidal, 25–30 × 13–15 μm. Pycnidia
common, immersed; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), dissectic acid (submajor), 16β- acetoxyhopane-6α,22-diol (major),
leucotylin (minor), 6α-acetoxyhopane-16β,22-diol (major), 6α,16β-diacetoxyhopane-22-ol (minor), norstictic acid
(major), connorstictic acid (minor), salazinic acid (minor or trace).
Distribution and habitat:—Occurs on bark and rocks in lower montane scrub, hill evergreen forest and lower
montane rainforest, also in Australia, India, Nepal, Vietnam, China and Japan; new addition to the Thai lichen
biota.
Remarks:—Characterized by the narrow lobes with marginal rhizines, a lower surface with a pseudocortex,
rhizines sparse or absent, the absence of isidia, phyllidia and soralia and the presence of norstictic and dissectic
acids.
Material from Thailand examined:—Phitsanulok: Phu Hin Rongkla, forest protection unit Man Daeng, in
lower montane forest, on rock, 1610 m, 16°55'51'' N, 101°03'04'' E, 16 July 2004, P. Mongkolsuk RU-PM 255/1,
(RAMK 21811); Loei: Phuluang Wildlife Sanctuary, along the trail in Kok Promachan area, in lower montane
scrub, on rock, 1560 m, 17°16'39'' N, 101°31'38'' E, 28 August 2008, S. Meesim & K. Buaruang, RU-MS 32.1,
(RAMK 21031); Nan: Doi Phu Kha, area of view point at Doi Phu Kha, in hill evergreen forest, on bark of Toona
ciliata, 1470 m, 19°18'13'' N, 101°09'55'' E, 10 May 2006, S. Meesim RU-DK 26, (RAMK 21842).
3. Heterodermia antillarum (Vain.) Swinscow & Krog, Lichenologist 8: 114 (1976) (Fig. 6E)
Anaptychia granulifera (Ach.) Vain. var. antillarum Vain., Ann. Acad. Sci. Fenn., Ser. A 6(7): 63 (1915). Type:—INDIAS OCC.
[Lesser Antilles]. Guadalupa [Guadeloupe]: Camp-Jacob, sur les Inga laurifolia, 500–800 m, Duss 459 pr. p. (lectotype, selected
by Kurokawa 1962, TUR-Vain 07737!); Anaptychia antillarum (Vain.) Kurok., J. Hattori Bot. Lab. 37: 596 (1973).
Thallus foliose, orbicular to irregularly spreading, adnate, comparatively small, up to 4 cm wide. Lobes 0.5–1.0
mm wide, to ca. 1.5–2.5 mm wide at the tips, ± plane, sublinear-elongate, usually richly sympodially branched, not
ascending, eciliate, isidiate. Upper surface whitish gray to brownish gray, darker at the lobe apices, occasionally
sparsely pruinose; isidia usually marginal, more rarely laminal, marginal isidia prominent at the periphery of the
thallus, flattened to cylindrical, eventually becoming coralloid. Medulla white, partly orange (decomposed
salazinic acid?). Lower surface corticate, whitish to pale brown, rarely dark gray, smooth to corrugated. Lower
surface rhizines frequent, evenly distributed on the lower surface, simple or irregularly branched, usually short (ca.
1 mm long), pale to dark brown or black. Apothecia rare, laminal, adnate to substipitate, 0.5–3.0 mm wide; margin
densely mealy pruinose, becoming isidiate; disc concave, dark brown, epruinose. Ascospores Pachysporaria-type,
ellipsoidal, 23–30 × 12–18 μm. Pycnidia ± common, immersed; conidia bacilliform, 4–6 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (minor), leucotylin (minor), salazinic
acid (major), hypoconstictic acid (submajor), 3-O-methylconsalazinic acid (minor), consalazinic acid (minor).
Distribution and habitat:—Usually on bark in ± open situations in dry evergreen forest with dominating
Fagaceae, also rarely on rock, also in Australia, Central America, the Caribbean, East Africa, South Africa and the
Galapagos Islands.
Remarks:—characterized by the lobes with marginal and more rarely additional laminal isidia and the
presence of salazinic acid and hypoconstictic acid in the medulla. The chemistry of this species is ± the same as that
of H. albicans and it may be considered its non-sorediate, isidiate counterpart. H. tropica also has the same
chemistry, but this species lacks any vegetative propagules.
Material reported from Thailand (Aptroot et al. 2007: 104):—Chiang Mai: Doi Suthep-Pui National Park,
Doi Suthep ridge, on bark of Fagaceae, Aptroot 55218.
Specimen examined: GUATEMALA. Retalhuleu: on free standing deciduous trees alongside a street SW of
Retalhuleu, 500 m, 14°32’ N, 91°41’ W, 14 January 1979, K. Kalb & G. Plöbst (herb. Kalb 40303).
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4. Heterodermia coralloidea Elix, Australas. Lichenol. 69: 12 (2011b) (Fig. 6F)
Type:—AUSTRALIA. Australian Commonwealth Territory: Jervis Bay, Stony Creek, 34 km SE of Nowra, 2 m, 35°10' S, 150°45' E,
on sandstone in dry sclerophyll forest beside creek with numerous shrubs and Livistonia, 8 November 1990, J. A. Elix 26421
(holotype CANB!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 2–7 cm wide. Lobes 0.7–1.5 mm wide, ± plane,
sublinear-elongate, sympodially or irregularly branched, ± discrete to contiguous at the periphery, with short lateral
lobes; apices not ascending, eciliate, phyllidiate. Upper surface whitish gray; phyllidia mostly marginal, more
rarely laminal, dissected and erumpent, forming dense, coralloid pseudoisidia that cover most of the upper surface,
rarely becoming granular. Medulla white. Lower surface with a pseudocortex, white to cream or pale tan near the
centre. Lower surface rhizines rare, whitish to cream, concolorous with the thallus or becoming pale to dark brown
towards the apices, simple to irregularly branched, marginal rhizines numerous, mostly simple, becoming
irregularly branched with age, 0.3–1.0 mm long, ± projecting beyond the lobe margins. Apothecia and pycnidia not
seen.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-
acetoxyhopane-16β,22-diol (trace), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (trace or absent), norstictic
acid (major), connorstictic acid (trace).
Distribution and habitat:—Rare, known only from one collection on rock in a montane forest, also in
Australia. While the only other collection originates from a dry coastal sclerophyll forest in Australia, the Thai
specimen was collected at 1745 m above sea level; new to SE Asia and the Northern Hemisphere.
Remarks:—This lichen has a pale lower surface and marginal rhizines and marginal and laminal, phyllidiate
isidia that erupt to form dense, coralloid pseudoisidia; the medulla contains norstictic acid, 16β-acetoxyhopane-
6α,22-diol and zeorin as major metabolites.
Material from Thailand examined:—Phitsanulok: Phu Hin Rongkla National Park, along the trail around the
Phu Man Khaow, in montane forest, on rock, 1745 m, 16°54'23'' N, 101°05'38'' E, 31 March 2004, T. Deansa RU-
TD 2, (RAMK 21647).
5. Heterodermia diademata (Taylor) D.D. Awasthi, Geophytology 3(1): 113 (1973) (Fig. 7A)
Parmelia diademata Taylor, London J. Bot. 6: 165 (1847). Type:—NEPAL. Wallich (holotype FH-Tayl., fide Kurokawa 1962);
Physcia hypoleuca var. diademata (Taylor) Müll. Arg., Flora 63: 277 (1880); Physcia speciosa f. diademata (Taylor) Müll. Arg.,
Flora 71: 197 (1888); Pseudophyscia hypoleuca f. diademata (Taylor) Hue, Nuov. Arch. Mus. Hist. Nat., sér. 4, 1: 113 (1899);
Anaptychia diademata (Taylor) Kurok., Beih. Nova Hedwigia 6: 28 (1962); Physcia major Nyl., Flora 41: 379 (1858).
Type:—MEXICO. Orizaba: F. Müller (holotype H-Nyl!); Heterodermia major (Nyl.) Trevis., Atti Soc. Ital. Sci. Nat. 11: 618
(1869 [1868]); Physcia speciosa var. major (Nyl.) Müll. Arg., Flora 70: 60 (1887); Anaptychia major (Nyl.) Vain., Dansk. Bot.
Ark. 4(11): 12 (1926).—For further synonymy, see Kurokawa (1962, 1998) and Swinscow & Krog (1976).
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, to 15 cm wide. Lobes 0.5–2.5 mm
wide, plane to weakly convex or weakly concave, sublinear-elongate, sympodially to irregularly branched,
contiguous or discrete at the periphery, radiating; apices not ascending, with short lateral lobes, eciliate; isidia,
phyllidia and soralia absent. Upper surface greenish white, whitish or gray, pruinose or not, rarely with marginal
lobules. Medulla white. Lower surface corticate, whitish to pale brown to dirty brown towards the centre, smooth
to strongly corrugated. Lower surface rhizines and marginal rhizines comparatively sparse, the latter sometimes
projecting from the thallus, concolorous with the thallus or darkening at the apices, simple or irregularly or
squarrosely branched. Apothecia laminal, sessile to substipitate, 1.5–5.5 mm wide; margin entire, crenulate to
lobulate; disc concave, brown to brown-black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 25−40 ×
10−17 µm. Pycnidia common, immersed or slightly protruding; conidia bacilliform, 3–4 × 1 µm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-acetoxyhopane-16β,22-diol (minor),
6α,16β-diacetoxyhopane-22-ol (minor or trace).
Distribution and habitat:—Grows on rock and more often on the bark of trees (Cassia, Betula and
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Lagerstroemia are reported) in hill evergreen forest, plantations, dry evergreen forest, lower montane forest, lower
montane scrub, coniferous forest and dry dipterocarp forest from 700–2000 m above sea level, also in Australia,
North, Central and South America, Africa and Asia; new for Thailand.
Remarks:—characterized by the narrow, sublinear-elongate lobes, the lack of isidia, phyllidia and soralia and
the presence of 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor) and 6α-acetoxyhopane-16β,22-diol
(minor) in the medulla.
Material from Thailand examined:—Nan: Doi Phu Kha, area of view point at Doi Phu Kha, in hill evergreen
forest, on bark of Cassia bakeriana, 1470 m, 19°18'13'' N, 101°09'55'' E, 14 July 1995, S. Meesim RU-DK 10,
(RAMK 21384); ibid., S. Meesim RU-DK 30.1, (RAMK 21669); Chiang Mai: Doi Suthep-Pui, along the trail from
Research centre Suan Song Saeng Kasetsart Uiversity, National guard units 1 Pha Dum, in evergreen forest, on
bark of unidentified tree, 1415 m, 18°45'04'' N, 98°53'16'' E, 30 April 2005, W. Hongsachart RU-WD 58, (RAMK
21524); ibid., W. Hongsachart RU-WD 73, (RAMK 21713); Doi Inthanon National Park, area along trail Doi
Inthanon, in hill evergreen forest, on bark of unidentified tree, 2000 m, 18°35'22'' N, 98°29'15'' E, 4 March 1998, P.
Mongkolsuk 11813, (RAMK 13724); ibid., area along trail Siriphum waterfall, in hill evergreen forest, on bark of
unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9 September 2005, N. Juntawong RU-NJ 285, (RAMK
21496); Loei: Na Haeo National Park, area of Wat Pha Chaleam Phrakeid, in dry evergreen forest, on rock, 750 m,
17°30'15'' N, 100°56'25'' E, 14 July 1995, P. Mongkolsuk 5684, (RAMK 21885); ibid., area of Wat Pha Chaleam
Phrakeid, in dry evergreen forest, on bark of unidentified tree, 750 m, 17°30'15'' N, 100°56'25'' E, 14 July 1995, P.
Mongkolsuk 5656, (RAMK 21884); ibid., area of check point 1, in hill evergreen forest, on Castanopsis sp., 1260
m, 17°30'15'' N, 100°56'25'' E, 12 July 1995, P. Mongkolsuk 4318, (RAMK 21879); Loei: Phuluang Wildlife
Sanctuary, area Phu khon of Forest protection unit, in dry dipterocarp forest, on rock, 700 m, 17°14'33'' N,
101°18'28'' E, 21 May 2009, S. Meesim & K. Buaruang, RU-MSPL 535, (RAMK 21115); Loei: Phuluang Wildlife
Sanctuary, area Lon Tae of Forest protection unit, along trail to Pa Kob, in coniferous forest, on bark of
unidentified tree, 1250 m, 17°12'00'' N, 101°33'48'' E, 12 November 2009, S. Meesim & K. Buaruang, RU-MSPL
603, (RAMK 21126); Loei: Phuluang Wildlife Sanctuary, area Kok Nok Kraba, in lower montane scrub, on bark of
unidentified tree, 1490 m, 17°16'49'' N, 101°31'29'' E, 28 August 2008, S. Meesim & K. Buaruang, RU-MS 32.1,
(RAMK 21031); Phitsanulok: Phu Hin Rong Kla National Park, along trail to Lan Hin Taek area, in lower montane
scrub, on rock, 925 m, 17°00'27'' N, 101°59'36'' E, 4 February 2003, C. Chansuk, RU-CS 6, (RAMK 21665); ibid.,
along trail to Phu Man Khaow area, in lower montane forest, on bark of unidentified tree, 1745 m, 16°54'23'' N,
101°05'38'' E, 31 March 2004, T. Deansa, RU-MS 35, (RAMK 21484); Kamphaeng Phet: Maewong, along trail
Chong Yen, in hill evergreen forest, on Betula alnoides, 1340 m, 16°06'03'' N, 99°06'27'' E, 2 May 2006, S. Meesim
RU-MW 9, (RAMK 21485); Nakon Ratchasima, Khao Yai National Park, Royal Forest Department training
centre, in plantation, on Lagerstroemia speciosa, 825 m, 14°24'50'' N, 101°22'28'' E, 21 September 2001, W.
Polyiam RU-22282, (RAMK 7144); Phetchaburi: Kaeng Krachan, along trail Khao Phanoenthung before km 36, in
hill evergreen forest, on bark of unidentified tree, 1210 m, 12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim
RU-MS 119, (RAMK 21483).
6. Heterodermia dissecta (Kurok.) D.D. Awasthi, Geophytology 3(1): 113 (1973) (Fig. 7B)
Anaptychia dissecta Kurok., J. Jap. Bot. 34(6): 182 (1959). Type:—JAPAN. Mikawa: Mt. Horaiji, 7 January 1956, Kurokawa 56029
(holotype TNS, fide Kurokawa 1959; isotype H!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–10 cm wide. Lobes 0.7–2.0 mm wide, plane to
weakly convex or weakly concave, sublinear-elongate, sympodially to subdigitately branched; apices not
ascending, ± discrete to contiguous at the periphery, with short lateral lobes, eciliate. Upper surface gray-white to
gray, epruinose, with marginal phyllidiate isidia or dorsiventral lobules that are ± granular; marginal phyllidia
usually minutely dissected, often granular near the tips, or entire phyllidia becoming granular and appearing
sorediate. Medulla white. Lower surface with a pseudocortex, white (sometimes rose-red spotted if norstictic acid
is decomposing) to grayish or pale brown at the centre, smooth, rarely slightly corrugated. Lower surface rhizines
rare or even absent, whitish to cream, concolorous with the thallus or becoming pale to dark brown towards the
apices, simple to irregularly branched, marginal rhizines numerous, mostly simple, becoming irregularly branched
and black with age, 0.3–1.5 mm long, ± projecting beyond the lobe margins. Apothecia very rare, laminal, sessile
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to substipitate, 1–5 mm wide; margin lobulate-isidiate; disc concave, brown to brownblack, epruinose. Ascospores
Pachysporaria-type, ellipsoidal, 28–32 × 12–16 μm. Pycnidia rare, initially immersed, becoming emergent, visible
as black dots; conidia bacilliform, 4–5 × 1 μm.
FIGURE 7. A. Heterodermia diademata (scale 1 cm). B. Heterodermia dissecta (scale 5 mm). C. Heterodermia firmula (scale 5 mm).
D. Heterodermia granulifera (scale 1.5 mm). E. Heterodermia isidiophora (scale 5 mm). F. Heterodermia isidiophorella (scale 2 mm).
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), 6α-acetoxyhopane-16β,22-
diol (trace), leucotylin (minor), dissectic acid (major to minor), norstictic acid (major), connorstictic acid (major to
trace).
Distribution and habitat:—Occurs on rocks and on the bark of unidentified trees in evergreen mountain
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
forest, lower montane scrub, tropical rainforest and lower montane forest from 765–1595 m above sea level, also in
Australia, Réunion, India, China and Japan; new addition to the Thai lichen biota.
Remarks:—This lichen is characterized by the minutely dissected marginal lobules and/or phyllidia that often
appear granular sorediate and the presence of norstictic and dissectic acids.
Material from Thailand examined:—Chiang Mai: Lumphun, Mae On, descent from Doi Mon Larn to Mae
Kam Pong village, in evergreen mountain forest, on rock over mosses, 1500 m, 18°51'22'' N, 99°22'02'' E, 19
March 2008, K. Kalb & al. (hb. KK 37038); Loei: Phuluang Wildlife Sanctuary, area of Kok Nok Kraba, along trail
to channel 7, in lower montane scrub, on bark of unidentified tree, 1470 m, 17°16'53'' N, 101°31'23'' E, 13
November 2008, S. Meesim & K. Buaruang, RU-MS 0275, (RAMK 20972); Phitsanulok, Phu Hin Rong Kla
National Park, area of Political and military schools, in tropical rainforest, on bark of Castanopsis sp., 1165 m,
16°59'33'' N, 100°00'42'' E, 5 June 2003, P. Mongkolsuk RU-PM 140, (RAMK 21411); ibid., on rock, 1165 m,
16°59'33'' N, 100°00'42'' E, 5 June 2003, P. Mongkolsuk RU-PM 136, (RAMK 21767); ibid., along trail to the Tub
Boek area, in lower montane forest, on bark of unidentified tree, 1595 m, 16°55'33'' N, 100°05'58'' E, 1 April 2004,
P. Mongkolsuk RU-PM 262/1, (RAMK 21815); ibid., P. Mongkolsuk RU-PM 262, (RAMK 21814); Nakon
Ratchasima: Khao Yai National Park, 'Pha Klua Mai' nature trail to Heo Suwat waterfall, in tropical rainforest, bark
of unidentified tree, 760 m, 14°26'19'' N, 101°24'25'' E, 30 October 1997, T. Pooprang RU-9140, (RAMK 5550).
7. Heterodermia firmula (Nyl.) Trevis., Atti Soc. Ital. Sci. Nat. 11: 615 (1868) (Fig. 7C)
Physcia firmula Nyl., Syn. Lich. 1: 418 (1860). Type:—INDIA. Supra terram in Himalaya, 1220–1520 m, J. D. Hooker & Thomson
2017 (holotype H-Nyl, fide Kurokawa 1962); Physcia speciosa var. firmula (Nyl.) Linds., Trans. Roy. Soc. Edinburgh 22: 248
(1859); Anaptychia firmula (Nyl.) Dodge et Awasthi, in Awasthi, J. Ind. Bot. Soc. 39: 423 (1960).
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, 3–7 cm wide. Lobes 0.3–1.2 mm wide,
plane to weakly convex or weakly concave, sublinear-elongate, sympodially to irregularly branched, contiguous or
discrete at the periphery, radiating; apices not ascending, eciliate, lacking isidia, phyllidia and soralia, with distinct
snow-white pseudocyphellae along the margins of the lobes (best seen if the thallus is somewhat darkened). Upper
surface grayish white to grayish brown, not pruinose. Medulla white, patchily or entirely orange-yellow to orange-
brown. Lower surface corticate, whitish to pale brown to dirty brown towards the centre, smooth to slightly
corrugated. Lower surface rhizines and marginal rhizines numerous, the latter sometimes projecting from the
thallus, concolorous with the thallus or darkening at the apices or totally black, simple or irregularly or squarrosely
branched. Apothecia rare, laminal, sessile to substipitate, 0.5–2.2 mm wide; margin entire, crenulate to lobulate
with punctiform pseudocyphellae; disc concave, brown to brown-black, epruinose. Ascospores Pachysporaria-
type, ellipsoidal, 20−27 × 10−11 µm. Pycnidia rare, immersed or slightly protruding; conidia bacilliform, 3–4 × 1
μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ purple, C–, P–; containing atranorin (major),
zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-acetoxyhopane-16β,22-diol (minor),
7-chloroemodin (major), flavoobscurins (minor).
Distribution and habitat:—Grows on the bark of trees (Acer pseudoplatanus, Rhododendron, Alnus
nepalensis, Pinus) and more rarely on rocks in hill evergreen forest and lower montane scrub forest, also in Nepal,
India and southwestern China from 1260 m in Thailand to 2500 m in Nepal; new addition to the Thai lichen biota.
Remarks:—characterized by the narrow, sublinear-elongate lobes with distinct pseudocyphellae along the
margins, the lack of isidia, phyllidia and soralia and the presence of an orange-yellow pigment in the medulla.
Heterodermia diademata is very similar, but differs in lacking the pigment in the medulla and in having smaller
ascospores.
Material from Thailand examined:—Chiang Mai: Queen Sirikit Botanic Garden, on south facing slope
opposite the entrance, in hill evergreen forest, on rock, 1260 m, 18°53'46'' N, 98°51'37'' E, 12 July 1995, P.
Mongkolsuk 4221, (RAMK 21873); Loei: Phuluang Wildlife Sanctuary, near office of forest protection unit Lon
Tae substation, on Acer pseudoplatanus in lower montane shrub forest, 1260 m, 17°14'18'' N, 101°33'19'' E, 29
December 2009, S. Meesim RU-MSPL 660 (RAMK).
Further selected specimens examined: INDIA. United Province: Dehradun; Rajpur, on Mangifera indica by
the road, ca. 1000 m, 26 February 1964, G. Degelius As-82 (GZU)—NEPAL. Ostnepal, Vorhimalaya: Höhe östlich
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über Panepa, an Steilflächen harten Gesteins, August 1962, J. Poelt (GZU); Kathmandu valley, NW of Kathmandu,
Rani-Ban-Forest, dry Rhododendron forest, 1600–1800 m, 17 March 1979, U. Kirschbaum (GZU).
Heterodermia granulifera (Ach.) W.L. Culb., Bryologist 69: 482 (1967) (Fig. 7D)
Parmelia granulifera Ach., Syn. Meth. Lich.: 212 (1814). Type:—NORTH AMERICA. Muhlenberg (holotype H!); Anaptychia
granulifera (Ach.) A. Massal., Mem. Lich.: 41 (1853); Physcia speciosa var. granulifera Tuck., Proc. Amer. Acad. Arts and Sci.
4: 390 (1860); Squamaria granulifera (Ach.) A. Massal., Atti Ist. Veneto Sci. 5: 250 (1860); Physcia speciosa var. hypoleuca f.
isidiophora Nyl., Syn. Meth. Lich. 1(2): 417 (1860); Heterodermia hypoleuca f. isidiophora Trevis., Atti Soc. Ital. Sci. Nat. 11:
618 (1869 [1868]); Physcia granulifera (Ach.) Tuck., Syn. North Amer. lich. 1: 69 (1882); Anaptychia speciosa f. isidiophora
(Nyl.) Zahlbr., Bot. Mag. (Tokyo) 41: 364 (1927).
Thallus foliose, orbicular to irregularly spreading, loosely adnate at the margins, 2–10 cm wide. Lobes 0.3–1.2 mm
wide, ± plane, or slightly concave, short, sympodially or irregularly branched, ± discrete to contiguous at the
periphery, isidiate; apices not ascending, eciliate. Upper surface whitish gray, usually pruinose; isidia mostly
laminal, more rarely marginal, papillar, granular, rarely long cylindrical, forming dense, coralloid aggregates that
cover most of the upper surface. Medulla white. Lower surface corticate, white to cream to lead-gray or pale tan
near the centre. Lower surface rhizines rare, whitish to cream, concolorous with the thallus or becoming pale to
dark brown towards the apices, simple to irregularly branched, marginal rhizines numerous, whitish to cream,
mostly simple, becoming irregularly branched and black with age, 0.3–1.0 mm long, ± projecting beyond the lobe
margins. Apothecia rare, laminal, subsessile, 0.4–2.0 wide, margin minutely crenate, pruinose and isidiate; disc
dark brown, epruinose; Ascospores Pachysporaria-type, pale brown, ellipsoid, somewhat constricted at the centre,
20–25 ×10–13 μm; pycnidia common, immersed in thallus, visible as black wart-like dots, conidia bacilliform, 4–5
× 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), salazinic
acid (major), consalazinic acid (major), ± hybocarpone (minor).
Nomenclature notes:—Zahlbruckner (1927), Awasthi (1973), Kashiwadani et al. (1990), Elix (2011c),
Weerakoon & Aptroot (2013) have erroneously used Physcia speciosa (Wulfen) Nyl. f. isidiophora Nyl. as the
basionym of Anaptychia/Heterodermia isidiophora although Nylander (1860: 417) stated under Physcia speciosa
var. hypoleuca: Parmelia granulifera Ach. Syn. p. 212 est forma isidiophora. Consequently, the combinations
Anaptychia speciosa (Wulfen) Nyl. f. isidiophora (Nyl.) Zahlbr. and Heterodermia isidiophora (Nyl.) D.D.
Awasthi are not synonymous with Heterodermia isidiophora (Vain.) D.D. Awasthi.
Distribution and habitat:—Known from North and Central America, including Mexico and Guatemala.
Remarks:—This lichen has a pale lower surface and marginal rhizines and mostly laminal, isidia that form
dense, coralloid clusters; the medulla contains salazinic acid and 16β-acetoxyhopane-6α,22-diol as major
metabolites, hybocarpone is sometimes found on the lower surface.
Specimens examined:—U.S.A. West Virginia: near Cedarville, Gilmer Co., on Quercus alba, May 1956, M.
Hale (Lichenes Americani Exsiccati 51, M!).—MEXICO. Chiapas: Ca. 25 km NW of Comitan de Dominguez, in a
shady, dry Quercus-Pinus forest, 2150 m, 25 January 1979, K. Kalb & G. Plöbst (hb. Kalb
40530).—GUATEMALA. Chimaltenango: ca. 5 km SE of Patzún, in a light, dry Cupressus forest, 2000 m, 17
January 1979, K. Kalb & G. Plöbst (hb. Kalb 40304).
8. Heterodermia isidiophora (Vain.) Swinscow & Krog, comb. nov. (1976) ut (Vainio) Awasthi (Fig. 7E).
Mycobank MB 814604
Basionym: Anaptychia isidiophora Vain., sp. nov. (1901) ut (Nyl.) Wain.
Physcia domingensis f. isidiophora Nyl., Acta Soc. Sci. Fennicae 7: 440 (1863). Type:—COLOMBIA. Bogota, ad saxa locis
subumbrosis, Lindig 2534 (holotype H); Anaptychia isidiophora (Nyl.) Vain., Cat. Afric. Plants 2(2): 409 (1901). Not Physcia
speciosa var. hypoleuca f. isidiophora Nyl. (1860: 417); see remarks under H. granulifera.
Thallus foliose, orbicular to irregularly spreading, loosely adnate, forming extensive colonies to 20 cm wide. Lobes
0.7–2.5 mm wide, ± plane, sublinear-elongate, sympodially or irregularly branched, ± discrete to contiguous at the
periphery, with short lateral lobes; apices not ascending, eciliate. Upper surface whitish gray, esorediate, isidiate;
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
isidia marginal and ± laminal, cylindrical or occasionally somewhat flattened, simple to coralloid-branched.
Medulla white. Lower surface corticate, white to grayish to pale brown near the centre. Rhizines numerous,
concolorous with the thallus or becoming dark to brown-black towards the apices, irregularly branched, mainly
marginal, forming a dense black mat. Apothecia rare, laminal, adnate to substipitate 1.5–5.0 mm wide; margin
isidiate; disc concave, brown to brown-black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 21–31 ×
12–17 μm. Pycnidia rare, immersed; conidia bacilliform, 4–6 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), 6α-acetoxyhopane- 16β,22-diol (minor or trace), leucotylin
(minor), 6α,16β-diacetoxyhopane-22-ol (minor or trace).
Nomenclature notes:—Vainio (1901: 409) did not refer to Physcia speciosa f. isidiophora Nyl. as basionym,
Syn. Meth. Lich. 1(2): 417 (1860) as indicated by Kurokawa (1962) and some subsequent authors (see remark
under P. granulifera), but to Physcia domingensis f. isidiophora Nyl., Acta Soc. Sci. Fennicae 7: 440 (1863), and
he explicitly described the differences between Anaptychia isidiophora and A. granulifera. Nevertheless there
exists some ambiguity about the basionym, because it was not listed directly below the new combination as in all of
his other new combinations, but after the description. However, when describing new species in this paper, Vainio
always added sp.n. after the new name and this is not the case with Anaptychia isidiophora. Furthermore, Vainio
published the name as Anaptychia isidiophora (Nyl.) Vain. and thus indicated that he was making a new
combination and not describing the new species Anaptychia isidiophora Vain .
Distribution and habitat:—Occurs on bark of unidentified trees and, more rarely, on rocks in hill evergreen
forest, mixed deciduous forest and lower montane forest from 880–1595 m above sea level; also in Australia,
Central and South America, Africa, Asia and New Zealand.
Remarks:—This species is characterized by the laminal and marginal, simple to coralloid-branched isidia, the
corticate lower surface with concolorous to brown-black rhizines and the lack of depsidones.
Material from Thailand examined:—Chiang Mai: Doi Inthanon National Park, along trail to Siriphum
waterfall, in hill evergreen forest, on bark of unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9 September
2005, N. Juntawong RU-NJ 273, (RAMK 21587); Loei: Phuluang Wildlife Sanctuary, Phuluang Wildlife Research
Station, area of Maenam Sarn, in mixed deciduous forest, on rock, 880 m, 17°20'36'' N, 1ß1°30'17'' E, 11
November 2009, S. Meesim & K. Buaruang, RU-MS 0289, (RAMK 20976); ibid., S. Meesim & K. Buaruang, RU-
MS 0290, (RAMK 20977); ibid., S. Meesim & K. Buaruang, RU-MS 0093, (RAMK 20923); Phitsanulok: Phu Hin
Rong Kla National Park, along trail area the Tub Boek, in lower montane forest, on bark of unidentified tree,
1595 m, 16°55'33'' N, 101°05'58'' E, 1 April 2004, T. Diagsa, RU-TD 17/1, (RAMK 21598); ibid., P. Mongkolsuk
RU-PM 261, (RAMK 21588); Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on bark
of unidentified tree, 1340 m, 16°06'03.42'' N, 99°06'26.95'' E, 2 May 2006, S. Meesim RU-MW 5, (RAMK 21849).
Heterodermia isidiophorella Elix, Australas. Lichenol. 69: 13 (2011b) (Fig. 7F)
Type: AUSTRALIA. Queensland; Cook district, Mt. Windsor Tableland, 45 km NW of Mossman, 1200 m, 16°15' S, 145°01' E, on
Flindersia in stunted open rainforest, J. A. Elix 16465 (holotype CANB!).
Thallus small-foliose, orbicular to irregularly spreading, adnate, 2–5 cm wide. Lobes 0.15–0.5 mm wide, ± plane,
sublinear-elongate, sympodially or irregularly branched, ± discrete to contiguous at the periphery, with short lateral
lobes; apices slightly ascending and recurved, showing the sorediate-granular lower surface, eciliate. Upper surface
whitish gray, phyllidiate; phyllidia marginal, flat, simple to sparingly branched, eventually becoming granular and
appearing sorediate-isidiate-granular. Medulla white. Lower surface corticate, becoming ecorticate at the periphery
or only with corticated strands, white to pale tan or brown near the centre. Marginal rhizines and lower surface
rhizines white, concolorous with the thallus or becoming pale to dark brown towards the apices, simple to
irregularly branched, numerous, 0.5–1.0 mm long, ± projecting beyond the lobe margins. Apothecia not seen.
Pycnidia immersed; conidia bacilliform, 4–6 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 6α-acetoxyhopane-16β,22-diol (major), leucotylin (minor or trace), 16β-acetoxyhopane-6α,22-diol
(trace), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (trace).
Distribution and habitat:—This rare species occurs on bark, twigs and, rarely, on rocks in Australia where it
was considered to be endemic. However, recently we found it in Borneo (K.K.) and the Azores (F.S.). It was
reported by v.d. Boom et al. (2011) from Réunion.
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Remarks:—Heterodermia isidiophorella is characterized by the diminutive thallus with very narrow lobes,
which are partly peripherally recurved showing the ecorticate and sorediate lower surface with white to brown
rhizines and having simple to sparingly branched phyllidia that become granular with age, the partly peripherally
recurved lobes showing the ecorticate and sorediate lower surface with white to brown rhizines. H. isidiophorella is
very similar to H. koyana, but differs mainly by the lack of dissectic acid and some minor morphological
properties, e.g. smaller thalli and smaller lobes. Contrary to the statement regarding presence of isidia in the taxon
(Elix 2011b), these vegetative propagules are generally absent but phyllidia are present, differing from isidia by the
lack of a lower cortex.
Specimens examined: MALAYSIA. Borneo: Sabah; Crocker Ranges, Kota Kinabalu National Park, ca. 50 km
NE of Kota Kinabalu, in a tropical montane rainforest, 06°09' N, 116°39' E, 1600 m, 2 August 2014, K. Kalb & A.
Mertens (hb. Kalb 40473).—PORTUGAL. Azores: Terceira, junction EN3-1 (from Angra do Heroismo to
Biscoitos) to Veredas, Cryptomeria japonica forest, 38°41.738' N, 27°14.224' W, 400 m, 11 July 2008, F. Schumm
(herb. Schumm 14038).
9. Heterodermia koyana (Kurok.) Elix, Australas. Lichenol. 66: 61 (2010a) (Fig. 8A)
Anaptychia dissecta Kurok. var. koyana Kurok., J. Jap. Bot. 34(6): 183 (1959); Type: JAPAN. Kii: Mt. Koya, 4 April 1956, Kurokawa
56027 (holotype TNS, fide Kurokawa 1959); Heterodermia dissecta Kurok. var. koyana (Kurok.) D.D. Awasthi, Geophytology 3:
113 (1973).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–10 cm wide. Lobes 0.7–2.0 mm wide, plane to
weakly convex or weakly concave, sublinear-elongate, sympodially to subdigitately branched; apices not
ascending, ± discrete to contiguous at the periphery, with short lateral lobes, eciliate. Upper surface gray-white to
gray, epruinose, with marginal phyllidiate isidia or dorsiventral lobules, rarely becoming granular; marginal lobules
occasionally minutely dissected, rarely granular near the tips, or the entire lobule becoming granular and appearing
sorediate. Medulla white. Lower surface with a pseudocortex, white to grayish or pale brown in the centre. Lower
surface rhizines absent or very rare, marginal rhizines numerous and partly projecting beyond the lobe margins,
irregularly branched, concolorous with the thallus or darkening and forming a dense black marginal mat. Apothecia
very rare, laminal, sessile to substipitate, 1–5 mm wide; margin lobulate-phyllidiate; disc concave, brown to
brown-black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 28–32 × 12–16 μm. Pycnidia initially
immersed, then becoming emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), 6α-acetoxyhopane- 16β,22-diol (trace), leucotylin (minor),
dissectic acid (minor).
Distribution and habitat:—Occurs on rocks and bark of unidentified trees in mixed deciduous forest, lower
montane scrub and hill evergreen forest from 880–1470 m above sea level; also in Australia, Central America?,
India and Japan; new addition to the Thai lichen biota.
Remarks:—characterized by the minutely dissected marginal lobules and/or phyllidia, the pseudocorticate
lower surface, the presence of dissectic acid and the absence of norstictic acid. Heterodermia dissecta is
morphologically very similar, but it contains additionally norstictic acid and more commonly develops granular-
sorediate phyllidiate isidia.
Material from Thailand examined:—Loei: Phuluang Wildlife Sanctuary, Phuluang Wildlife Research
Station, area of Maenam Sarn, in mixed deciduous forest, on rock, 880 m, 17°20'36'' N, 1ß1°30'17'' E, 11
November 2009, S. Meesim & K. Buaruang, RU-MS 0294, (RAMK 20980); Loei: Phuluang Wildlife Sanctuary,
area of Kok Nok Kraba along trail to channel 7, in lower montane scrub, on bark of unidentified tree, 1470 m,
17°16'46'' N, 101°31'08'' E, 30 August 2005, W. Hongsachart WD 207, (RAMK 21899); Loei: Phuluang Wildlife
Sanctuary, area of Kok Nok Kraba along trail to Pha Som Ded and Pha Taloen, in lower montane scrub, on bark of
unidentified tree, 1470 m, 17°16'46'' N, 101°31'08'' E, 24 June 2008, S. Meesim & K. Buaruang, RU-MS 14,
(RAMK 20998); Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on bark of
unidentified tree, 1340 m, 16°06'04'' N, 99°06'27'' E, 2 May 2006, S. Meesim, RU-MW 18, (RAMK 21825).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
FIGURE 8. A. Heterodermia koyana (scale 5 mm). B. Heterodermia koyanoides (scale 5 mm). C. Heterodermia lepidota (scale 5
mm). D. Heterodermia pseudospeciosa (scale 5 mm). E. Heterodermia rubescens (scale 5 mm). F. Heterodermia rugulosa (scale
5 mm).
10. Heterodermia koyanoides Elix, Australas. Lichenol. 69: 14 (2011b) (Fig. 8B)
Type:—AUSTRALIA. Queensland: Mount Baldy, 4 km SW of Atherton, 17°16’ S, 145°23’ E, 1080 m, on sapling along margin of
regrowth rainforest, 25 June 1984, J.A. Elix 16308 & H. Streimann (holotype, CANB!).
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Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–10 cm wide. Lobes 0.7–2.0 mm wide, plane to
weakly convex or weakly concave, sublinear-elongate, sympodially to subdigitately branched, ± discrete to
contiguous at the periphery, with short lateral lobes; apices not ascending, eciliate. Upper surface whitish gray,
sorediate; soralia marginal and at the apices of the lateral lobes, capitate, 0.5–1.2 mm wide; soredia granular.
Medulla white. Lower surface with a pseudocortex, white to grayish or pale brown in the centre. Lower surface
rhizines absent or very rare, marginal rhizines numerous, predominantly marginal, concolorous with the thallus, ±
irregularly branched, 0.3–1.0 mm long, ± projecting beyond the lobe margin. Apothecia and pycnidia not seen.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P+ yellow; containing atranorin
(major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-acetoxyhopane-16β,22-diol
(trace), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (trace or absent), dissectic acid (major or minor).
Distribution and habitat:—This very rare, corticolous lichen was known only from the type locality in a
montane forest in north-eastern Queensland, but was now collected also from bark of trees (reported are Osbeckia
stellata and Betula alnoides) in hill evergreen forest and lower montane forest in Thailand from 1215–1420 m
above sea level; new addition to the Thai lichen biota and the Northern Hemisphere.
Remarks:—H. koyanoides is characterized by the pseudocorticate lower surface, the capitate soralia at the
lobe margins and the apices of short lateral lobes, the absence of dissected marginal phyllidia and the presence of
dissectic acid and 16β-acetoxyhopane-6α,22-diol in the medulla.
Material from Thailand examined:—Chiang Mai: Doi Suthep-Pui, along trail junction to Pan Ma-Doi-Pui,
km 5, in hill evergreen forest, 1420 m, 18°48'17'' N, 98°53'46'' E, 30 April 2005, W. Hongsachart RU-WD 38,
(RAMK 21862); Phitsanulok: Phu Hin Rong Kla National Park, along trail from km 28 to Nakronthai 36 along trail
from Paradron waterfall, in lower montane forest, on Osbeckia stellata, 1215 m, 16°59'56'' N, 101°00'50'' E, 2
April 2004, S. Meesim RU-SM 49, (RAMK 21855); Phitsanulok: Phu Hin Rong Kla National Park, along trail area
the Tub Boek, in lower montane forest, on bark of unidentified tree, 1595 m, 16°55'33'' N, 101°05'58'' E, 1 April
2004, T. Diagsa, RU-TD 14, (RAMK 21810); ibid., T. Diagsa, RU-TD 17, (RAMK 21819); Kamphaeng Phet:
Maewong, along trail Chong Yen, on Betula alnoides in hill evergreen forest, 1340 m, 16°06'03.42'' N,
99°06'26.95'' E, 2 May 2006, S. Meesim 2, (RAMK 21826).
Heterodermia lepidota Swinscow & Krog, Lichenologist 8: 122 (1976) (Fig. 8C)
Type:—UGANDA. Karamoja District: Mathenico County, Mt. Moroto, near Sogolomon, 2°30' N, 34°45' E, 2500 m, 12 June 1970,
Swinscow 2U 36/12 (holotype BM!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, forming extensive colonies to 20 cm wide. Lobes
0.7–2.5 mm wide, ± plane to convex, sublinear, subdichotomously to irregularly branched, ± discrete to contiguous
at the periphery, with short lateral lobes; apices not ascending, eciliate. Upper surface whitish gray, phyllidiate,
with sparse pseudocyphellae along the margins which are best seen from below; phyllidia mostly marginal, very
rarely also ± laminal, simple to dissected, ecorticate on the lower surface. Medulla white. Lower surface corticate,
white to grayish to pale brown near the centre. Lower surface rhizines sparse, marginal rhizines concolorous with
the thallus or becoming dark brown to brown-black towards the apices, simple to sparingly branched and projecting
beyond the lobes. Apothecia rare, laminal, adnate to substipitate 0.5–2.0 mm wide; margin crenulate to phyllidiate;
disc concave, brown to brown-black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 24–33 × 12–17 μm.
Pycnidia immersed, mainly in the phyllidia; conidia bacilliform, 4–6 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), 6α-acetoxyhopane-16β, 22-diol (minor or trace), leucotylin
(minor), 6α,16β-diacetoxyhopane-22-ol (minor or trace).
Distribution and habitat:—Rare on bark of deciduous trees in open situations in Australia, East Africa (from
1850–3000 m above sea level), the Galapagos Islands and Venezuela.
Remarks:—characterized by the simple to dissected phyllidia, and the presence of only triterpenes in the
medulla.
Specimens examined: KENYA. Central Province: Nanyuki district, between Naru Moru and Mount Kenya,
2300 m, 14 August 1985, K. Kalb & A. Schrögl (herb. Kalb 13578); Rift Valley: Kajiado district, Ngong hills, ca.
35 km SW of Nairobi, 1850 m, 7 August 1985, K. Kalb & A. Schrögl (herb. Kalb 13820).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
11. Heterodermia pseudospeciosa (Kurok.) W.L. Culb., Bryologist 69: 484 (1967 [1966]) (Fig. 8D)
Anaptychia pseudospeciosa Kurok., J. Jap. Bot. 34(6): 176 (1959). Type:—JAPAN. Honshu: Prov. Idzu, Suishochi, Amagi Pass, 20
August 1956, Asahina (holotype TNS, isotype M!).
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, 2–5 cm wide. Lobes comparatively
small, 0.7–1.5 mm wide, plane to weakly convex, sublinear-elongate, sympodially to subdigitately branched,
usually discrete at the periphery, radiating; apices not ascending, eciliate, with distinct pseudocyphellae along the
margins of the lobes or on small short lateral branches. Upper surface grayish white to gray to brownish gray, ±
darker at the lobe tips, pruinose or not, sorediate; soralia white to bluish gray, originating mostly from bursting
pseudocyphellae, developing into subcapitate or labriform soralia. Medulla white. Lower surface corticate, whitish
to pale brown, rarely dark gray in the centre. Lower surface rhizines sparse, mostly simple or sparsely branched,
marginal rhizines frequent, concolorous with the thallus or becoming dark or black near the apices, irregularly
branched, to 1 mm long, projecting beyond the lobes. Apothecia rare, laminal, adnate to sessile, 1–3 mm wide;
margin initially crenulate, later sorediate; disc concave, brown to brown-black, epruinose. Ascospores
Pachysporaria-type, ellipsoidal, 26–32 × 12–14 μm. Pycnidia immersed or slightly protruding; conidia
bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-
acetoxyhopane-16β,22-diol (trace), norstictic acid (major), connorstictic acid (minor), salazinic acid (trace).
Distribution and habitat:—This species occurs on bark of trees (Magnolia henryi, Dipterocarpus costatus,
Castanopsis and Acer pseudoplantanus are reported) and, more rarely, on mossy rocks in hill evergreen forest, dry
dipterocarp forest, lower montane scrub, tropical rainforest, lower montane rainforest and mixed deciduous forest
from 700–1610 m above sea level; also in North, Central and South America, Africa, Asia, Australia and New
Zealand.
Remarks:—This lichen is characterized by the bursting pseudocyphellae along the lobe margins developing
into soralia and the presence of norstictic acid in the medulla. Usually the development of the soralia starts in a
very early stage and it may be difficult to see the few remaining pseudocyphellae. But sometimes the development
of the soralia starts late and those specimens are very similar to Heterodermia angustiloba, which however, is
easily separated by the presence of dissectic acid.
Material from Thailand examined:—Nan: Doi Phu Kha, along trail to Doi Phu Kha, in hill evergreen forest,
on bark of Magnolia henryi, 1595 m, 19°18'13'' N, 101°09'55'' E, 10 May 2006, S. Meesim RU-DK 13, (RAMK
21821); ibid., S. Meesim RU-DK 6, (RAMK 21822); Chiang Mai: Doi Suthep-Pui, along trail junction to Pan Ma-
Doi-Pui, km 5, in hill evergreen forest, on bark of Dipterocarpus costatus, 1420 m, 18°48'17'' N, 98°53'46'' E, 30
April 2005, W. Hongsachart RU-WD 39, (RAMK 21841); ibid., W. Hongsachart RU-WD 39, (RAMK 21863);
Chiang Mai: Doi Inthanon National Park, area along trail Siriphum waterfall, in hill evergreen forest, on bark of
unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9 September 2005, N. Juntawong RU-NJ 443, (RAMK
21827); Loei: Na Haeo National Park, area of check point 1, in hill evergreen forest, on Castanopsis sp., 1260 m,
17°30'15'' N, 100°56'25'' E, 12 July 1995, P. Mongkolsuk 4462, (RAMK 21875); Loei: Na Haeo National Park, area
of War Pha Chaleam Phrakeid, in dry dipterocarp forest, bark of unidentified tree,705 m, 17°30'15'' N, 100°56'25''
E, 5 November 1995, P. Mongkolsuk 6637, (RAMK 21872); Loei: Phuluang Wildlife Sanctuary, area of Kok Nok
Kraba, in lower montane scrub, on rock, 1495 m, 17°16'49'' N, 101°31'29'' E, 29 August 2005, W. Hongsachart,
RU-WD 148, (RAMK 21179); ibid., W. Hongsachart, RU-WD 122, (RAMK 21164); ibid., S. Meesim & K.
Buaruang RU-MS 85, (RAMK 21089); Loei: Phuluang Wildlife Sanctuary, area of Kok Nok Kraba along trail to
channel 7, in lower montane scrub, on rock, 1470 m, 17°16'46'' N, 101°31'08'' E, 29 August 2005, W. Hongsachart
WD 221, (RAMK 21900); Loei: Phuluang Wildlife Sanctuary, office of forest protection unit, Phu Ho, in dry
dipterocarp forest, on bark of Acer pseudoplatanus, 700 m, 17°14' 33'' N, 101°18'28'' E, 10 November 2009, S.
Meeesim RU-MSPL 653, (RAMK 21147); Phitsanulok: Phu Hin Rong Kla National Park, area of political and
military schools, in tropical rainforest, on rock, 1165 m, 16°59'33'' N, 100°00'42'' E, 5 June 2003, P. mongkolsuk,
RU-PM 135, (RAMK 21700); Phitsanulok: Phu Hin Rong Kla National Park, area of Kang Han Nam water fall, in
lower montane rain forest, on rock, 1165 m, 16°59'31'' N, 100°00'44'' E, 6 June 2003, P. Mongkolsuk RU-PM
special , (RAMK 21817); Phitsanulok: Phu Hin Rong Kla, forest protection unit Man Daeng, in lower montane
forest, on rock, 1610 m, 16°55'51'' N, 101°03'04' E, 7 June 2004, P. Mongkolsuk RU-PM 256, (RAMK 21832);
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ibid., P. Mongkolsuk RU-PM 256/2, (RAMK 21833); Phitsanulok: Phu Hin Rong Kla National Park, along trail to
Lan Hin Taek area, in mixed deciduous forest, on rock, 825 m, 17°00'27'' N, 100°59'36'' E, 6 February 2003, C.
Chansuk RU-CS 19, (RAMK 21816).
12. Heterodermia rubescens (Räsänen) D.D. Awasthi, Geophytology 3(1): 114 (1973) (Fig. 8E)
Anaptychia hypoleuca f. rubescens Räsänen, Arch. Soc. Zool. Bot. Fenn. 'Vanamo' 5(1): 27 (1950). Type:—INDIA. Darjeeling: E.
Himalayas, 1830 m, June 1948, D.D. Awasthi 161 (holotype H!); Anaptychia esorediata f. rubescens (Räsänen) D.D. Awasthi, J.
Indian Bot. Soc. 39(3): 423 (1960); Anaptychia rubescens (Räsänen) Kurokawa, Beih. Nova Hedwigia 6: 31 (1962).
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, to 15 cm wide. Lobes 1.0–1.5 mm
wide, plane to weakly convex or weakly concave, sublinear-elongate, sympodially to irregularly branched,
contiguous or discrete at the periphery, radiating; apices not ascending, without short lateral lobes, eciliate; isidia,
phyllidia and soralia absent. Upper surface whitish gray or gray, not pruinose. Medulla white. Lower surface
corticate, whitish to pale brown to dirty brown towards the centre, smooth to weakly corrugated. Lower surface
rhizines and marginal rhizines comparatively sparse, the latter sometimes projecting from the thallus, concolorous
with the thallus or darkening at the apices, simple or irregularly or squarrosely branched. Apothecia laminal, sessile
to substipitate, 1.0–5.0 mm wide; margin entire, crenulate to lobulate; disc concave, brown to brown-black,
epruinose. Ascospores Pachysporaria-type, ellipsoidal, 23−30 × 11−14 μm. Pycnidia rare, immersed or slightly
protruding; conidia bacilliform, 3–4 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow; containing
atranorin (major), zeorin (major), norstictic acid (major), connorstictic acid (submajor) and 16β-acetoxyhopane-
6α,22-diol (major).
Distribution and habitat:—Grows on bark of trees (Betula alnoides and Quercus incana are reported) in
India and Thailand in lower montane forest and hill evergreen forest from 1470–1745 m above sea level; new
addition to the Thai lichen biota.
Remarks:—characterized by the narrow, sublinear-elongate lobes, the lack of isidia, phyllidia and soralia and
the presence of norstictic acid (major), connorstictic acid (submajor) and 16β-acetoxyhopane-6α,22-diol (minor) in
the medulla. Kurokawa (1962) also found salazinic acid, but this might have been confused with connorstictic acid.
H. diademata is very similar in overall habit, but is readily separated by the lack of norstictic acid.
Material from Thailand examined:—Nan: Doi Phu Kha, area of view point at Doi Phu Kha, in hill evergreen
forest, on bark of Betula alnoides, 1470 m, 19°18'13'' N, 101°09'55'' E, 10 May 2006, S. Meesim RU-DK 4,
(RAMK 21476); ibid., S. Meesim RU-DK 11, (RAMK 21386); Phitsanulok: Phu Hin Rongkla National Park, along
the trail around the Phu Man Khaow, in lower montane forest, on bark of Betula alnoides, 1745 m, 16°54'23'' N,
101°05'38'' E, 2 April 2004, T. Deansa RU-TD 53, (RAMK 21451).
Heterodermia rugulosa (Kurok.) Wetmore, Bryologist 79: 304 (1976) (Fig. 8F)
Anaptychia rugulosa Kurok., Beih. Nova Hedwigia, 6: 41 (1962). Type:—MEXICO. Jalisco: Guadalajara, on Bursera trees, 29 July
1902, Pringle 15372 (holotype US, isotype TNS; fide Kurokawa 1962).
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, to 10 cm wide. Lobes 0.7–2 mm wide,
plane to weakly convex or weakly concave, sublinear-elongate, dichotomously to irregularly branched, contiguous
or discrete at the periphery, radiating; apices not ascending, eciliate; isidia, phyllidia and soralia absent. Upper
surface greenish white, densely white pruinose, centrally with short lateral flat or knobby lobules. Medulla white,
patchily yellow. Lower surface corticate, whitish to pale brown to dirty brown towards the centre, smooth to
strongly corrugated. Lower surface rhizines and marginal rhizines numerous, the latter projecting from the thallus,
concolorous with the thallus or darkening at the apices, simple or irregularly or squarrosely branched. Apothecia
laminal, sessile to substipitate, 1.5–6 mm wide; margin entire, crenulate to lobulate; disc concave, brown to brown-
black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 19−28 × 11−13 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), leucotylin (minor), yellow pigment (7-chloroemodin).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Distribution and habitat:—Grows on the bark of trees in North America and Mexico.
Remarks:—characterized by the narrow, sublinear-elongate lobes, the lack of isidia, phyllidia and soralia and
the presence of 7-chloroemodin in the medulla. Heterodermia diademata is very similar, but differs in lacking the
yellow pigment in the medulla.
13. Heterodermia speciosa (Wulfen) Trevis., Atti Soc. Ital. Sci. Nat. 11: 614 (1868) (Fig. 9A).
Lichen speciosus Wulfen, in Jacquin, Coll. Bot. 3: 119 (1789); Type:—AUSTRIA. Carinthia: In sylvarum Montis Calvariae saxis (not
seen); Anaptychia speciosa (Wulfen) A. Massal.; Mem. Lich.: 36 (1853); Physcia speciosa (Wulfen) Nyl., Actes. Soc. Linn.
Bordeaux 21: 307 (1856); Pseudophyscia speciosa (Wulfen) Müll. Arg., Bull. Herb. Boissier 2, app. 1: 40 (1894). Pseudophyscia
speciosa (Wulfen) Müll. Arg., Bull. Soc. Roy. Bot. Belgique 32: 129 (1893), invalidly published (Art. 35.1., Art. 38.1., Art. 38.5.;
no description provided for the generic name and the genus was not monotypic).
Thallus foliose, orbicular to irregular, adnate to loosely adnate, 2–4 cm wide, often coalescing to form colonies up
to 15 cm wide. Lobes 0.5–1.5 mm wide, plane to weakly convex, sublinear-elongate, often slightly broader towards
the apices, dichotomously branched, ± discrete at the periphery, radiating; apices not ascending, eciliate, with
distinct pseudocyphellae along the margins of the lobes or on small short lateral branches. Upper surface grayish
white to gray to brownish gray, ± darker at the lobe tips, pruinose or not, sorediate; soralia white to bluish gray,
originating mostly from bursting pseudocyphellae, developing into subcapitate or labriform soralia. Medulla white.
Lower surface corticate, whitish to pale brown, rarely dark gray in the centre. Marginal rhizines frequent,
concolorous with the thallus or becoming dark or black near the apices, irregularly branched, to 1 mm long,
projecting beyond the lobes, lower surface rhizines sparse, mostly simple or sparsely branched. Apothecia rare,
laminal, adnate to sessile, 1–3 mm wide; margin initially crenulate, later sorediate; disc concave, brown to brown-
black, epruinose. Apothecia rare, laminal, sessile to substipitate, 3–8 mm wide; margin crenulate at first,
subsequently sorediate; disc concave, brown to brown-black, epruinose. Ascospores Pachysporaria-type,
ellipsoidal, 25–37 × 14–18 μm. Pycnidia immersed or slightly protruding; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing
atranorin (major), zeorin (major), 6α-acetoxyhopane-16β,22-diol (minor), 6α-acetoxy-22-hydroxyhopane-25-oic
acid (minor or trace), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (minor), leucotylin (trace).
Distribution and habitat:—Scattered in open situations over mosses on rocks and trees in hill evergreen
forest and lower montane scrub from 1200–1320 m above sea level; also in Australia, Europe, North and South
America, Africa, Asia, New Zealand.
Remarks:—This lichen is characterized by the striking contrast between the blue-gray soralia and the paler
thallus. Actually it is most similar to Heterodermia pseudospecia, but lacks norstictic acid in the medulla.
European specimens of this species reveal a yellowish pigment on TLC plates with Rf values 36, 21, 21 in solvents
A, B' and C. This spot is not seen from tropical material tested. The taxonomic importance of this finding can only
be solved by molecular methods. Nevertheless, we consider it rather unlikely that a species which is primarily
restricted to mountainous areas in Europe (Moberg 2004b), should also occur in the tropics. Similarly, Kurokawa
(1962) doubted the occurrence of H. speciosa outside of Europe and transferred specimens from the tropics to H.
pseudospeciosa, but this species contains norstictic acid.
Material from Thailand examined:—Chiang Mai: Doi Inthanon National Park, along trail to Siriphum
waterfall, in hill evergreen forest, on bark of unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9 September
2005, N. Juntawong RU-NJ 444, (RAMK 21860); ibid., N. Juntawong RU-NJ 329, (RAMK 21859); ibid., N.
Juntawong RU-NJ 466, (RAMK 21861); Phitsanulok: Phu Hin Rong Kla National Park, along trail to Lan Hin Pum
area, in lower montane scrub, on rock, 1200 m, 16°59'11'' N, 100°59'28'' E, 30 March 2004, S. Meesim RU-MS 30,
(RAMK 21672).
Further specimens examined: AUSTRIA. Upper Austria: mountain range called "Totes Gebirge", Dietlgut
near Hinterstoder, ± 650 m, 2 October 1984, R. Türk; Plantae Graecenses, Lich. 389 (GZU)—SWITZERLAND.
Valais: Pennine Alps above Zermatt, Sunnegga, over mosses on Serpentinit, 2200 m, 2 May 1974, K. Kalb (herb.
Kalb 2334).
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FIGURE 9. A. Heterodermia speciosa (scale 5 mm). B. Heterodermia tremulans (scale 5 mm). C. Heterodermia tropica (scale
5 mm). D. Heterodermia verdonii (scale 5 mm). E. Leucodermia borphyllidiata (scale 5 mm). F. Leucodermia borphyllidiata, lateral
phyllidia, seen from below (scale 1 mm).
Heterodermia tremulans (Müll. Arg.) W.L. Culb., Bryologist 69: 485 (1967) (Fig. 9B)
Physcia hypoleuca var. tremulans Müll. Arg., Flora 63: 277 (1880); Anaptychia speciosa var. tremulans (Müll. Arg.) Müll. Arg., Bot.
Jahrb. 15: 505 (1893); Pseudophyscia speciosa var. tremulans (Müll. Arg.) Müll. Arg., Bull. Soc. Roy. Bot. Belgique 32: 130
(1893), invalidly published (Art. 35.1., Art. 38.1., Art. 38.5.; no description provided for the generic name and the genus was not
monotypic). Type:—BRAZIL. Rio de Janeiro: near Petropolis on mosses, Deventer (holotype G, fide Kurokawa 1962).—For
further synonymy, see Kurokawa (1962).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Thallus foliose, orbicular to irregularly spreading, adnate to loosely adnate, 2–7 cm wide, often coalescing and
forming colonies to 15 cm wide. Lobes 0.7–1.5 mm wide, plane to slightly convex, sublinear, ± flexuose,
sympodially to subdigitately branched, usually discrete at the periphery, radiating; apices not ascending, with short
lateral lobes, eciliate. Upper surface grayish white to gray to brownish gray, ± darker at the apices, pruinose or not,
sorediate; soralia whitish or grayish, granular, forming labriform soralia at the tips of the main and lateral lobes.
Medulla white. Lower surface corticate, whitish to pale brown, rarely dark gray in the centre. Lower surface
rhizines sparse, mostly simple or sparsely branched, marginal rhizines frequent, concolorous with the thallus or
becoming dark or black near the apices, irregularly branched, to 1 mm long, projecting beyond the lobes.
Apothecia rare, laminal, sessile to substipitate, 1–3 mm wide; margin initially crenulate, becoming sorediate; disc
concave, brown to brown-black, epruinose. Ascospores Pachysporaria-type, ellipsoidal, 23–32 × 12–14 μm.
Pycnidia immersed or slightly protruding; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing
atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), 6α-acetoxyhopane-16β,22-diol (major),
6α,16β-diacetoxyhopane-22-ol (minor), leucotylin (minor), chloroatranorin (minor).
Distribution and habitat:—This pantropical and pantemperate species occurs on bark, dead wood and rocks,
but is not yet reported from Thailand. It might, however occur in the country and not distinguished from H.
speciosa.
Remarks:—Although this lichen has commonly been confused with P. speciosa, the latter has linear-elongate
lobes and larger spores (25–37 × 14–18 μm), whereas P. tremulans has short, often flexuose lobes and smaller
spores (23–32 × 12–14 μm). The specimen from São Paulo, cited below, was collected not far from the type
locality at ± the same elevation above sea level.
Specimens examined: BRAZIL. Rio de Janeiro: Serra da Mantiqueira; Itatiaia, Parque National do Itatiaia,
near the Maromba waterfall, in a humid and dark primary rainforest, 1050 m, 22°20’ S, 44°35’ W, 21 July 1978,
(herb. Kalb 39806); Rio de Janeiro: Serra da Mantiqueira; between Registro do Picú and Agulhas Negras, in a
humid swamp with small bushes, 1850 m, 22°20’ S, 44°40’ W, 14 March 1980, K. Kalb & G. Plöbst (herb. Kalb
39807); São Paulo: Serra do Mar; between Areias and Silveiras, near Cruzeiro, 700 m, 22°40’ S, 44°45’ W, 4
November 1978, K. Kalb & G. Plöbst (herb. Kalb 40531).
Heterodermia tropica (Kurok.) Sipman, Tropical Bryology 12: 206 (1996) (Fig. 9C)
Anaptychia tropica Kurok., Beih. Nova Hedwigia 6: 36 (1962). Type:—MEXICO. Chiapas: Road to El Suspiro, 5–7 km N of
Berriozábal, 920 m, Hale 20214 (holotype US, fide Kurokawa 1962); Pseudophyscia speciosa mexicana de Lesd., Lichens du
Mexique: 4 (1914). Type:—Mexico. Puebla: Acatzingo, 2100 m, July 1907, Amable 4107 (lectotype [selected here] UPS!);
Heterodermia tropica (Kurok.) Sipman, Tropical Bryology 12: 206 (1996); Heterodermia tropica (Kurok.) Kurok., Folia
Cryptogamica Estonica 32: 24 (1998), illegimate (see remarks).
Thallus foliose, orbicular to irregularly spreading, comparatively small, adnate, to 5 cm wide. Lobes to 3 mm long,
0.5–1.0 mm wide, to ca. 1.5–2.5 mm wide at the tips, ± plane to weakly convex, sublinear-elongate, usually richly
subdichotomously branched; lobe apices not ascending, eciliate, lacking isidia, phyllidia and soralia, with
pseudocyphellae along the margins of the lobes, best seen from below. Upper surface whitish gray to brownish
gray, darker at the apices, occasionally sparsely pruinose. Medulla white. Lower surface corticate, whitish, lead-
gray to pale brown, blackish towards the centre, smooth to corrugated. Lower surface rhizines frequent, evenly
distributed on the lower surface, simple or irregularly branched, usually short (ca. 1 mm long), pale to dark brown
or black, marginal rhizines frequent, simple or irregularly branched, concolorous with the thallus, darkening near
the apices or black. Apothecia common, laminal, substipitate, 0.5–4.0 mm wide; disc concave, dark brown,
epruinose. Ascospores Pachysporaria-type, narrowly ellipsoidal, 20–25 × 10–13 μm. Pycnidia ± common,
immersed, with black weakly protruding apices; conidia bacilliform, 4–6 × 1 µm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (minor), leucotylin (minor), 6α-
acetoxyhopane-16β,22-diol (trace), salazinic acid (major), hypoconstictic acid (submajor), 3-O-methylconsalazinic
acid (minor), consalazinic acid (minor).
Nomenclature notes:—Two more combinations are cited in MycoBank,
http://www.mycobank.org/BioloMICS.aspx?Table=Mycobank&Rec=186653&Fields=All
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Heterodermia tropica (Kurok.) Trass, Censo de Macrolíquenes Venezolanos de los Estados Falcón, Lara,
Mérida, Táchira y Trujillo: 180 (1986) and Heterodermia tropica (Kurok.) Sipman, Tropical Bryology 6: 24
(1992). But Heterodermia tropica (Kurok.) Trass is not mentioned in the reference given. On p. 180 only
Heterodermia tropica (Kurok.) is cited without any further author and without a basionym. Furthermore the author
of the book is not Trass but Lopez Figueiras.—Heterodermia tropica (Kurok.) Sipman is not mentioned in Tropical
Bryology 6: 24 (1992) or on any other page in this article.—The first correct combination is made by Sipman in
Tropical Bryology 12: 206 (1996), thus making the later combination of Kurokawa (1998) illegitimate.
Distribution and habitat:—This ± rare species occurs on bark in Central and South America (Venezuela) in
(sub-)montane rainforests between 1000 and 2500 m.
Remarks:—Heterodermia tropica is characterized by the lack of vegetative propagules and the presence of
salazinic acid and hypoconstictic acids in the medulla. On a world basis, three Heterodermia species are known
with this chemistry, namely H. tropica without vegetative propagules, H. albicans with soralia and H. antillarum
with isidia.
Specimens examined:—MEXICO. Chiapas: ca. 5 km W of San Cristobal de las Casas, on deciduous trees
alongside a street, 1800 m, 19 January 1979, K. Kalb & G. Plöbst (herb. Kalb 40302)—GUATEMALA.
Chimaltenango: between Zaragoza and Patzùn, 2165 m, 14°37'51'' N, 90°55'12'' W, K. Kalb & G. Plöbst (herb.
Kalb 40485).
14. Heterodermia verdonii Elix, Australas. Lichenol. 68: 18 (2011) (Fig. 9D)
Type:—AUSTRALIA. New South Wales: 25 km NW of Coffs Harbour, below Waihou trig. stn., 30°06’S, 153°02’E, 340 m, on
sandstone outcrop in closed Tr is tania-Ficus forest, 12 Oct. 1978, D. Verdon 3814 (CANB!);—Chaudhuria indica Zahlbr., Ann.
Mycol. 30 (5–6): 434 (1932). Type:—INDIA. Darjeeling: supra muscos, 2286 m, G. L. Chopra (holotype W!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, to 5 cm wide, but often forming colonies to 10
cm wide. Lobes 0.5–1.2 mm wide, plane to weakly convex or weakly concave, sublinear-elongate, sympodially
branched, radiating; apices plane, usually discrete, eciliate; phyllidia, isidia and soredia absent, but with distinct
pseudocyphellae along the margins of the lobes. Upper surface greenish white, whitish to gray. Medulla white.
Lower surface corticate, whitish to pale brown, dirty brown near the centre, smooth. Rhizines mainly marginal,
concolorous with the thallus or darkening near the apices, irregularly branched, 1–2 mm long. Apothecia laminal,
substipitate, 0.5–2.5 mm wide; margin ± crenate with pseudocyphellae; disc concave, brown to blackish brown, ±
thinly gray-pruinose. Ascospores Pachysporaria-type, ellipsoidal, 20–30 × 13–15 μm. Pycnidia common, initially
immersed, becoming emergent, visible as black dots; pynoconidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex and medulla K+ yellow, C–, KC–, P+ yellow; containing atranorin (major), zeorin (major),
dissectic acid (submajor), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-acetoxyhopane-16β,22-
diol (major), 6α,16β-diacetoxyhopane-22-ol (minor).
Nomenclature notes:—Zahlbruckner (1932) described a new monotypic genus for this species, because he
was puzzled by the bluish-green discoloration of the thallus of the specimen available to him and thus presuming it
contained a cyanobacterium as the photobiont. But the study of the type specimen revealed green algae as
photobiont and a typical Heterodermia chemistry, i.e. atranorin (major), zeorin (major), dissectic acid (major), 16β-
acetoxyhopane-6α,22-diol (major), leucotylin (minor), japonene (trace) and 6α,16β-diacetoxyhopane-22-ol (trace).
Kurokawa (1962) who did not see the type, placed Chauduria indica into synonymy with Heterodermia
diademata. Indeed that species is superficially very similar to Heterodermia verdonii, but is easily separated by a
different terpene array and especially by the lack of dissectic acid. Heterodermia angustiloba is also similar
morphologically, but contains additional norstictic acid.
However, Chaudhuria indica, the older name cannot be combined into Heterodermia because another species
has been described as Heterodermia indica (H. Magn.) D.D. Awasthi.
Distribution and habitat:—This rare lichen occurs on bark (Betula alnoides is reported) and rock in hill
evergreen forest, lower montane forest and dry evergreen forest from 800–1610 m above sea level; also in India
and Australia.
Remarks:characterized by the narrow, eciliate lobes, the lack of phyllidia, isidia and soredia, and the
occurrence of dissectic acid and triterpenes.
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Material from Thailand examined:—Nan: Doi Phu Kha, area of view point at Doi Phu Kha, in hill evergreen
forest, on bark of Betula alnoides, 1470 m, 19°18'13'' N, 101°09'55'' E, 10 May 2006, S. Meesim RU-DK 15,
(RAMK 21382); ibid., S. Meesim RU-DK 7, (RAMK 21477); Chiang Mai: Doi Inthanon National Park, along trail
to Siriphum waterfall, in hill evergreen forest, on bark of unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9
September 2005, N. Juntawong RU-NJ 464, (RAMK 21689); ibid., Doi Suthep-Pui, along trail from research
centre of Suan Song Saeng Kasetsart University to National guard units 1 Pha Dum, in hill evergreen forest, on
bark of unidentified tree, 1415 m, 18°45'04'' N, 98°53'16'' E, 30 April 2005, W. Hongsachart RU-WD 57, (RAMK
21710); Phitsanulok: Phu Hin Rongkla, forest protection unit Man Daeng, in lower montane forest, on rock, 1610
m, 16°55'51'' N, 101°03'04'' E, 16 July 2004, P. Mongkolsuk RU-PM 256/4, (RAMK 21812); Nakon Ratchasima:
Khao Yai National Park, around Ban Chom View, in dry evergreen forest, on bark of unidentified tree, 800 m,
14°25'19'' N, 101°23'51'' E, 11 January 2003, S. Udompaisan RU-21567, (RAMK 7172).
II. The lichen genus Leucodermia
Leucodermia Kalb, gen. nov.
Mycobank MB 813825
Synonym: Anaptychia Körb. sect. Polyblastidium Kurok. ser. leucomelaenae Kurok., Beih. Nova Hedwigia 6: 8 (1962). Type
species:—Leucodermia leucomelos (L.) Kalb; Lichen leucomelos L., Sp. Pl., ed. 2, 2: 1613 (1763). America meridionali (LINN
1273.109, holotype n.v., fide Jørgensen et al. 1994: 329, 378).
Etymology:—The generic name has been combined from ser. leucomelaenae and Heterodermia.
The new genus is characterized by a loosely adnate or, in part, unattached thallus with ribbon-like, linear elongate,
dichotomously branched lobes with conspicuous long marginal rhizines, a paraplectenchymatous upper cortex and
a lower ecorticated surface. Apothecia stipitate with triangular lobules, disc brown, ± densely whitish to gray
pruinose. Ascospores brown, of Polyblastidia-type, with sporoblastidia.
Remarks:Leucodermia is closely related to Heterodermia comosa-group, but differs mainly in having linear
elongate, dichotomously branched lobes and a foliose to subfruticose thallus. In contrast, the lobes in H. comosa-
group are usually ascending, spathulate or paddle-shaped.
Key to Thai species of the genus Leucodermia
(TLC at least in solvent A is necessary for identification)
1. Medulla with salazinic acid............................................................................................................... L. leucomelos (L.) Kalb
- Medulla without depsidones .................................................................................................................................................. 2
2. Thallus with squamiform phyllidia ................................................................................... L. borphyllidiata Kalb & Meesim
- Thallus without vegetative propagules..................................................................................................... L. boryi (Fée) Kalb
1. Leucodermia borphyllidiata Kalb & Meesim, sp. nov. (Figs. 9E, 9F).
Mycobank MB 813836
Similar to Leucodermia boryi, but differs in having lobes with marginal phyllidia.
Type:—BRAZIL. Minas Gerais: Serra da Mantiqueira; some km outside Vila Monte Verde, ca. 40 km E of Camanducaia, in a montane
rainforest, 1800 m, 22°45’ S, 46°05’ W, 29 November 1980, K. Kalb 39847 (CGMS, holotype).
Etymology:—The name is composed of bor[yi], indicating the similarity to Leucodermia boryi, and phyllidiata, indicating the
presence of phyllidia in this taxon as they are a very useful diagnostic character.
Thallus foliose to subfruticose, in loose rosettes or forming tangled mats, loosely adnate or, in part, unattached,
5–15 cm wide. Lobes 0.3–0.5 mm wide, separate, ± plane, linear-elongate, ribbon-like, tangled, dichotomously
branched, with circinately revolute tips to the distal lobes, ornated with dorsiventral phyllidia; with conspicuous
gray to black simple or sparingly branched marginal rhizines, 5–15 mm long. Upper surface ivory to gray-white,
smooth. Medulla white. Lower surface mostly ecorticate, canaliculate, powdery, but not sorediate, white
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throughout or partially pinkish brown; pigments absent; lower margins thickened, corticated. Apothecia and
pycnidia not seen
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing
atranorin (major), zeorin (major), japonene (minor or trace).
Distribution and habitat:—Probably a rare species with a pantropical distribution. As far as we can estimate
from the few collections we have, L. borphyllidiata occurs on the bark of trees or over mosses in montane tropical
rainforests up to the timberline.
Remarks:As in other phyllidiate species, in L. borphyllidiata too, the phyllidia readily break off and leave
soralia-like white spots which develop new phyllidia with age. These phyllidia may be very narrow and resemble
isidia.
Specimens examined:—SPAIN. Canary Islands: La Gomera; Schumm 9271; BRAZIL. Minas Gerais: Serra
da Mantiqueira; Vila Monte Verde, ca. 30 km E of Camanducaia, on a Podocarpus tree in a dark and humid
rainforest along a rivulet, 1500 m, 22°50’ S, 46°00’W, 20 August 1978, K. Kalb & G. Plöbst (hb. Kalb
37795);—KENYA. Central Province: Nyeri District, Aberdare National Park, on the ground over mosses between
shrubs, 3300 m, 11 August 1985, K. Kalb & A. Schrögl (hb. Kalb 13918);—CHINA. Xizang: SE Tibet; Tsangpo
Vy., Nangxian-Mainling, Vy. 10 km W of Gyemdong (High Camp 13), undisturbed Betula-Abies forest on N-facing
steep boulder slope, on mosses on bark, 3570 m, 28°56' N, 93°14' E, 5 August 1994, Miehe 94-149-41/01 &
Wündisch (GZU);—THAILAND. Kamphaeng Phet: Maewong, along trail Chong Yen, on an unidentified tree in
an evergreen montane rainforest, 1340 m, 16°06'03.42'' N, 99°06'26.95'' E, 2 May 2006, S. Meesim 18, (RAMK
21721); ibid., S. Meesim 19, (RAMK 21722).
2. Leucodermia boryi (Fée) Kalb, comb. nov. (Fig. 10A)
Mycobank MB 813828
Basionym: Borrera boryi Fée, Essai Cryptog. Écorc. Exot. Officin. Introd. XCVI et tab. II, fig. 23 (1825) and Essai Cryptog. Écorc.
Exot. Officin., Suppl. 148 (1837); Anaptychia boryi (Fée) A. Massal., Mem. Lichenogr. 41 (1853). Type:—ÎLE BOURBON
[Réunion], A.L.A. Fée s.n.; (syntype PC n.v., fide Swinscow & Krog 1976: 124); Anaptychia neoleucomelaena Kurok., J. Jap.
Bot. 36: 51 (1961); Heterodermia neoleucomelaena (Kurok.) Follmann & Redón, Willdenowia 6: 446 (1972). Type:—INDIA.
below Jamunotri Tehri Garhwal, 9000–9500 ft, on tree trunks, 6 June 1951, D.D. Awasthi 902; (holotype LWG-LWU n.v.);
Heterodermia boryi (Fée) K.P. Singh & S.R. Singh, Geophytology 6: 33 (1976).
For further synonymy, see Kurokawa (1998).
Thallus foliose to subfruticose, often in loose rosettes or forming tangled mats, loosely adnate or, in part,
unattached, 5–15 cm wide. Lobes 0.4–3.0 mm wide, separate, ± plane, linear-elongate, ribbon-like, tangled,
dichotomously branched, often ascending at the apices or, occasionally, the apices reflexed or with circinately
revolute tips to the distal lobes, with conspicuous gray to black simple or sparingly branched marginal rhizines,
5–15 mm long. Upper surface ivory to gray-white, smooth. Medulla white. Lower surface mostly ecorticate,
canaliculate, arachnoid or powdery, but not sorediate, white throughout or partially pinkish brown or rarely purple;
yellow pigments absent; lower margins thickened, corticated. Apothecia ± common, subapical to apical,
substipitate to distinctly stipitate, 1–5 mm wide; margin crenulate to lobulate, lobules triangular, to 2 mm long,
often with sparse short black cilia; disc concave, dark brown, ± distinctly white-pruinose. Ascospores
Polyblastidia-type, ellipsoidal, with numerous small and/or large sporoblastidia, 35–53 × 18–25 μm. Pycnidia rare,
immersed, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing
atranorin (major), zeorin (major) and japonene (minor or trace).
Distribution and habitat:—This species occurs on bark and more rarely on mossy rocks in hill evergreen
forest and lower montane forest; it is widespread in tropical to warm temperate regions.
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Remarks:—Although we have not studied the type of this species, we were able to investigate specimens which
were collected recently by the authors in Réunion (type locality). They
agree very well with the clear drawings published in Fée (1825, Tab. 2,
Fig. 23). Heterodermia boryi is distinguished by the tangled mass of
rather narrow, elongate, linear lobes having long black marginal cilia; a
largely ecorticate lower surface and thickened, corticate margins. The
type is fertile and esorediate, but without a molecular investigation it is
not possible to decide whether the sorediate specimens deserve
taxonomic recognition.
Material from Thailand examined:—Chiang Mai: Doi Inthanon National Park, along trail Ang Ka Doi Inthanon,
in hill evergreen forest, on bark of unidentified tree, 2240 m, 18°48'27'' N, 98°53'77'' E, 30 April 2005, W.
Hongsachart, WD 116, (RAMK 21732); Phitsanulok: Phu Hin Rong Kla National Park, along trail area the Tub
Boek, in lower montane forest, on bark of unidentified tree, 1595 m, 16°55'33'' N, 101°05'58'' E, 1 April 2004, T.
Diagsa, RU-TD 23, (RAMK 21730); ibid., S. Meesim RU-MS 40/8, (RAMK 21717).
3. Leucodermia leucomelos (L.) Kalb, comb. nov. (Fig. 10B)
Mycobank MB 813827
Basionym: Lichen leucomelos L., Sp. Pl., 2nd edn, 2: 1613 (1763); Parmelia leucomela (L.) Ach., Methodus 256 (1803); Physcia
leucomelos (L.) Michx., Fl. Bor.-Amer. 2: 326 (1803); Borrera leucomela (L.) Ach., Lichenogr. Univ. 499 (1810); Anaptychia
leucomelaena (L.) A. Massal., Mem. Lichenogr. 35 (1853); Heterodermia leucomelos (L.) Poelt, Nova Hedwigia 9: 31 (1965).
Type:—AMERICA MERIDIONALIS [SOUTH AMERICA]; (LINN 1273.109, holotype n.v., fide Jørgensen et al. 1994: 329,
378).
Remarks:Many synonyms as well as varieties and forms are listed for this species in Kurokawa (1962) and elsewhere. They are not
repeated here, because they may well represent independent species when modern chemical and molecular methods are applied to
their delimitation.
Thallus foliose to subfruticose, often in loose rosettes or forming tangled mats, loosely adnate or, in part,
unattached, 5–15 cm wide. Lobes 0.5–4.0 mm wide, separate, ±plane, linear-elongate, ribbon-like, tangled,
dichotomously branched, often ascending at the apices or, occasionally, the apices reflexed, with conspicuous long,
gray to black simple or squarrosely branched marginal rhizines, 5–15 mm long. Upper surface ivory to gray-white,
smooth, epruinose. Medulla white. Lower surface predominantly ecorticate, but with thickened corticated margins,
canaliculate, arachnoid or powdery and then becoming sorediate, often with subapical soralia, white throughout or
partially pinkish brown or, rarely, purple; yellow pigments absent. Apothecia rare, subapical to apical, substipitate
to distinctly stipitate, 2–5 mm wide; margin crenulate to lobulate, the lobules triangular, to 3 mm long, often with
sparse short black cilia along the margins of the lobules; disc concave, dark brown to black, ± white-pruinose.
Ascospores Polyblastidia-type, ellipsoidal, with numerous small or large sporoblastidia, 31–50 × 15–24 μm.
Pycnidia rare, immersed, visible as black dots; conidia
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), japonene (minor or trace), salazinic acid (minor), consalazinic acid
(trace).
Distribution and habitat:—Occurs on bark and on mossy rocks in tropical rainforest. Widespread in tropical
to warm-temperate regions.
Remarks:Leucodermia leucomelos is characterized by the tangled mass of elongate, linear lobes with long
black marginal rhizines, the ecorticate lower surface with thickened, corticate margins, subapical soralia and the
presence of salazinic acid.—Like in L. boryi, the holotype of L. leucomelos is fertile, but in both species, sorediate
specimens do occur. Whether they deserve taxonomic recognition, remains an unresolved question.
Material from Thailand examined:—Loei: Phuluang Wildlife Sanctuary, Forest protection unit office,
Lontae, along trail Pa Kob, before exit Tropical forest, in tropical rainforest, on bark of unidentified tree, 1254 m,
17°11'32'' N, 101°33'43'' E, 12 November 2009, S. Meesim & K. Buaruang RU-MSPL 581, (RAMK 21123); ibid.,
S. Meesim & K. Buaruang RU-MSPL 706, (RAMK 21155).
MONGKOLSUK ET AL.
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FIGURE 10. A. Leucodermia boryi, fertile specimen (scale 5 mm). B. Leucodermia leucomelos, sorediate specimen (scale 5 mm). C.
Polyblastidium appendiculatum (scale 5 mm). D. Polyblastidium fragilissimum (scale 5 mm). E. Polyblastidium hypocaesium (scale
5 mm). F. Polyblastidium hypoleucum (scale 5 mm).
Material belonging to the following species was also studied and found to pertain to this genus.
Leucodermia appalachensis (Kurok.) Kalb, comb. nov.
Mycobank MB 813829
Phytotaxa 235 (1) © 2015 Magnolia Press
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Basionym: Anaptychia appalachensis Kurok., Beih. Nova Hedwigia 6: 83 (1962). Type:—U.S.A. West Virginia: Hardy Co., Fabius,
on base of chestnut oak, July 1956, Hale 12069 (holotype, US); Heterodermia appalachensis (Kurok.) W.L. Culb., Bryologist 69:
479 (1967 [1966]).
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing atranorin (major),
zeorin (major), 6α-acetoxyhopane-16β,22-diol (submajor), appalachin (major).
Remarks:Although we did not see the type, we did study authentic material from West Virginia and from other
localities in the Appalachian Mountains. Appalachin is a new lichen substance of unknown composition; it is not a
terpene most probably not the yellowish pigment reported for this species. Although we have recognized the latter
in many specimens, we could not detect it on TLC plates after the solvents were evaporated. Appalachin seems to
be a very rare compound in Heterodermia sens.lat., for despite analyzing more than one thousand specimens, we
have only found it in L. appalachensis and one other specimen (RAMK 21767), which we tentatively have
identified as Polyblastidium hypoleucum.
Selected specimens examined:—U.S.A. West Virginia: Pendleton Co.; 6 mi. W of Sugar Grove, base of trees,
mountain sloppes, C.F. Reed 75688 (NY); North Carolina: Swain Co.; Great Smoky Mountains National Park,
Sunkota Ridge, Ridge top forest with hardwoods dominant, on Quercus base, 2200–3400 ft., 35°29'42'' N,
83°24'46'' W, 21 June 2011, J. Lendemer 29427 (NY).
Leucodermia arsenei (Kurok.) Kalb, comb. nov.
Mycobank MB 813830
Basionym: Anaptychia arsenei Kurok., Beih. Nova Hedwigia 6: 89 (1962). Type:—Mexico. Puebla: Esperanza, 2400 m, 1907, Arsène
8089 (holotype US, n.v.). Heterodermia arsenei (Kurok.) Trass, Folia Cryptog. Estonia 29: 7 (1992).
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange; containing atranorin (major),
zeorin (major), salazinic acid (major), consalazinic acid (major), 16β-acetoxyhopane-6α,22-diol (minor).
Remarks:Although we have not seen the type material, we have investigated several collections from Chiapas,
SE of Puebla which matched the description given by Kurokawa (1962). The species was placed in ser.
Podocarpae by Kurokawa (1962), but it seems more closely related to Leucodermia appalachensis. It differs from
L. appalachensis in having broader lobes and by containing salazinic acid.
Specimen examined:—MEXICO. Chiapas: ca. 25 km NW of Comitan de Dominguez, in a dark and dry
forest, dominated by Quercus spec. and Pinus spec., 2150 m, 16°27’ N, 92°19’ W, 25 January 1979, K. Kalb & G.
Plöbst (herb. Kalb 39870, 39872).
Leucodermia circinalis (Zahlbr.) Kalb, comb. nov.
Mycobank MB 813831
Basionym: Anaptychia leucomelaena (L.) A. Massal. var. multifida (Meyen & Flot.) Vainio f. circinalis Zahlbr., Beih. Bot. Centralbl.
19, 2. Abt.: 84 (1905); Type:—ECUADOR. [Cimborazo]: El Altar; Paramo region, 4200 m, Meyer 399. (W, n.v., lectotype,
selected by Kurokawa 1962: 79). Heterodermia circinalis (Zahlbr.) W.A. Weber, Mycotaxon 13: 101 (1981).
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P– or P+ pale yellow; containing atranorin (major),
zeorin (major) and leucotylin (minor to trace).
Remarks:—Zahlbruckner (1905) and Kurokawa (1962) treated this species as a form of Heterodermia leucomelos
or H. boryi. However, it differs from the former by lacking depsidones and from the latter by the more circinately
enrolled tips of the lobe apices. Furthermore, its main distribution is in the paramos of South America, above 3000
m.
Specimen examined:—ECUADOR. Cañar: Side road from the Panamericana to Sangay National Park, in
Paramo vegetation, on mosses and soil on a steep slope along the road, 3100 m, 02°21' S, 78°59,7' W, 31 August
2011, K. Kalb & H. Jonitz (herb. Kalb 40500).
Leucodermia fertilis (Moberg) Kalb, comb. nov.
Mycobank MB 813831
Basionym: Heterodermia fertilis Moberg, Nord. J. Bot. 29: 136 (2011).
Type:—ECUADOR. Azuay, road Cuenca-Moleturo, 25 km west of Cuenca, 2°46' S, 79°12' W, 3600 m, gras paramo and shrub
vegetation, epiphyte. 8 July 1983, L. & A. Arvidsson 4158 holotype GB, isotype UPS).
MONGKOLSUK ET AL.
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Phytotaxa 235 (1) © 2015 Magnolia Press
Remarks:—The clear figure and description provided by Moberg (2011) undoubtedly places this species in
Leucodermia.
Leucodermia lutescens (Kurok.) Kalb, comb. nov.
Mycobank MB 813833
Basionym: Anaptychia lutescens Kurok., J. Jap. Bot. 36(2): 54 (1961). Type:—MEXICO. Chiapas: Monte Ovando, 25 March 1932, E.
Matuda 46 (TNS, holotype, n.v.); Heterodermia lutescens (Kurok.) Follmann, Philippia II(2): 73 (1974).
Chemistry: Atranorin (major), zeorin (major), japonene (major) and an unknown yellow pigment.
Nomenclature notes:—Kurokawa (1962: 79) gave the reference for this species as Journ. Jap. Bot. 36: 55 (1955)
and this was copied by Follmann (loc. cit.) when making the combination into Heterodermia. However, the correct
reference is that given above. Nevertheless the combination made by Follmann is validly published (ICN 41.6.).
Selected specimens examined (both collections are fertile):—BRAZIL. Rio de Janeiro: Serra da Mantiqueira;
Itatiaia, between Registro do Picú and Agulhas Negras, at the edge of a dense rainforest, on Podocarpus, 1900 m,
22°20’ S, 44°45’ W, 1 November 1978, K. Kalb & G. Plöbst (herb. Kalb 39844); BRAZIL. Rio de Janeiro: Serra da
Mantiqueira; Itatiaia, between Registro do Picú and Agulhas Negras, on soil in a humid swamp with small bushes,
1850 m, 22°20’ S, 44°40’ W, 14 March 1980, K. Kalb & G. Plöbst (herb. Kalb 39849).
Leucodermia vulgaris (Vain.) Kalb, comb. nov.
Mycobank MB 813834
Basionym: Anaptychia leucomelaena (L.) A. Massal. var. vulgaris Vain., Acta Soc. Fauna Fl. Fenn. 7(1): 128; Anaptychia vulgaris
(Vain.) Kurok., Beih. Nova Hedwigia 6: 81 (1962). Heterodermia vulgaris (Vain.) Follmann & Redón, Willdenowia 6: 447
(1972). Type:—BRAZIL. Minas Gerais; Lafayette. Vain. Lich. Bras. Exs. 289 (Lectotype, selected by Kurokawa loc. cit.: 81,
TUR-Vain. 7855!).
Remarks:—TLC was performed on the rather large type specimen, as well as our collections made near the type
locality, as well as from Rio de Janeiro and São Paulo States and Peru. All specimens tested exhibit similar
chemistry but japonene may be present (as in the type) or it may be absent.
Chemistry: Atranorin (major), zeorin (major), 7-chloroemodin (submajor), 5,7-dichloroemodin (minor), ±
japonene (minor) and two unknown orange and violet pigments. In solvent A, the two pigments have the same R
f
value, and the common spot appeared violet after the plate was dried, while "vulgaris low" alone appeared pink.
After H
2
SO
4
spray and charring, the common spot in A appeared pink like the "vulgaris low". However after H
2
SO
4
spray and charring, the "vulgaris high" appears very faintly violet or almost invisible in all three solvents, but
"vulgaris low" appears pink in B' and C.
Selected specimens examined:—PERU. Cuzco: surroundings of Machu Picchu, on trees and shrubs alongside
the railway track from Aguas Calientes to Cuzco, 20 October 1995, B. Marbach P 34 (herb. Kalb 40534);
BRAZIL. Minas Gerais: Serra da Mantiqueira; Vila Monte Verde, ca. 30 km E of Camanducaia, on Podocarpus, at
the edge of a dark and humid rainforest along a rivulet, 1500 m, 22°50’ S, 46°00’W, 07–11 September 1978, K.
Kalb & G. Plöbst (herb. Kalb 40532); BRAZIL. Rio de Janeiro: Serra da Mantiqueira; Itatiaia, between Registro do
Picú and Agulhas Negras, on bark of a tree at the edge a dense rainforest, 1900 m, 22°20’ S, 44°45’ W, 1. XI. 1978,
K. Kalb & G. Plöbst (herb. Kalb 40533); BRAZIL. São Paulo: Serra do Mar; Serra de Paranapiacaba, ca 60 km SW
of São Paulo, above Juquitiba, on deciduous trees in a light and humid rainforest along the Rio Juquiá, 700 m,
23°50’ S, 47°10’ W, 13-14 May 1978, K. Kalb & G. Plöbst (herb. Kalb 39841).
III. The lichen genus Polyblastidium
Polyblastidium Kalb, gen. nov.
Mycobank MB 813826
Similar to Heterodermia s.str., but differing in having an arachnoid lower surface and mostly ascospores of Polyblastidia-type. Type
species:—Polyblastidium japonicum (M. Satô) Kalb; Anaptychia dendritica var. japonica M. Satô, J. Jap. Bot. 12: 427 (1936);
Phytotaxa 235 (1) © 2015 Magnolia Press
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Anaptychia japonica (M. Satô) Kurok., Beih. Nova Hedwigia 6: 58 (1962). Type:—TAIWAN. Mt. Arisan, Mingestu, 24 January
1936, M. Satô (Taiwan 10) (holotype TI n.v.); Heterodermia japonica (M. Sato) Swinscow & Krog, Lichenologist 8: 122 (1976).
Etymology:—Among other features, ascospores often with many (Gr. = polys) sporoblastidia distinguish this genus from
Heterodermia s.str.
The new genus is characterized by a thallus with lobes appressed to the substrate which do not or only ascend
slightly towards the tips, a paraplectenchymatous upper cortex, an ecorticate ± compact or arachnoid lower surface
without rhizines (marginal rhizines are present); apothecia which are laminal, brown, ascospores thick-walled, 1-
septate with 1–3 sporoblastidia (Polyblastidia-type), very rarely without sporoblastidia (e.g. P. hypoleuca and P.
microphyllum). Polyblastidium contains many species of Anaptychia Körb. sect. Polyblastidium Kurok. ser.
Polyblastidium Kurok., but the type of this series, Physcia speciosa (Wulfen) Nyl. *obscurata Nyl., has a lower
surface with a superimposed layer of cotton-woolly hyphae, impregnated by yellow to orange-brown pigments,
hence this species together with similar ones belongs in another group, unrelated to Polyblastidium (see below).
Key to Thai species of Polyblastidium
(TLC in solvents A and B is necessary for precise identification)
1. Thallus without vegetative propagules................................................................................................................................... 2
- Thallus with soralia, phyllidia................................................................................................................................................ 3
2. Ascospores without sporoblastidia.............................................................................................. P. hypoleucum (Ach.) Kalb
- Ascospores with 1–3 sporoblastidia................................................................................................ P. togashii (Kurok.) Kalb
3. Thallus with soralia................................................................................................................................................................ 4
- Thallus with phyllidia ............................................................................................................................................................ 7
4. Medulla with salazinic acid as major depsidone, norstictic acid and/or hybocarpone sometimes present as minor to trace
substances............................................................................................................................... P. hypocaesium (Yasuda) Kalb
- Medulla with norstictic acid as major depsidone or without depsidones and hybocarpone.................................................. 5
5. Medulla with norstictic acid as major depsidone.................................................................. P. propaguliferum (Vain.) Kalb
- Medulla without depsidones .................................................................................................................................................. 6
6. Medulla with japonene as major terpene, 6α-acetoxyhopane-16β,22-diol absent......................... P. japonicum (Satô) Kalb
- Medulla without japonene, 6α-acetoxyhopane-16β,22-diol present.............................................. P. chilense (Kurok.) Kalb
7. Medulla with norstictic acid.............................................................................................. P. appendiculatum (Kurok.) Kalb
- Medulla without norstictic acid.............................................................................................................................................. 8
8. Lower surface ± whitish to the centre of the thallus, zeorin present as the only major terpene ..............................................
............................................................................................................................................... P. microphyllum (Kurok.) Kalb
- Lower surface white at the lobe tips, but colored violet-gray to the centre........................................................................... 9
9. Medulla with japonene, 16β-acetoxyhopane-6α,22-diol ± trace or absent........................... P. fragilissimum (Kurok.) Kalb
- Medulla with 16β-acetoxyhopane-6α,22-diol, japonene absent ................................................. P. violostriatum (Elix) Kalb
1. Polyblastidium appendiculatum (Kurok.) Kalb, comb. nov. (Fig. 10C)
Mycobank MB 813838
Basionym: Anaptychia appendiculata Kurok., Beih. Nova Hedwigia 6: 61 (1962). Type:—IVORY COAST, Cercle of Man, Mont
Tonkoui, Rocher aux Sacrifices, on Hymenodictyon floribundum, in the outskirt of a low forest, ca. 1100 m, 14 August 1954, R.
Santesson 10645a (holotype UPS!); Heterodermia appendiculata (Kurok.) Swinscow & Krog, Lichenologist 8: 114 (1976).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–12 cm wide. Lobes 1.0–1.5 mm wide,
broadening to ca. 2–3 mm wide at the tips, plane to weakly convex or weakly concave, sublinear-elongate,
dichotomously to digitately branched, radiating, the apices not ascending, contiguous to discrete, eciliate. Upper
surface greenish white, whitish gray to cream-colored, epruinose, with marginal dorsiventral phyllidia, more rarely
also laminal, usually minutely dissected, often granular near the tips or the whole phyllidia becoming granular and
appearing sorediate. Medulla white. Lower surface ecorticate, ± compact, violetgray in the centre, violet-striate
towards the apices; yellow pigments absent. Rhizines marginal, initially simple and concolorous with the lower
surface, later becoming black and squarrosely branched, 2–4 mm long. Apothecia laminal, adnate to substipitate,
1–4 mm wide; margin lobulate, the lobules becoming elongate and deeply dissected; disc concave, dark brown to
blackish brown, ± thinly gray-pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 0−2 sporoblastidia, 37–53
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× 18–25 μm. Pycnidia common, initially immersed, becoming emergent, visible as black dots; conidia bacilliform,
4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-
acetoxyhopane-16β,22-diol (major), 6α,16β-diacetoxyhopane- 22-ol (minor), chloroatranorin (minor), norstictic
acid (major), connorstictic acid (minor).
Distribution and habitat:—Rare on bark of deciduous trees (Betula alnoides is reported) and on rocks in
lower montane scrub, mixed deciduous forest, tropical rainforest and hill evergreen forest from 790 to 1470 m
above sea level; also in Australia, Central America and the Caribbean, West and East Africa, Malaysia and New
Zealand.
Remarks:—This lichen is characterized by having lobes with an ecorticate, non-pigmented lower surface,
marginal phyllidia, ascospores with sporoblastidia and the presence of terpenes and norstictic acid in the medulla.
Material from Thailand examined:—Loei: Phuluang Wildlife Sanctuary, area of Kok Nok Kraba along trail
to channel 7, in lower montane scrub, on bark of unidentified tree, 1470 m, 17°16'46'' N, 101°31'08'' E, 29 August
2005, W. Hongsachart WD 175, (RAMK 21896); ibid., W. Hongsachart WD 192, (RAMK 21898); area of Phu
Luang Wildlife protection unit at Mae Num Sarn, in mixed deciduous forest, on rock, 790 m, 17°20'36'' N,
101°30'17'' E, 11 November 2008, S. Meesim & K. Buaruang, RU-MS 0098, (RAMK 20929); area of Kok Toei,
Forest protection unit, in mixed deciduous forest, on rock, 1200 m, 17°19'55'' N, 101°31'26'' E, 27 June 2008, S.
Meesim & K. Buaruang, RU-MS 70, (RAMK 21074); Helicopter apron side station, black rest home, in lower
montane scrub, on rock, 1470 m, 17°16'46'' N, 101°31'08'' E, 26 September 2008, S. Meesim & K. Buaruang, RU-
MS 82, (RAMK 21086); Phitsanulok: Phu hin Rong Kla National Park, area of political and military schools, in
tropical rainforest, on rock, 1165 m, 16°59'33'' N, 100°00'42'' E, 5 June 2003, P. Mongkolsuk, RU-PM 139, (RAMK
21351); Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on bark of unidentified tree,
1340 m, 16°06'04'' N, 99°06'27'' E, 2 May 2006, S. Meesim, RU-MW14, (RAMK 21847); ibid., on bark of Betula
alnoides, 1340 m, 16°06'04'' N, 99°06'27'' E, 2 May 2006, S. Meesim, RU-MW14, (RAMK 21848); Nakon
Ratchasima: Khao Yai National Park, tropical rainforest SE of the main road no. 3077 between the park
headquarter and Khao Khaeo, on bark of Ficus sp., 875 m, 14°26'26'' N, 101°21'58'' E, 17 July 2003, W. Polyiam &
N. Osathanon, RU-22407, (RAMK 7646); Phetchaburi: Kaeng Krachan, along trail Khao Phanoenthung, in hill
evergreen forest, on bark of unidentified tree, 1207 m, 12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim,
RU-MW120, (RAMK 21851); ibid., S. Meesim, RU-MW116, (RAMK 21844).
2. Polyblastidium chilense (Kurok.) Kalb, comb. nov.
Mycobank MB 813839
Basionym: Anaptychia chilensis Kurok., Beih. Nova Hedwigia 6: 65 (1962). Type:—CHILE. Valparaiso; Alto del Puerto, on soil at a
small stream in a creek, 14 August 1940, R. Santesson 2919; (S!); Heterodermia chilensis (Kurok.) Swinscow & Krog,
Lichenologist 8: 115 (1976).
Thallus foliose, orbicular to irregularly spreading, adnate, 2–5 cm wide. Lobes 0.7–1.5 mm wide, slightly more
narrow at the apices, plane to convex, sublinear to sublinear-elongate, dichotomously to digitately branched,
ascending at the apices, with a few short lateral branches, eciliate. Upper surface whitish gray to pale yellow-brown,
uneven, pruinose at the apices, sorediate; soralia apical, labriform, originating from the reflexed underside. Medulla
white. Lower surface ecorticate, ± compact or granular towards the tips, white, lacking pigments, canaliculate, with
thick corticate margins. Rhizines marginal, concolorous with the thallus to brown-black, simple to digitately divided,
0.7–2.0 mm long. Apothecia laminal, sessile to substipitate, 1.0–1.5 mm wide; margin pruinose; disc brown-black,
Ascospores Polyblastidia-type, ellipsoidal, with 0–2 small sporoblastidia, 25–39 × 15–22 μm. Pycnidia initially
immersed, later weakly emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), 6α-acetoxyhopane-16β,22-diol (minor), 6α-acetoxy-22- hydroxyhopane-25-oic acid (minor), 6α-
acetoxy-16β,22-dihydroxyhopane-25-oic acid (minor).
Distribution and habitat:—Occurs on mossy rocks, rarely on tree bases in Australia, South America, South
and East Africa and New Zealand and probably also in Thailand.
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Remarks:This lichen is characterized by the saxicolous or terricolous habitat, the apical, white, labriform
soralia, the white, ecorticate lower surface with corticate margins, and by the presence of the speciosa
chemosyndrome of triterpenes, i.e. 6α-acetoxyhopane-16β,22-diol, 6α-acetoxy-22-hydroxyhopane-25-oic acid and
6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid. The only report for Thailand is from the RAMK checklist. We
did not find this species in the material investigated, but as it is possible that it occurs in the country, we have
included it in the key and provided a description from the type specimen.
3. Polyblastidium fragilissimum (Kurok.) Kalb, comb. nov. (Fig. 10D)
Mycobank MB 813840
Basionym: Anaptychia fragilissima Kurok., Beih. Nova Hedwigia 6: 60 (1962). Type:—JAPAN. Prov. Kii: Mt. Koya, Nuymaziri 679;
(TNS n.v., isotype CANB!); Heterodermia fragilissima (Kurok.) Wei & Jiang, Lich. Xizang 111 (1986).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–12 cm wide. Lobes 1.0–1.5 mm wide, ca.
2–3 mm wide at the tips, plane to weakly convex or weakly concave, sublinear-elongate, dichotomously to
digitately branched, radiating; apices not ascending, contiguous to discrete, eciliate. Upper surface greenish white,
whitish gray to cream-colored, epruinose, with marginal dorsiventral phyllidia, very rarely also laminal, usually
minutely dissected, often granular near the tips, or entire phyllidia becoming granular and appearing sorediate.
Medulla white. Lower surface ecorticate, ± compact, violet-gray in the centre, violet-striate towards the apices;
yellow pigments absent. Rhizines marginal, at first simple and concolorous with the lower surface, then black and
squarrosely branched, 2–4 mm long. Apothecia laminal, adnate to substipitate, 1–4 mm wide; margin lobulate, the
lobules becoming elongate and deeply dissected; disc concave, dark brown to blackish brown, ± thinly gray-
pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 2 or 3 sporoblastidia at either ends, 36–50 × 16–20 μm.
Pycnidia common, immersed, then becoming emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P+ pale yellow; containing
atranorin (major), zeorin (major), japonene (major), anaptychin-5 (minor), anaptychin-1 (trace), 6α-acetoxy-
16β,22-dihydroxyhopane-25-oic acid (trace).
Distribution and habitat:—Occurs on trees in Chiang Mai; also on rocks in Australia, Guatemala, Tanzania,
China and Japan.
Remarks:characterized by the minutely dissected marginal phyllidia, the ecorticate lower surface and the
presence of atranorin and triterpenes. This species is very similar to P. violostriatum from which it is distinguished
by the longer, Polyblastidia-type ascospores with more numerous sporoblastidia and in containing the japonica-
rather than the leucotylin-chemosyndrome of triterpenes (Elix 2011a: 20). The North American P. squamulosum is
similar, but the spores are smaller in that species and laminal phyllidia are much more common than in P.
fragilissimum.
Material reported from Thailand:—Chiang Mai: Doi Inthanon National Park (Wolseley & Aguirre-Hudson
(1997: 333).
Specimen examined:—MASCARENE ISLANDS. Réunion: between le Dix Septième and Plain d'Affouches,
ca. 8 km SSW of St-Denis, in remnants of a rainforest, 1000 m, 20°56' S, 55°24'30'' E, 14 August 1991, K. & A.
Kalb (herb. Kalb 33752).
4. Polyblastidium hypocaesium (Yasuda) Kalb, comb. nov. (Fig. 10E)
Mycobank MB 813842
Basionym: Anaptychia hypocaesia Yasuda, In: Räsänen, J. Jap. Bot. 16: 139 (1940). Type:—JAPAN. Prov. Sagami; Hakone, ad cort.
Pini densiflorae, 15 Oct. 1922, Ya s u d a 674 (H!, holotype; H!, isotype); Heterodermia hypocaesia (Yasuda) D.D. Awasthi,
Geophytology 3: 113 (1973).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, to 5 cm wide, but often coalescing to form
colonies up to 15 cm wide. Lobes 0.5–1.2 mm wide, to c. 2–3 (–5) mm wide at the tips, plane to weakly convex or
weakly concave, sublinear-elongate, dichotomously branched, radiating; apices ascending, usually discrete, ±
lobulate along the lobe margins, the lobules ± rounded or sparingly branched, 0.3–0.8 mm wide, 0.5–1.0 mm long.
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Upper surface greenish white, whitish to cream-colored, sorediate; soredia farinose to granular, in labriform to
capitate soralia on lateral or terminal lobes, occasionally spreading along the lobe margins. Medulla white. Lower
surface ecorticate, ± compact, purple-black in the centre, white, ochraceous or yellow towards the apices. Marginal
rhizines common, gray to black, simple or squarrosely branched, 1–3 (–5) mm long. Apothecia very rare, laminal,
substipitate, 1–4 mm wide; margin lobulate, the lobules ecorticate, partly yellow on the lower surface, apically
sorediate; disc concave, dark brown to blackish brown, sometimes very thinly gray-pruinose. Ascospores
Polyblastidia-type, ellipsoidal, with 2 or 3 small sporoblastidia, 35–45 × 16–20 μm. Pycnidia common, initially
immersed, then becoming emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow; pigmented
medulla K– or K+ brownish green or K+ violet; containing atranorin (major), zeorin (major), japonene (major),
salazinic acid (major), ± norstictic acid (minor), hybocarpone (minor), subhypocarpone (trace), 6α-acetoxy-22-
hydroxyhopane-25-oic acid (trace), 7-chloroemodin (minor).
Nomenclature notes:—Kurokawa (1962: 57) cited Y. Asahina as collector. But on the labels of the holo- and
one isotype, A. Yasuda is mentioned as the collector.
Distribution and habitat:—This species occurs on bark of trees (Acacia auriculiformis and Alstonia scholaris
are reported), dead wood and on rock in lower montane scrub, lower montane forest and hill ever green forest from
975 to 1595 m above sea level; also in Australia, India, Nepal, China, Japan and the Hawaiian Islands.
Remarks:characterized by lobes with an ecorticate, pigmented lower surface, labriform to capitate soralia,
ascospores with sporoblastidia and the presence of medullary salazinic acid, hybocarpone and ± 7-chloroemodin in
the lower pigmented medulla. A member of the P. japonicum complex, it is distinguished by the presence of
salazinic acid.
Material from Thailand examined:—Phitsanulok: Phu Hin Rong Kla National Park, along trail to Lan Hin
Taek area, in lower montane scrub, on bark of Acacia auriculiformis, 975 m, 17°00'27'' N, 100°59'36'' E, 4
February 2003, C. Chansuk RU-CS79, (RAMK 21450); ibid., along trail to Tub Boek area, in lower montane
forest, on bark of unidentified tree, 1595 m, 16°55'33'' N, 101°05'58'' E, 1 April 2004, S. Meesim RU-MS39,
(RAMK 21818); Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on bark of
unidentified tree,, 1340 m, 16°06'03'' N, 99°06'27'' E, 2 May 2006, S. MeesimRU-MW 7, (RAMK 21625); Nakon
Ratchasima: Khao Yai National Park, along main road ca. 100–200 m to view point, in hill evergreen forest, on
bark of unidentified tree, 1375 m, 14°21'36'' N, 101°23'47'' E, 23 June 2000, W. Polyiam, RU-20632, (RAMK
5094); ibid., 5 February 2000, W. Polyiam, RU-16732, (RAMK 7128); Phetchaburi: Kaeng Krachan, along trail
from Khao Phanoenthung to the guesthouse area, in hill evergreen forest, on bark of unidentified tree, 1207 m,
12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim RU-MS 96, (RAMK 21622); ibid., on Alstonia scholaris,
S. Meesim RU-MS 98, (RAMK 21407).
5. Polyblastidium hypoleucum (Ach.) Kalb, comb. nov. (Fig. 10F)
Mycobank MB 813843
Basionym: Parmelia speciosa b. hypoleuca Ach., Syn. Lich.: 211 (1814). Type:—U.S.A. Pennsylvania, 29 November 1805,
Muhlenberg (lectotype, selected by Moberg & Nash 1999: 5, H) [for collection locality and date, see Lendemer & Hewitt 2002:
187]; Anaptychia hypoleuca (Ach.) A. Massal., Atti Soc. Ital. Sci. Nat. 11:615 (1869); Heterodermia hypoleuca (Ach.) Trevis.,
Atti Soc. Ital. Sci. Nat. 11: 615 (1869).
Thallus foliose, rarely orbicular, usually irregularly spreading, loosely adnate, to 9 cm wide. Lobes 0.3–1.0 mm
wide, to ca. 1.5–2.5 mm wide at the tips, plane to weakly convex or weakly concave, sublinear-elongate, irregularly
branched, radiating; apices not ascending, usually discrete, ± lobulate along the lobe margins, the lobules ±
rounded, 0.1–0.2 mm wide, eciliate. Upper surface whitish to pale gray, without soralia, isidia or phyllidia. Medulla
white. Lower surface ecorticate, ± compact, white to beige in the centre; yellow pigments absent. Marginal rhizines
totally black or grayish at the base, simple or squarrosely branched, 1–3 mm long. Apothecia common, laminal,
substipitate, 1–7 mm wide; margin crenulate to lobulate, the lobules ecorticate on the lower surface; disc concave,
dark brown to brown-black, ± thinly gray pruinose. Ascospores Polyblastidia-type, ellipsoidal, mostly without
polyblastidia, albeit in some specimens, almost one third of the spores are with polyblastidia, 23–33 × 12–17 μm.
Pycnidia not seen.
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), japonene (major).
Nomenclature notes:Some authors, e.g. Kurokawa (1962: 42) quote Parmelia hypoleuca Muhl., Cat. Pl.
Amer. Sept. 1813:105 as basionym. But the genus Parmelia is outlined on p. 102 and no Parmelia hypoleuca is
mentioned there.
Distribution and habitat:—Occurs on bark of trees (Betula alnoides and Acer pseudoplatanus are reported)
in lower montane scrub, in lower montane rainforest and in hill evergreen forest in northern Thailand from 1185 to
1340 m above sea level; also recorded from North, Central and South America, Africa and Asia, e.g. Japan and
China.
Remarks:—The species is characterized by ± radiating lobe apices, with a whitish lower surface lacking lower
cortex, the absence of vegetative propagules, ± small ascospores mostly without sporoblastidia and the lack of
depsidones. Polyblastidium magellanicum is very similar, but is readily distinguished by larger ascospores
invariably with polyblastidia.
Material from Thailand examined:Loei: Phuluang Wildlife Sanctuary, trail to Pa Kob Lon Tae forest
protection, in lower montane scrub, on bark of unidentified tree, 1245 m, 17°11'32'' N, 101°33'43'' E 17°16'39'' N,
101°31'38'' E, 28 August 2008, S. Meesim & K. Buaruang, RU-MSPL 710, (RAMK 21156); Loei: Phuluang
Wildlife Sanctuary, Forest protection unit, area of Lon Tae, in lower montane scrub, on Acer pseudoplatanu, 1260
m, 17°14'18'' N, 101°33'19'' E, 12 November 2009, S. Meesim & K. Buaruang, RU-MSPL 565,, (RAMK 21122);
Phitsanulok: Phu Hin Rong Kla National Park, area of Kang Han Nam water fall, in lower montane rain forest, on
bark of unidentified tree, 1185 m, 16°59'31'' N, 100°00'44'' E, 4 June 2003, T. Dia, RU-TD 25, (RAMK 21585);
Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on Betula alnoides, 1340 m, 16°06'03''
N, 99°06'27'' E, 2 May 2006, S. Meesim RU-MW 16, (RAMK 21734); Nakon Ratchasima: Khao Yai National Park,
along the trail Pha Deaw Dai, in hill evergreen forest, on bark of unidentified tree,1280 m, 14°22'28'' N, 101°23'04''
E, 10 January 1999, R. Noicharoen RU-16065, (RAMK 5559).
6. Polyblastidium japonicum (M. Satô) Kalb, comb. nov. (Fig. 11A)
Mycobank MB 813844
Basionym: Anaptychia dendritica var. japonica M. Satô, J. Jap. Bot. 12: 427 (1936); Anaptychia japonica (Satô) Kurok., Beih. Nova
Hedwigia 6: 58 (1962). Type:—TAIWAN. Mt. Arisan, Mingestu, 24 Jan. 1936, M. Satô (Taiwan 10); TI n.v. Heterodermia
japonica (M. Satô) Swinscow & Krog, Lichenologist 8: 122 (1976).
Thallus foliose, irregularly spreading to ± orbicular, loosely adnate, to 5 cm wide, often coalescing to form colonies
up to 15 cm wide. Lobes 0.7–2.0 mm wide, to ca. 2–4(–5) mm wide at the tips, plane to weakly convex or weakly
concave, sublinear-elongate, dichotomously branched, radiating; apices ascending, usually discrete, ± lobulate
along the lobe margins, lobules ±rounded or sparingly branched, 0.3–0.8 mm wide, rarely with small soralia. Upper
surface greenish white, whitish to pale gray, sorediate; soredia farinose to granular, soralia labriform to capitate, on
lateral and terminal lobes, occasionally spreading along lobe margins. Medulla white. Lower surface ecorticate, ±
compact, white to violet-black in the centre; yellow pigments absent. Rhizines marginal, black, simple or
squarrosely branched, 1–3(–7) mm long. Apothecia very rare, laminal, substipitate, 1–8 mm wide; margin lobulate,
lobules ecorticate, partly yellow on the lower surface, apically sorediate; disc concave, dark brown to brown-black,
± gray pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 2–3 sporoblastidia, 40–45 × 20–23 μm. Pycnidia
common, immersed, then becoming emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), japonene (major), 16β-acetoxyhopane-6α,22-diol (trace), 6α-acetoxy-22-hydroxyhopane-25-oic
acid (trace), anaptychin 5 (minor).
Distribution and habitat:—This is a very rare species in Thailand. It was collected on bark of Cinchona
pubescens and on bark of Fagaceae in hill evergreen forest. A pantropical and pantemperate species.
Remarks:—This species is characterized by sorediate upper surface, ± radiating lobe apices, the absence of a
lower cortex, the white to blackish violet lower surface, ascospores with sporoblastidia and the presence of
atranorin and triterpenes. It was not present amongst the material we investigated, but has been reported by Aptroot
et al. (2007: 105). As no chemistry was given in that paper, the data may refer to the depsidone containing, P.
propaguliferum.
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Material reported from Thailand:—Chiang Mai: Doi Suthep-Pui National Park, in a medicinal garden on
bark of Cinchona pubescens, 1100 m, 18°48'17'' N, 98°54'43'' E, 12 October 2002, A. Aptroot 55188 (ABL); Doi
Suthep-Pui National Park, Doi Suthep ridge, Doi Suthep, San Khu summit, in montane oak forest on ridge, on bark
of Fagaceae, 1500 m, 18°48'56'' N, 98°53'41'' E, 13 October 2002, A. Aptroot 55214 (ABL).
Selected specimens examined:—PHILIPPINES. Mindanao: Cotabato province; between Lake Venado and
the peak of Mt. Apo, 2200–2800 m, 07°00'–06°59' N, 125°16.17–125°20' E, 12 June 2000, F. Schumm & Schwarz
(herb. Schumm 6175)—MASCARENE ISLANDS. Réunion: Street from le Vingt-Septième and Piton de la
Furnaise, in a small population of Sophora denudata and Philippia montana, 2250 m, 21°13'' S, 55°39' E, 29
August 1991, K. & A. Kalb (herb. Kalb 33709)—BRAZIL. Rio de Janeiro: Serra da Mantiqueira; Itatiaia, Parque
National do Itatiaia, at the headquarters of the N.P. on palm tree, 800 m, 22°20’ S, 44°35’ W, 21 July 1978, K. Kalb
& G. Plöbst (herb. Kalb 40306)—Costa Rica. Puntarenas: Fila Cruces Ridge, surroundings of Las Cruces Tropical
Botanical Garden, ca. 4 km SSE of San Vito, in a premontane rainforest, 1300 m, 31 December 1978, K. Kalb & G.
Plöbst (herb. Kalb 37785).
7. Polyblastidium microphyllum (Kurok.) Kalb, comb. nov. (Fig. 11B)
Mycobank MB 813845
Basionym: Anaptychia hypoleuca var. microphylla Kurok., J. Jap. Bot. 34: 123 (1959); Anaptychia microphylla (Kurok.) Kurok., Beih.
Nova Hedwigia 6: 44 (1962). Type:—JAPAN. Prov. Shinano: Higashi-Tikuma-gun, Sakakita-mura, 28 July 1953, Yamazaki s.n.;
(holotype, TNS n.v., isotype, TNS n.v.).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–15 cm wide. Lobes 0.5–2.0 mm wide, plane to
weakly convex or weakly concave, sublinear-elongate, irregularly to subdigitately branched; apices not ascending,
± discrete to contiguous at the periphery, with short lateral lobes, eciliate. Upper surface gray-white to gray,
epruinose, with marginal dorsiventral lobules, rarely also laminal, usually minutely dissected, often granular near
the tips, or entire phyllidia becoming granular and appearing sorediate. Medulla white. Lower surface ecorticate,
white to grayish or pale brown in the centre. Rhizines numerous, mainly marginal, concolorous with the thallus or
darkening and forming a dense marginal mat, irregularly branched. Apothecia rare, laminal, sessile to substipitate,
3–5 mm wide; margin phyllidiate, the phyllidia becoming ±sorediate; disc concave, brown to brown-black,
epruinose. Ascospores Polyblastidia-type, ellipsoidal, mostly without polyblastidia, albeit in some specimens,
some of the spores are with polyblastidia (Swinscow & Krog 1988: 98), 25–35 × 12–18 μm. Pycnidia initially
immersed, becoming emergent, visible as black dots; conidia bacilliform, 3–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P+ pale yellow; containing
atranorin (major), zeorin (major), aipolic acid (trace), 6α-acetoxyhopane-16β,22- diol (trace), japonene (trace), 6α-
acetoxy-16β,22-dihydroxyhopane-25-oic acid (minor), ±16β- acetoxyhopane-6α,22-dihydroxyhopane-25-oic acid
(trace), anaptychin 1 (trace).
Distribution and habitat:—Known from hill evergreen forest; also in South America, Australia, East Africa,
South Africa, South, South-east and East Asia and New Zealand.
Remarks:—This species is characterized by minutely dissected marginal phyllidia, ecorticate lower surface,
Polyblastidia-type ascospores and presence of the japonica-chemosyndrome of triterpenes. It was not found
amongst the material investigated, but may well occur in the country.
Material reported from Thailand:—Chiang Mai: Doi Inthanon National Park; Nakhon Nayok: Khao Yai
National Park (Boonpragob et al. 1998: 217).
Selected specimens examined: BRAZIL. Rio de Janeiro: Serra do Mar; between Paratí and Cunha, in a light
and humid coastal rainforest, 10 m, 23°10’ S, 44°50’ W, 2 November 1979, K. Kalb & G. Plöbst (herb. Kalb
40309); BRAZIL. São Paulo: Serra do Mar; between Taubaté and Ubatuba, ca. 150 km E of São Paulo, at the edge
of a humid and dense rainforest, 800 m, 23°15’ S, 45°15’ W, 18 June 1978, K. Kalb & G. Plöbst (herb. Kalb
40283)—KENYA. Rift Valley: Mount Longonot, ca. 70 km NW of Nairobi, 9 August 1985, K. Kalb & A. Schrögl
(herb. Kalb 14277)—PHILIPPINES. Cebu: Tops Lookout, ca. 7 km NW of Cebu City, 640 m, 10°22'15'' N,
129°52'16'' E, 14 August 2000, F. Schumm & Schwarz (herb. Schumm 7659).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
FIGURE 11. A. Polyblastidium japonicum (scale 5 mm). B. Polyblastidium microphyllum (scale 5 mm). C. Polyblastidium
propaguliferum (scale 5 mm). D. Polyblastidium queenslandicum (scale 5 mm). E. Polyblastidium togashii (scale 5 mm). F.
Polyblastidium violostriatum (scale 5 mm).
8. Polyblastidium propaguliferum (Vain.) Kalb, comb. nov. (Fig. 11C)
Mycobank MB 813846
Basionym: Anaptychia hypoleuca ssp. dendritica var. propagulifera Vain., Philipp. J. Sci., C, 8: 107 (1913); Type:—PHILIPPINES.
Benguet; Baguio, 150 m, May 1911, C.H. Robinson 14071; (TUR-VAIN 8003!); Anaptychia subheterochroa var. propagulifera
(Vain.) Kurok., J. Jap. Bot. 35: 241 (1960); Heterodermia dendritica var. propagulifera (Vain.) Poelt, Nova Hedwigia 9: 31
MONGKOLSUK ET AL.
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(1965); Heterodermia propagulifera (Vain.) Dey, In: Parker & Roane, Dist. Hist. Biota S. Appal. 4: 403 (1977); Anaptychia
japonica var. reagens Kurok., J. Jap. Bot. 35(8): 354 (1960). Type:—JAPAN. Sagami Prov.: Mt Kintoki, Hakone, 26 Apr. 1958,
S. Kurokawa 58064; (TNS, holotype n.v.; M, TNS, US, isotype n.v.); Heterodermia reagens (Kurok.) Elix, Australas. Lichenol.
67: 6 (2010).
Thallus foliose, usually irregularly spreading to rarely orbicular, loosely adnate, to 5 cm wide, but often coalescing
to form colonies up to 15 cm wide. Lobes 0.7–2.0 mm wide, to ca. 2–4 (–5) mm wide at the tips, plane to weakly
convex or weakly concave, sublinear-elongate, irregularly branched, radiating; apices ascending, usually discrete,
±lobulate along the lobe margins, lobules ± rounded or sparingly branched, 0.3–0.8 mm wide, developing small
soralia. Upper surface greenish white, whitish to pale gray, sorediate; soredia farinose to granular; soralia on lateral
or terminal lobes, occasionally spreading along the lobe margin, labriform to capitate . Medulla white. Lower
surface ecorticate, arachnoid, white to violet-black in the centre, lacking yellow pigments. Rhizines marginal,
black, simple or squarrosely branched, 1–3(–7) mm long. Apothecia very rare, laminal, substipitate, 1–8 mm wide;
margin lobulate, lobules ecorticate, partly yellow on the lower surface, apically sorediate; disc concave, dark brown
to brown-black, ±gray pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 2–3 sporoblastidia, 40–45 ×
20–23 μm. Pycnidia common, initially immersed, becoming emergent, visible as black dots; conidia bacilliform,
4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ dark yellow;
containing atranorin (major), zeorin (major), japonene (major), 16β-acetoxyhopane- 6α,22-diol (trace), 6α-
acetoxy-22-hydroxyhopane-25-oic acid (trace), norstictic acid (minor), connorstictic acid (trace), ± salazinic acid
(minor or trace), anaptychin-5 (minor), anaptychin-1 (trace).
Nomenclature notes:Kurokawa (1962) and all subseqent authors that we have checked (e.g. Poelt 1965,
Dey 1977), cited Anaptychia dendritica (Pers.) Vain. var. propagulifera Vain. as basionym. However, Vainio (1890:
134) as well as Vainio (1913: 106) wrote: *A. dendritica (Pers.) Wain. and the entry in 1913 was followed by the
footnote: "* An asterisk indicates a subspecies". So the correct basionym is that given above.—Elix (2011c) treated
this species as synonym with Heterodermia japonica, but TLC of the holotype revealed the presence of norstictic
acid, thus Heterodermia reagens becomes a synonym of P. propaguliferum.
Distribution and habitat:—This species occurs on bark of trees (Cassia bakeriana, Cinnamomum porrectum,
Acer pseudoplatanus, Betula alnoides, Lithocarpus polystachyus and Castanopsis are reported) and, more rarely,
on mossy rocks in lower montane rainforest, hill evergreen forest, dry evergreen forest, lower montane scrub and
lower montane forest from 900–2210 m above sea level; also in Australia, Central and South America, Africa and
Asia. Interestingly, this common and widespread species had not previously been reported for Thailand, hence it is
a new addition to the Thai lichen biota.
Remarks:—This lichen is characterized by the sorediate lobe apices, an ecorticate lower surface that lacks
yellow pigments, and the presence of norstictic acid. This is the depsidone-containing counterpart of P. japonicum.
Material from Thailand examined:—There are too many collections to be listed individually, but a few
representative specimens are cited. The species is known from the provinces of Nan: Doi Phu Kha, area of view
point Doi Phu Kha, on Cassia bakeriana in hill evergreen forest, 1470 m, 19°18'12.83'' N, 101°09'54.97'' E, S.
Meesim RU-DK28.1 (RAMK 21606)—Chiang Mai: Queen Sirikit Botanic Garden; Doi Suthep-Pui, along trail
from research centre of Suan Song Saeng Kasetsart University to National guard units 1 Pha Dum, in hill evergreen
forest, on bark of Cinnamomum porrectum, 1420 m, 18°45'04'' N, 98°53'16'' E, 30 April 2005, W. Hongsachart
RU-WD 55, (RAMK 21655); Doi Inthanon National Park, along trail to Kew Mae Parn, on bark of unidentified tree
in hill evergreen forest, 2210 m, 18°33'26.87'' N, 98°25'56.32'' E, 27 October 2005, N. Juntawong RU-NJ 1999
(RAMK 21626—Loei: Na Haeo National Park, area of check point 1, in hill evergreen forest, on rock, 1260 m,
17°30'15'' N, 100°56'25'' E, 12 July 1995, P. Mongkolsuk 4180, (RAMK 21871); Phuluang Wildlife Sanctuary,
forest protection unit office, Lon Tae substation, in lower montane scrub forest, on bark of Acer pseudoplatanus,
1260 m, 17°14'18'' N, 101°33'19'' E, 12 November 2009, S. Meesim & K. Buaruang RU-MSPL 651, (RAMK
21140)—Phitsanulok: Phu Hin Rong Kla National Park, area of Kang Han Nam water fall, in lower montane rain
forest, on rock, 1165 m, 16°59'31'' N, 100°00'44'' E, 6 June 2003, P. Mongkolsuk RU-PM 151, (RAMK
21769)—Kamphaeng Phet: Maewong, along trail Chong Yen, in hill evergreen forest, on bark of Betula alnoides,
1340 m, 16°06'03'' N, 99°06'27'' E, 2 May 2006, S. Meesim RU-MW 8, (RAMK 21408)—Nakon Ratchasima: Khao
Yai National Park, north of the main road no. 3077 from the park's headquarters to Khao Khaeo, in hill evergreen
forest, on bark of unidentified tree, 1390 m, 14°21'40'' N, 101°23'35'' E, 4 November 1997, K. Boonpragob RU-
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
9719, (RAMK 8983)—Phetchaburi: Kaeng Krachan, along trail Khao Phanoenthung, in hill evergreen forest, on
bark of Lithocarpus polystachyus, 1210 m, 12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim, RU-MW 100,
(RAMK 21610).
9. Polyblastidium queenslandicum (Elix) Kalb, comb. nov. (Fig. 11D)
Mycobank MB 813847
Basionym: Heterodermia queenslandica Elix, Australas. Lichenol. 67: 5 (2010b).
Type:—AUSTRALIA. Queensland: Mt. Baldy, 4 km SW of Atherton, 17°17’ S, 145°27’ E, 1080 m, on sapling along margin of
regrowth rainforest, 25 June 1984, J.A. Elix 16310 & H. Streimann (CANB!, holotype).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, to 8 cm wide. Lobes 0.5–2.0 mm wide, plane to
weakly convex or weakly concave, sublinear-elongate, irregularly to subdichotomously branched, radiating; apices
plane to subascending, usually discrete, with small lateral lobules; isidia and soredia absent. Upper surface greenish
white, whitish or gray. Medulla white. Lower surface ecorticate, whitish to pale brown or, rarely, purple-black in
the centre. Rhizines predominantly marginal or submarginal, pale gray to black, ±densely squarrosely branched,
1.0–4.5 mm long. Apothecia laminal, substipitate to stipitate, 2.5–5.5 mm wide; margin densely lobulate, lobules to
2.5 × 2.0 mm; disc concave, pale to dark brown, epruinose or sparsely gray-pruinose. Ascospores Polyblastidia-
type, ellipsoidal, with 0–3 sporoblastidia, 40–45 × 20–24 μm. Pycnidia common, immersed, visible as black dots;
conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ yellow-orange;
containing atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (hopane-6α,
16β,22-triol) (major), 6α-acetoxyhopane-16β,22-diol (minor), norstictic acid (major), salazinolide (major),
consalazinolide (minor).
Distribution and habitat:—Rare on bark in evergreen montane rainforest: also in Australia.
Remarks:—The species is characterized by narrow, sublinear-elongate lobes with an ecorticate lower surface,
the absence of soredia and isidia but the presence of marginal lobules and the presence of atranorin, zeorin,
norstictic acid, salazinolide and consalazinolide.
Material from Thailand examined:—Chiang Mai: Lumphun, Mae On, descent from Doi Mon Larn to Mae
Kam Pong village, in an evergreen montane rainforest, on bark of an unidentified tree, 1500 m, 18°51'22'' N,
99°22'02'' E, 19 March 2008, K. Kalb et al. (herb. Kalb 36966).
10. Polyblastidium togashii (Kurok.) Kalb, comb. nov. (Fig. 11E)
Mycobank MB 813848
Basionym: Anaptychia togashii Kurok., Beih. Nova Hedwigia 6: 68 (1962). Type:—India. Sikkim: Jongri, 4000 m, 21 May 1960, M.
Tog a s h i (holotype TI, n.v., isotypes KrK, US); Heterodermia togashii (Kurok.) D.D. Awasthi, Geophytology 3(1): 114 (1973).
Thallus foliose, rarely orbicular, usually irregularly spreading, loosely adnate, to 9 cm wide. Lobes 0.3–1.0 mm
wide, to ca. 1.5–2.5 mm wide at the tips, plane to weakly convex or weakly concave, sublinear-elongate, irregularly
branched, radiating; apices ascending, usually discrete, not lobulate along the lobe margins, eciliate. Upper surface
whitish to cream or pale gray, without soralia, isidia or phyllidia. Medulla white. Lower surface ecorticate, ±
arachnoid, white to beige in the centre; yellow pigments absent. Marginal rhizines totally black or grayish at the
base, simple, but basally squarrosely branched and forming a dense mat, 1–3 mm long. Apothecia not common,
laminal, substipitate, 1–5 mm wide; margin crenulate to lobulate, the lobules ecorticate on the lower surface; disc
concave, dark brown to brown-black, ± thinly gray pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 2 or
3 ± large sporoblastidia, 33–50 × 15–25 μm. Pycnidia rare, immersed, then becoming emergent, visible as black
dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; containing atranorin (major),
zeorin (major), japonene (major).
Distribution and habitat:—Occurs on bark of trees (Castanopsis is reported) or over mosses and liverworts
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on small branches in hill evergreen forest in northern Thailand from 1413 to 2565 m above sea level, also recorded
from India and Nepal; new addition to the Thai lichen biota.
Remarks:This species is characterized by the lack of vegetative propagules, ± radiating lobe apices, the
absence of a lower cortex, the whitish lower surface, ± large ascospores with sporoblastidia and the lack of
depsidones. Polyblastidium togashii is very similar to P. hypoleucum, but is readily distinguished by larger
ascospores (23–30 × 11–16 μm in P. hypoleucum). Furthermore, polyblastidia are very rare in P. hypoleucum. It is
distinguished from P. magellanicum by the marginal rhizines which are richly branched at the base.
Material from Thailand examined:—Chiang Mai: Doi Inthanon National Park, along trail Ang Ka Doi
Inthanon, in hill evergreen forest, on bark of unidentified tree, 2240 m, 18°48'27'' N, 98°53'27'' E, 30 April 2005,
W. Hongsachart, RU-WD 108, (RAMK 21458); ibid., W. Hongsachart, RU-WD 103, (RAMK 21454); Chiang Mai:
Doi Inthanon National Park, along trail to the peak of Doi Inthanon, in hill evergreen forest, on bark of unidentified
tree, 2565 m, 18°35'22'' N, 98°29'15'' E, 7 September 2005 N. Juntawong, RU-NJ 2244, (RAMK 21337); ibid., 29
October 2005 N. Juntawong, RU-NJ 2267, (RAMK 21746); Chiang Mai: Doi Suthep-Pui, along tourist trail to
Agricultural Centre Ban Doi-Pui, in hill evergreen forest, on Castanopsis sp., 1413 m, 18°45'4" N, 98°53'16'' E, 30
April 2005, W. Hongsachart, RU-WD 102, (RAMK 21453).
11. Polyblastidium violostriatum (Elix) Kalb, comb. nov. (Fig. 11F)
Mycobank MB 813849
Basionym: Heterodermia violostriata Elix, Australas. Lichenol. 69: 16 (2011).
Type:—AUSTRALIA. Queensland: Paluma Ra., Paluma Village, 40 km S of Ingham, 19°01’S, 146°13’E, 830 m, on stem of exotic
tree in maintained garden, 29 October 1995, H. Streimann 58057; (holotype, CANB!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, 5–12 cm wide. Lobes 1.0–1.5 mm wide, ca. 2–3
mm wide at the tips, plane to weakly convex or weakly concave, sublinear-elongate, irregularly to digitately
branched, radiating; apices not ascending, contiguous to discrete. Upper surface greenish white, whitish gray to
creamcolored, epruinose, with marginal to rarely laminal dorsiventral phyllidia, phyllidia minutely dissected, often
granular near the tips, or entire phyllidia becoming granular and appearing sorediate. Medulla white. Lower surface
ecorticate, arachnoid, violet-gray in the centre, violet-striate towards the apices; yellow pigments absent. Rhizines
marginal, at first simple and concolorous with the lower surface, later black and squarrosely branched, 2–4 mm
long. Apothecia laminal, adnate to substipitate, 1–4 mm wide; margin phyllidiate, the phyllidia becoming elongate
and deeply dissected; disc concave, dark brown to blackish brown, ±grayish pruinose. Ascospores Polyblastidia-
type, ellipsoidal, with 0–2 small sporoblastidia, 30–45 × 15–25 μm. Pycnidia common, immersed, then emergent,
visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P+ pale yellow; containing
atranorin (major), zeorin (major), 16β-acetoxyhopane-6α,22-diol (major), leucotylin (minor), 6α-acetoxyhopane-
16β,22-diol (major or minor), 6α,16β-diacetoxyhopane- 22-ol (minor), chloroatranorin (minor).
Distribution and habitat:—This very rare species occurs on rocks and bark of trees in a hill evergreen forest
in the north of Thailand at 1320 m above sea level; also in Australia; new addition to the Thai lichen biota.
Remarks:characterized by lobes with an ecorticate, violet-gray to violet lower surface, marginal phyllidia,
Polyblastidia-type ascospores and the presence of terpenes in the medulla. Previously misidentified as P.
appendiculatum or P. microphyllum; the former is distinguished from P. violostriatum by the presence of additional
norstictic acid, while P. microphyllum has a white lower surface and contains the japonica chemosyndrome of
triterpenes.
Material from Thailand examined:—Chiang Mai: Doi Inthanon National Park, area along trail to Siriphum
waterfall, in hill evergreen forest, on bark of unidentified tree, 1320 m, 18°32'50'' N, 98°30'56'' E, 9 September
2005, N. Juntawong RU-NJ 277, (RAMK 21760);
Further new combinations:
Polyblastidium magellanicum (Zahlbr.) Kalb, comb. nov., Mycobank MB 813850, Basionym: Anaptychia magellanica Zahlbr., Kungl.
Svenska Vetensk. Akad. Handl. 57: 54 (1917).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Polyblastidium dendriticum (Pers.) Kalb, comb. nov., Mycobank MB 813855, Basionym: Borrera dendritica Pers., In: Gaudichaud,
M.C. Voyage autour du monde: 207 (1826).
Polyblastidium squamulosum (Degel.) Kalb, comb. nov., Mycobank MB 813841, Basionym: Anaptychia squamulosa Degel., Ark.
Bot. 30A (3): 76 (1942 [1941]).
Chemistry:—Atranorin (major), zeorin (major), 6α-acetoxyhopane-16β,22-diol (submajor), leucotylin (minor to
trace).
Specimen examined:—U.S.A. Virginia: Giles Co., Mountain Lake, on Quercus alba, 1964, W.L. Culberson
11474 (S).
The Heterodermia comosa-group
This group of species is similar to Leucodermia, but differs in the main by having a foliose to subfruticose thallus
with typically ascending, spathulate or paddle-shaped lobes and white marginal rhizines. At present no isidiate or
phyllidiate species are known.
The group conforms to Anaptychia sect. Polyblastidium ser. Podocarpae (Kurokawa 1962: 84).
Key to the Thai species of the Heterodermia comosa-group
(TLC in solvents A and B' is necessary for precise identification; furthermore, it is strongly recommended to
perform cochromatography with a lichen containing testacein, e.g. Parmelia testacea Stirt., Pyxine retirugella Nyl.,
P. copelandii Vain. (Kalb 2004).
1. Thallus surface and exciple of apothecia ciliate (Figs. 4A, 4B) ............................. H. comosa (Eschw.) Follmann & Redón
- Thallus and exciple of apothecia eciliate ............................................................................................................................... 2
2. Medulla with dissectic acid.................................................................................................. H. pandurata (Kurok.) J.C. Wei
- Medulla without dissectic acid .............................................................................................................................................. 3
3. Medulla with norstictic acid............................................................................................. H. podocarpa (Bél.) D.D. Awasthi
- Medulla without norstictic acid.............................................................................................................................................. 4
4. Lower surface with superimposed layer of cotton-woolly hyphae which are impregnated with yellow to orange-brown pig-
ments ................................................................................................................... H. hypochraea (Vain.) Swinscow & Krog
- Lower surface ± white............................................................................................................................................................ 5
5. Thallus sorediate .................................................................................................................................................................... 6
6. Medulla with japonene, testacein absent....................................................................... H. galactophylla (Tuck.) W.L. Culb.
- Medulla with testacein, japonene absent......................................................................................................... H. archeri Elix
- Thallus esorediate, medulla without japonene....................................................................................................................... 7
7. Thallus attached to the substrate, only apothecia bearing lobes ascending ........................... H. stellata (Vain.) W.A. Weber
Thallus forming rosettes of ascending lobes, attached only at the central parts ................... H. incana (Stirt.) D.D. Awasthi
1. Heterodermia archeri Elix, Australas. Lichenol. 68: 16 (2011a). (Fig. 12A)
Type:—AUSTRALIA. Queensland; Flaggy Ck, Black Mountain Rd, 9 km NNW of Kuranda, 16°47’ S, 145°36’ E, 420 m, on felled
trees along the rainforest margin, 7 July 1984, J.A. Elix 17595 & H. Streimann (holotype, CANB!).
Thallus foliose, orbicular to irregularly spreading, loosely adnate, to 5 cm wide. Lobes 0.5–1.2 mm wide,
broadening to ca. 2−3 mm wide at the tips and becoming paddle-shaped, plane to weakly convex, sublinear-
elongate, irregularly to subpalmately branched, radiating; apices ± ascending, with lateral simple, white, 0.5–1.2
mm long rhizines visible from above. Upper surface greenish white, whitish to cream-colored, sorediate; soredia
farinose to granular, in labriform soralia, occasionally to 5 mm wide and spathulate. Medulla white. Lower surface
ecorticate, whitish at the margins, darkening to pale brown centrally. Rhizines marginal, simple or becoming richly
branched, white or creamish, 0.5–2.0 mm long. Apothecia and pycnidia not seen.
Chemistry:—Cortex and medulla K+ yellow, C–, KC–, P+ pale yellow; containing atranorin (major), zeorin
(major), 6α-acetoxyhopane-16β,22-diol (minor), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (minor),
testacein (minor).
Distribution and habitat:—Rare on twigs and on bark of trees (Shorea sp. is reported) in dry evergreen forest
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and hill evergreen forest from 300 to 1210 m above sea level; also in Australia and Papua New Guinea; new
addition to the Thai lichen biota.
Remarks:—characterized by the loosely adnate thallus with narrow, sublinear-elongate lobes with pale
marginal rhizines and spathulate soralia on some of the subascending lobe apices, a white, ecorticate lower surface
and the presence of testacein.
Material from Thailand examined:—Sakon Nakhon: Phu phan National Park; Tha are Pha Nang Mern, 300
m, 17°03'43'' N, 103°58'23'' E, on bark of Shorea sp. in a dry evergreen forest, 7 October 1997, P. Mongkolsuk 8974
(RAMK 21893)—Phetchaburi: Kaeng Krachan, along trail Khao Phanoenthung before km 36, in hill evergreen
forest, on bark of unidentified tree, 1210 m, 12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim RU-DK 118,
(RAMK 21845).
2. Heterodermia comosa (Eschw.) Follmann & Redón, Willdenowia 6: 446 (1972) (Figs. 1F, 4A, 4B)
Parmelia comosa Eschw., in K.F.P. von Martius, Icon. Pl. Cryptog. 2: 26 (1828); Anaptychia comosa (Eschw.) A. Massal., Mem.
Lichenogr. 39 (1853); Physcia comosa (Eschw.) Nyl., Syn. Meth. Lich. 1(2): 416 (1860). Type:—BRAZIL. Pará; near Porta de
Môz, on branches of tree; not located but based on illustration tab. XII, fig. 1, fide S.Kurokawa, Beih. Nova Hedwigia 6: 103
(1962). Epitype (designated by Moberg & Nash, 1999: 4):—BRAZIL. Minas Gerais, Caldas, ad arbores, August 1873, H. Mosén
(UPS!)
Thallus foliose to subfruticose, forming small rosettes or tufts of ascending lobes, 3–7 cm wide. Lobes 0.7–5.0
(–8.0) mm wide, sublinear, spathulate or paddle-shaped, convex, rarely branched, ascending or suberect, partially
imbricate, with rounded apices when sterile, ciliate; cilia prominent, usually simple, marginal and laminal, whitish,
2–4 mm long. Upper surface white to grayish white, sorediate; soredia on the undersides of lobe tips; marginal
rhizines whitish, mostly simple, sometimes furcate or irregularly ramified. Medulla white. Lower surface
ecorticate, arachnoid, uniformly white (South American material, including the epitype, partly yellow-brown),
apically sorediate when sterile, erhizinate. Apothecia subterminal to terminal on ascending lobes, substipitate,
1–5(–10) mm wide; margin crenate or lobulate, ciliate; disc dark brown, densely whitish gray pruinose. Ascospores
Polyblastidia-type, narrowly ellipsoid to fusiform, with 2–3 sporoblastidia, 31–35 × 13–16 μm. Pycnidia not seen.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; pigmented lower surface K+
violet; containing atranorin (major), zeorin (major), 6α-acetoxyhopane-16β,22-diol (minor), ± 7-chloroemodin
(minor to trace).
Distribution and habitat:—Occurs on the twigs or trees and shrubs in a garden situated in forest remnants at
750–800 m above sea level; also in Australia, North, Central and South America, Africa and Asia.
Remarks:This lichen is characterized by spathulate or paddle-shaped lobes with cilia on the upper surface,
and soralia beneath lobe apices. It is possible that the type race from South America, with the yellow-brown
pigmented lower surface, is a different species to the Palaeotropical taxon. However, specimens lacking the
anthraquinone (7-chloroemodin) are also known from the Neotropics. The anthraquinone is distinctly visible on the
epitype, which was collected in 1873.
Material reported from Thailand:—Chiang Mai: Mushroom Research Centre along road to Pae, on bark in
Camellia garden in forest remnants, 750–800 m, 19°07'11'' N, 98°44'01'' E, 1–3 July 2004, Aptroot 61359 (ABL).
(Aptroot et al. 2007: 104).
Selected specimens examined: BRAZIL. Minas Gerais: Serra da Mantiqueira; Vila Monte Verde, ca. 30 km E
of Camanducaia, on a deciduous tree at the edge of in a light and ± dry forest, 1850 m, 22°50’ S, 46°00’W, 7–11
September 1978, K. Kalb & G. Plöbst (herb. Kalb 37798); Brazil. Minas Gerais: Serra do Espinhaço; Serra do
Caraça, above the monastery Caraça, in an open cerrado, 1450 m, 20°10’ S, 43°30’ W, 8. July 1978, K. Kalb & G.
Plöbst (herb. Kalb 37787)—PARAGUAY. Presidente Hayes: Gran Chaco; ca. 50 km NW of Filadelfia, at the edge
of a dense forest with many cacti, 250 m, 22°07 S, 60°30' W, 6 June 1980, K. Kalb (herb. Kalb 37804).
3. Heterodermia galactophylla (Tuck.) W.L. Culb., Bryologist 69: 482 (1967) (Figs. 1E, 12B)
Parmelia ciliaris β galactophylla Tuck., Proc. Amer. Acad. Arts Sci. 1: 224 (1847).; Type:—U.S.A. Maine, ex herb. Oakes (holotype,
FH-Tuck n.v.); Anaptychia galactophylla (Tuck.) Trevis., Flora 44: 52 (1861).
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
Thallus foliose to subfruticose, usually attached to the substrate, forming rosettes or tufts of apically ascending
lobes, 3–5 cm wide, but sometimes coalescing to form colonies up to 17 cm. Lobes 0.4–2.5 (–8.0 at sorediate tips)
mm wide, sublinear, spathulate or paddle-shaped, convex, rarely branched, ascending or suberect, partially
imbricate, with rounded apices when non-sorediate. Upper surface white to grayish white or seldom glaucous gray,
sorediate; soredia on the undersides of reflexed and erect and widened lobe tips; marginal rhizines whitish to
somewhat darkened, usually simple to furcate, but becoming thyrsiformly ramified with age. Medulla white.
Lower surface ecorticate, arachnoid, uniformly white (South American material partly yellow-brown). Apothecia
very rare, not present in Thai material, subterminal to terminal on ascending lobes, substipitate, 1–2.5 (–3) mm
wide; margin crenate or lobulate, ciliate; disc dark brown, densely white pruinose. Ascospores Polyblastidia-type,
narrowly ellipsoidal to fusiform, with 2–3 sporoblastidia, 31–35 × 13–16 μm. Pycnidia not seen.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; pigmented lower surface K+
violet; containing atranorin (major), zeorin (major), japonene (submajor to minor).
Distribution and habitat:—Occurs on the twigs of trees and shrubs in a garden at 750–800 m above sea level;
also in North, Central and South America, Africa and Asia.
Remarks:—This lichen is characterized by the elongate lobes without cilia on the upper surface, soralia
beneath the widened and reflexed lobe apices and a chemistry without depsidones. We could find no mention of
apothecia for this species in the literature, but in our vast collections from South America we have located one
fertile specimen (herb. Kalb 37797). Probably, H. galactophylla is the sorediate counterpart of H. trichophora
(Kurok.) Trass, as the two species have the same chemistry.
Material reported from Thailand:—Chiang Mai: Mushroom Research Centre along road to Pae, on bark in
Camellia garden in forest remnants, 750–800 m, 19°07'11'' N, 98°44'01'' E, 1–3 July 2004, Aptroot 61331 (ABL).
(Aptroot & al. 2007: 104).
Specimens examined: BRAZIL. Minas Gerais: Serra da Mantiqueira; above Vila Monte Verde, ca. 17 km SE
of Camanducaia, in a very disturbed montane rainforest, 1800 m, 22°52' S, 46°02' W, 3 July 1979, K. Kalb & G.
Plöbst (herb. Kalb 37797)—BRAZIL. Mato Grosso: Serra dos Coroados; Buriti, nature reserve of the Protestant
school of Buriti, ca. 6 km SW of Buriti in a submontane rainforest, 600 m, 15°30’ S, 55°40’ W, 8 July 1980, K.
Kalb & m, Marcelli (herb. Kalb 37800)—BRAZIL. Rio de Janeiro: Serra da Mantiqueira; Itatiaia, Parque National
do Itatiaia, in a humid and dark primary rainforest, 1100 m, 22°20’ S, 44°35’ W, 21. July 1978, K. Kalb & G. Plöbst
(herb. Kalb 39358)—BRAZIL. São Paulo: Serra do Mar; Serra de Paranapiacaba, ca. 80 km SW of São Paulo, in a
light and humid rainforest along the Rio Juquiá, 700 m, 24°00’ S, 47°20’ W, 24 March 1978, K. Kalb & G. Plöbst
(herb. Kalb 39361).
4. Heterodermia hypochraea (Vain.) Swinscow & Krog, Lichenologist 8(2): 119 (1976) (Fig. 12C)
Anaptychia hypochraea Vain., Bot. Mag. (Tokyo) 35: 59 (1921); Anaptychia podocarpa var. hypochraea (Vain.) M. Satô, J. Jap. Bot.
12: 431 (1936). Type:—JAPAN. Prov. Inaba: Mt. Sensoku, ad corticem arboris, 25 September 1915, A. Yasuda 151 (holotype,
TUR-Vain. 7958!)
Thallus foliose to subfruticose, forming small rosettes or irregularly spreading, adnate to loosely adnate, 2–5 cm
wide. Lobes 0.5–2.0 mm wide, sublinear, convex, rarely flat or partly concave, irregularly branched, suberect or
ascending at the apices, partially imbricate, discrete to contiguous, non-ciliate, Upper surface grayish white or gray,
epruinose; soredia and isidia absent. Medulla white. Lower surface ecorticate, arachnoid, ± canaliculate, white,
partly or totally impregnated by an orange to orange-brown pigment. Marginal rhizines simple or irregularly
branched, whitish or somewhat darker at the tips, 1–4 mm long. Apothecia common (not seen in the holotype),
substipitate to stipitate, 1–5 mm wide, subterminal on ascending lobes; margin crenate or with well-developed
squamules, eciliate; disc brown to dark brown, often white-pruinose. Ascospores Polyblastidia-type, ellipsoidal,
with 2–3 small sporoblastidia, 30–45 × 17–22 μm. Pycnidia not seen.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ orange; orange
pigment on lower surface K+ violet; containing atranorin (major), zeorin (major), japonene (major), 7-
chloroemodin (minor) and dissectic acid (minor).
Distribution and habitat:—Rare on branches and bark of trees (Adenanthera pavonina is reported) in hill
evergreen forest and lower montane scrub from 1210 to 1470 m above sea level; also in Asia, Africa and South
America.
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FIGURE 12. A. Heterodermia archeri, holotype (scale 5 mm). B. Heterodermia galactophylla, apothecia with ciliate exciple (scale 1
mm). C. Heterodermia hypochraea, holotype (scale 5 mm). D. Heterodermia incana (scale 5 mm). E. Heterodermia podocarpa (scale
1.5 mm). F. Heterodermia stellata, lectotype (scale 1 mm).
Remarks:This species might be confused with H. flabellata, but differs by having a comosa-like growth
form and in containing dissectic acid and japonene (rather than 16β-acetoxyhopane-6α,22- diol as a major terpene).
Material from Thailand examined:—Nan: Doi Phu Kha, area of view point at Doi Phu Kha, in hill evergreen
forest, on bark of Toona ciliata, 1470 m, 19°18'13'' N, 101°09'55'' E, 10 May 2006, S. Meesim RU-DK 27, (RAMK
21425); Loei: Na Haeo National Park, area of check point 1, in hill evergreen forest, on Adenanthera pavonina,
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1260 m, 17°30'15'' N, 100°56'25'' E, 12 July 1995, P. Mongkolsuk 4508, (RAMK 11988); ibid., Phuluang Wildlife
Sanctuary, area of Kok Nok Kraba, in lower montane scrub, on bark of unidentified tree, 1490 m, 17°16'49'' N,
101°31'29'' E, 28 August 2005, W. Hongsachart, RU-WD 172, (RAMK 21197); Nakon Ratchasima, Khao Yai
National Park, north of the main road no. 3077 from the park's headquarters to Khao Khaeo, in hill evergreen
forest, on bark of unidentified tree, 1390 m, 14°21'40'' N, 101°23'35'' E, 21 August 1997, K. Boonpragob et al. RU-
8360, (RAMK 5553); Phetchaburi: Kaeng Krachan, along trail Khao Phanoenthung before km 36, in hill evergreen
forest, on bark of unidentified tree, 1210 m, 12°53'07'' N, 99°37'53'' E, 12 February 2006, S. Meesim RU-DK 114,
(RAMK 21428).
5. Heterodermia incana (Stirt.) D.D. Awasthi, Geophytology 3(1): 114 (1973) (Fig. 12D)
Physcia incana Stirt., Proc. Roy. Philos. Soc. Glasgow 11: 322 (1879); Anaptychia incana (Stirt.) Zahlbr., Cat. Lich. Univ. 7: 727
(1931); Type:—INDIA. Darjeeling; J.A. Thomson 13 (holotype, BM n.v.).
Thallus foliose, forming small rosettes or tufts of suberect to ascending lobes, 3–6 cm wide. Lobes 1–4 mm wide,
irregularly to subdichotomously branched, convex, epruinose or slightly pruinose at the tips, not ciliate. Upper
surface grayish white, sometimes darkening at the centre, soralia and isidia absent. Marginal rhizines whitish,
somewhat darkening at the tips, squarrosely or fruticosely branched. Medulla white. Lower surface ecorticate,
arachnoid, uniformly white. Apothecia commonly numerous, pedicellate, subterminal, 1–8 mm wide, margins
smooth to lacinulate, sometimes ciliate, disc darkbrown, densely white pruinose. Ascospores Polyblastidia-type, ±
constricted at the centre with 2–3 small sporoblastidia 52–56 × 22–27 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow, C–, P–; atranorin (major) and zeorin
(major).
Distribution and habitat:—This rare species occurs on bark of unidentified trees in lower montane scrub and
coniferous forest from 1470 to 1520 m above sea level; also in Taiwan, Java, India and Nepal.
Remarks:—H. incana is similar to H. himalayensis (D.D. Awasthi) D.D. Awasthi, but differs in lacking
norstictic acid.
Material from Thailand examined:—Loei: Phuluang Wildlife Sanctuary, area of Kok Nok Kraba and Ban Po
Ra Dok, in lower montane scrub, on bark of unidentified tree, 1470 m, 17°16'46'' N, 101°31'08'' E, 29 August 2005,
S. Meesim & K. Buaruang, RU-WD155, (RAMK 21183); area of Kok Nok Kraba, along trail from Pha Chang Parn
to Pha Taloen, in lower montane scrub, on bark of unidentified tree, 1470 m, 17°16'46'' N, 101°31'08'' E, 24 June
2008, S. Meesim & K. Buaruang, RU-MS 76.1, (RAMK 21081); ibid, in lower montane scrub, on bark of
unidentified tree, 1470 m, 17°16'46'' N, 101°31'08'' E, 13 November 2009, S. Meesim & K. Buaruang, RU-
MS0265, (RAMK 20682); area of Lan Suriyan, in lower montane scrub, on bark of unidentified tree, 1470 m,
17°16'44'' N, 101°31'07'' E, 25 June 2009, S. Meesim & K. Buaruang, RU-MS29.1, (RAMK 21022); forest
protection unit, along trail to Tae, area of Pak Dum, in coniferous forest, on bark of unidentified tree, 1520 m,
17°15'07'' N, 101°31'05'' E, 29 August 2005, S. Meesim & K. Buaruang, RU-MSPL 591, (RAMK 21124).
6. Heterodermia pandurata (Kurok.) J.C. Wei, An enumeration of lichens in China: 112 (1991)
Anaptychia pandurata Kurok., Beih. Nova Hedwigia 6: 95 (1962); Type:—JAPAN. Province Kii: Mt. Koya, 18 April 1926, Y.
Numaziri 597 (holotype, TNS, n.v.).
Thallus foliose to subfruticose, forming small rosettes or irregularly spreading, adnate to loosely adnate, 2–6 cm
wide. Lobes 0.5–2.0 mm wide, sublinear, convex, rarely flat or partly concave, irregularly branched, suberect or
ascending at the apices, partially imbricate, discrete to contiguous. Upper surface grayish white or gray, epruinose;
soredia and isidia absent. Medulla white. Lower surface ecorticate, arachnoid, ± canaliculate, white, partly or
totally impregnated by an orange to orange-brown pigment. Marginal rhizines simple or irregularly branched,
whitish or somewhat darker at the tips, 1–4 mm long. Apothecia common, substipitate to stipitate, 1–6 mm wide,
subterminal on ascending lobes; margin crenate or with well-developed squamules, eciliate; disc brown to dark
brown, often white pruinose. Ascospores Polyblastidia-type, ellipsoidal, with 2–3 sporoblastidia, 35–45 × 16–22
μm. Pycnidia not seen.
MONGKOLSUK ET AL.
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Phytotaxa 235 (1) © 2015 Magnolia Press
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ orange; orange
pigment on lower surface K+ violet; containing atranorin (major), zeorin, 7-chloroemodin and dissectic acid.
Distribution and habitat:—Rare on branches and bark of trees in Australia, Laos, Japan and probably in
Thailand.
Remarks:—At this time we cannot be sure that this species occurs in Thailand. It has been reported from Laos
and could eventually be found in Thailand as well.
Material reported from Thailand:—Hase Plantation, probably Laos (Wolseley et al. 2002: 28).
7. Heterodermia podocarpa (Bél.) D.D. Awasthi, Geophytology 3: 114 (1973) (Fig. 12E)
Parmelia podocarpa Bél., Voy. Indes-Orient., Bot. 2, Crypt.: 122, pl. 13, fig. 1 (1834); Anaptychia podocarpa (Bél.) A. Massal., Atti
Reale Ist. Veneto Sci. Lett. Arti, ser. 3, 5: 249 (1860). Type:—[MASCARENE ISLANDS]. Île de Bourbon [Réunion] Sainte-
Suzanne, sur les troncs pourris des arbres, pl. 13, fig. 1. Epitype (designated here): MASCARENE ISLANDS. Réunion: Plaine-
des-Palmistes, W-side of village, garden of Gîte rural 104 with Guave shrubs and Casuarina tree, on Psidium guajava, 1100 m,
55°37.1' E, 21°08.0' S, 1 June 2008, v.d. Boom & Sérusiaux 40294 (B!).—Physcia barbifera Nyl., Syn. Lich. 1: 416 (1860);
Anaptychia barbifera (Nyl.) Trevis., Flora 44: 52 (1861); Heterodermia barbifera (Nyl.) Kr.P. Singh, Bull. Bot. Surv. India 21:
221 (1981 [19 February]) [1979]. Type:—PERU. Near Tatanara, on branches of We in m a n n i a, August 1854, Lechler 2562
(holotype, PC n.v.).
Thallus foliose to subfruticose, forming small rosettes or irregularly spreading, adnate to loosely adnate, 2–5 cm
wide. Lobes 0.5–3.0 mm wide, sublinear, convex, irregularly branched, suberect or ascending at the apices,
partially imbricate, discrete to contiguous, ciliate, with whitish simple cilia 1–2 mm long scattered along the
margins, not mat-forming. Upper surface grayish white or gray, epruinose; soredia and isidia absent. Medulla
white. Lower surface ecorticate, arachnoid, ±canaliculate, whitish. Rhizines simple or irregularly branched,
whitish, 1–2 mm long. Apothecia common, substipitate to stipitate, 1–5 mm wide, subterminal on ascending lobes;
margin crenate or with well-developed squamules, eciliate; disc brown to dark brown, often white-pruinose.
Ascospores Polyblastidia-type, ellipsoidal, with 2 or 3 small sporoblastidia, 36–51 × 17–25 μm. Pycnidia common,
immersed, then becoming emergent, visible as black dots; conidia bacilliform, 4–5 × 1 μm.
Chemistry: Cortex K+ yellow, C–, KC–, P+ yellow; medulla K+ yellow then red, C–, P+ orange; yellow
pigment on lower surface K+ violet; containing atranorin (major), zeorin (major), 6α-acetoxyhopane-16β,22-diol
(minor), 6α-acetoxy-16β,22-dihydroxyhopane-25-oic acid (trace), leucotylin (trace), norstictic acid (major),
connorstictic acid (minor).
Nomenclature notes:Moberg & Nash (1999: 9) selected an epitype from Tanzania, Africa. However, TLC
studies of this material revealed that it lacks depsidones. This chemistry differs from that commonly accepted for
this species and for all six specimens collected from different places in Réunion—all contained norstictic and
connorstictic acids as major depsidones. Therefore, we designate a new epitype. Furthermore, we place H.
barbifera into synonymy, because all specimens from Réunion exhibit densely branches rhizines which form a mat
along the margins, the prime distinguishing character of H. barbifera according to Kurokawa (1962) . Only near
the tips of the lobes are the rhizines more or less simple. Although we have not seen the type of H. barbifera, we
have seen authentic material from Peru which was identified by Kurokawa.
Distribution and habitat:—Very rare on bark of trees (Castanopsis and Acer pseudoplatanus are reported) in
lower montane rainforest and lower montane scrub from 1125 to 1475 m above sea level; also in Asia, Australia,
North, Central and South America and Africa.
Remarks:characterized by the ascending or suberect lobes, the subterminal, substipitate or stipitate
apothecia, eciliate apothecial margins, the white, ecorticate lower surface, ascospores with sporoblastidia, the
presence of norstictic acid and the lack of japonene.
Material from Thailand examined:—Phitsanulok: Phu Hin Rong Kla National Park, area at air raid shelter,
in lower montane rainforest on bark of Castanopsis sp., 1125 m, 16°59'27'' N, 100°00'45'' E, 6 June 2003, P.
Mongkolsuk RU-PM 194, (RAMK 21443); ibid., P. Mongkolsuk RU-PM 204, (RAMK 21445); Phuluang Wildlife
Sanctuary, Forest protection unit office, Lon Tae substation, in lower montane scrub forest, on bark of Acer
pseudoplatanus, 1263 m, 17°14'18'' N, 101°33'19'' E, 12 November 2009, S. Meesim & K. Buaruang RU-MSPL
651, (RAMK 21146); ibid., S. Meesim & K. Buaruang RU-MSPL 648, (RAMK 21144); Loei: Phuluang Wildlife
Sanctuary, area of Kok Nok Kraba along trail to channel 7, in lower montane scrub, on bark of Castanopsis sp.,
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THE LICHEN FAMILY PHYSCIACEAE IN THAILAND—II
1470 m, 17°16'49'' N, 101°31'29'' E, 30 August 2005, W. Hongsachart WD 194, (RAMK 21205); Loei: Phuluang
Wildlife Sanctuary, walking trail to Pa Chang Parn, in lower montane scrub, on bark of unidentified tree, 1475 m,
17°16'51'' N, 101°31'10'' E, 24 June 2008, S. Meesim & K. Buaruang, RU-MS 14.4, (RAMK 21002).
8. Heterodermia stellata (Vain.) W.A. Weber, Mycotaxon 13: 102 (1981) (Fig. 12F)
Anaptychia podocarpa var. stellata Vain., Acta Soc. Fauna Fl. Fenn. 7(1): 131 (1890); A. stellata (Vain.) Kurok., Beih. Nova Hedwigia
6: 90 (1962). Type—Brazil. Minas Gerais; Sitio, E.A.Vainio [Lich. Bras. Exs. No. 1080]; (lectotype, designated here: TUR-Vain.
7938!).
Thallus foliose, forming small rosettes to irregularly spreading, adnate to loosely adnate, 2–5 cm wide. Lobes
0.5–3.0 mm wide, plane to convex, short, sublinear, irregularly branched; apices plane to suberect or ascending,
partially im