Tetramelas thiopolizus comb. nov. with a key to all known species of Tetramelas

Abstract

The study of the type and additional material of Buellia andicola, B. hypophana and B. reagens has shown that these taxa are conspecific with B. thiopoliza. Buellia thiopoliza is a muscicolous species growing at middle to high altitudes in the mountains of both hemispheres. It is easily recognized by its K+ purple-red yellow pigment mainly concentrated in the hypothecium, and by the presence of xanthones in the thallus and the apothecial tissues. The new combination Tetramelas thiopolizus is proposed. A key to all known Tetramelas species is included. The also muscicolous Buellia hypoleuca is reduced to synonymy with Phaeorrhiza sareptana var. sphaerocarpa.

Full text

Tetramelas thiopolizus comb. nov. with a key to all known
species of Tetramelas
Mireia GIRALT and Philippe CLERC
Abstract: The study of the type and additional material of Buellia andicola, B. hypophana and B.
reagens has shown that these taxa are conspecific with B. thiopoliza. Buellia thiopoliza is a muscicolous
species growing at middle to high altitudes in the mountains of both hemispheres. It is easily
recognized by its K+ purple-red yellow pigment mainly concentrated in the hypothecium, and by the
presence of xanthones in the thallus and the apothecial tissues. The new combination Tetramelas
thiopolizus is proposed. A key to all known Tetramelas species is included. The also muscicolous Buellia
hypoleuca is reduced to synonymy with Phaeorrhiza sareptana var. sphaerocarpa.
Key words: Buellia s. lat, Caliciaceae, Lecanoromycetes, lichenized fungi, Physciaceae, taxonomy,
terricolous lichens
Introduction
As part of a revision of the genus Buellia in
the Iberian Peninsula, several specimens pre-
viously identified as Buellia hypophana and B.
reagens have been studied. These are well
characterized by their xanthone content, by
the presence of a K+ purple-red yellow pig-
ment mainly located in the hypothecium
and the medulla below it, and by the rather
large, 1-septate ascospores with additional
pseudosepta when overmature. A thorough
literature search (e.g. Poelt 1969; Clauzade
& Roux 1985; Wirth 1995b; Marbach 2000;
Nordin 2000) revealed that, in addition to
the two species mentioned above, other taxa
might be involved in the Iberian flora, such
as B. andicola Müll. Arg. ex Zahlbr., B.
hypoleuca H. Magn. and B. thiopoliza (Nyl.)
Boistel. A study of the type material of these
taxa has shown that they are all conspecific
and well accommodated in the genus Tetra-
melas Norman, except B. hypoleuca which is a
synonym of Phaeorrhiza sareptana var. sphae-
orocarpa (Th. Fr.) H. Mayrhofer & Poelt.
A second aim of this study is to provide a
key for all known Tetramelas species, which
are so far treated in many dispersed contri-
butions (Marbach 2000; Kalb 2004; Nordin
2004; Nordin & Tibell 2005; Giralt et al.
2009).
Material and Methods
This study is based on herbarium material from G, H,
LISU, M, PO and from the private herbaria of J. Etayo
(Spain), K. Kalb (Germany) and P. P. G. van den Boom
(The Netherlands). Lichen morphology was examined
by standard techniques using stereo and compound
microscopes. Only free, mature ascospores lying outside
the asci have been measured. Measurements were made
on material mounted in water at ×1000 magnification.
Mean value (x
¯
) and standard deviation (SD) were
calculated and the results are given as (minimum value
observed) x
¯
± SD (maximum value observed). x
¯
,SD
and n(the total number of ascospores measured) are
given in parentheses. The terminology used here for the
asci follows Rambold et al. (1994), Nordin (2000) for
the excipulum-type, Nordin (1997) for the ascospore
septation and Giralt (2010) for the ascospore and
ontogeny-types.
Chemical constituents were identified by thin-layer
chromatography (e.g. Culberson & Ammann 1979;
Culberson et al. 1981; Culberson & Johnson 1982) and
high performance liquid chromatography (HPLC) (Elix
et al. 2003).
M. Giralt: Departament de Bioquı´mica i Biotecnologia
(Àrea de Botànica), Facultat d’Enologia de Tarragona,
Universitat Rovira i Virgili, Marcel·lı´ Domingo s/n,
43007, Tarragona, Spain. Email: mireia.giralt@urv.cat
P. Clerc: Conservatoire et jardin botaniques de la Ville
de Genève, Case postale 60, CH-1292 Chambésy (GE),
Switzerland.
The Lichenologist 43(5): 417–425 (2011) © British Lichen Society, 2011
doi:10.1017/S0024282911000363

The key to species has been constucted from our own
data and the following literature: Nordin (1999, 2000,
2004), Giralt et al. (2000, 2009), Marbach (2000),
Foucard et al. (2002), Kalb (2004), Nordin & Tibell
(2005) and Bungartz et al. (2007).
The Species
Tetramelas thiopolizus (Nyl.) Giralt &
Clerc comb. nov.
MycoBank No.: MB561208
Lecidea thiopoliza Nyl., Flora 56: 244 (1878); type:
France, Arvernia, sur les rochers qui dominent la
Bourboule prés du Mont Doré, Lamy, 1877 (H-NYL
10368!—lectotype, selected here; H-NYL 10367—
isolectotype!). Two additional specimens (H-Nyl—
10366! and G—290893!) could also be original material
but the collection date is 1878 or is lacking,
respectively).—Buellia thiopoliza (Nyl.) Boistel, Nouv.
Flore Lich.2: 230 & 233 (1903).
New synonyms: Buellia hypophana (Nyl.) Zahlbr.,
Cat. Lich. Univ.7: 367. (1931).—Lecidea hypophana
Nyl. apud Hue, Bull. Soc. Bot. France 34: 473 (1887);
type: France, Auvergne, Cantal, à la Roquevieille, 1886,
Fuzet (H-NYL 10648—holotype!).
Hypoflavia andicola (Müll. Arg. ex Zahlbr.) Marbach,
Bibliotheca Lichenologica 74: 292 (2000).—Buellia andi-
cola Müll. Arg. ex Zahlbr., Acta Horti Gothob.2: 25
(1925–1926); type: Chile, Anden, über Moosen, ex
Herbarium Hampe 1877 (G290798—holotype!).
Buellia reagens H. Magn., Broteria IV: 148 (1947);
type: Portugal, Beira Alta, Serra da Estrela-Caldas de
Manteigas, 850 m, sur le granite moussu, 1944, Tavares
(LISU).
(Figs1&2)
Thallus subsquamulose, areolate; areoles
discrete and bullate to usually contiguous,
sublobate, some almost lobate-effigurate,
plane to verrucose, whitish grey, ochraceous,
yellowish, yellowish orange to dull brown;
basal part of some areoles include a yellow
pigment reacting K+ purple-red. Medulla
normally I+ weakly violet (microscope slide!)
rarely I−.
Apothecia lecideine, adnate to sessile,
black, up to 1·25 mm diam., often confluent.
Proper margin initially thick and prominent,
becoming thinner and finally ±excluded.
Disc plane to convex, epruinose. Excipulum
proprium very well developed, up to 100 m
thick, including some C+ orange crystals
(microscope slide!), prosoplectenchyma-
tous, outer part brown, N−, inner part paler
F. 1.Tetramelas thiopolizus (G 290893), habitus. Scale = 500 m.
418 THE LICHENOLOGIST Vol. 43

(see Nordin 2000, Fig. 4A & B). Epihymenium
brown, N−, with C+ orange crystals (epip-
samma). Hymenium colourless, 60–100 m
high, without oil droplets, often including
rows of some C+ orange crystals. Hypothe-
cium dark brown, basal part with a yellow
pigment reacting K+ purple-red. Paraphyses,
apical cells strongly swollen, up to 7 m
diam., brown. Asci Bacidia-type. Ascospores
(14–)17–22·5(–27) × (6·5–)7·4–9·1(–11) m
[
x
¯= 20 × 8·2 m; SD = 2·5 × 0·8 m; n=
190], 1-septate, apices pointed, straight or
slightly curved, smooth at ×1000; when
young with subapical inner wall thickenings
(Callispora-type, Fig. 2B), when mature
without any inner wall thickening (Buellia-
type, Fig. 2C), some showing a pseudo-
septum or endospore septa (see Nordin
1997, Fig. 11) in each cell when overmature
(better seen in K) (Fig. 2D); ontogeny type C
(subapical inner wall thickenings are distinct
before the septum is developed, Fig. 2A).
Conidia bacilliform 4–6 × 1 m.
Chemistry.6-O-methylarthothelin, artho-
thelin and an unknown yellow pigment in
all specimens examined except in the two
known Andine specimens where arthothelin
is absent [corresponding to B. andicola (G-
holotype and hb. Kalb-29314)]. Low con-
centrations of several additional unknowns
have also sometimes been detected. Thallus
C+ pale to deep orange, K+ yellow-orange to
orange, KC+ orange to deep orange and
UV+ orange (the distinctness of these spot
tests varies in a single thallus and must be
dependent upon the concentration of xan-
thones); basal part of the hypothecium and
medulla below it always including the K+
purple-red yellow pigment; basal part of the
medulla of the areolae sometimes including
the K+ purple-red yellow pigment. This yel-
low pigment was tentatively identified by
HPLC as cinnamomeic acid B (cf. Paz-
Bermúdez & Giralt 2010).
Observations. Phylogenetic studies based
on genetic data have shown that the genus
Tetramelas Norman constitutes a well
founded segregate of Buellia s. lat. (Helms
et al. 2003; Nordin & Tibell 2005). The
genus was resurrected by Marbach (2000).
Later, it was emended with the addition of
new diagnostic characters: the presence of
6-O-methylarthothelin (Kalb 2004), the
relative thickness of the parts of the asco-
spore wall (proper wall >1/2 thickness of the
perispore) (Nordin 2004) and the inner wall
thickenings of the ascospores (±Callispora-
type) (Giralt et al. 2009). Since then, thirteen
species have been combined into this genus
and one new species has been described.
Buellia thiopoliza contains 6-O-methyl-
arthothelin and has ascospores with
Callispora-type thickenings. The spores are
often curved, have pointed apices and may
become tri-septate at maturity. The paraphy-
ses have strongly swollen apical cells and the
conidia are bacilliform. This set of charac-
ters fits well with the definition of the genus
F. 2.Tetramelas thiopolizus, ascospore ontogeny and
variability (hb. v. d. Boom 19535). A, immature non-
septate ascospore with subapical thickenings; B, young
Callispora-type ascospores; C, mature Buellia-type asco-
spores; D, overmature ascospore with a pseudoseptum
per cell. Scale = 10 m.
2011 Tetramlas thiopolizus—Giralt & Clerc 419

Tetramelas and therefore the new combina-
tion Tetramelas thiopolizus is proposed here.
According to Kalb (2004), in addition to
the presence of xanthones, other diagnostic
characters for Tetramelas include the typically
muscicolous habitat, the large ascospores
and the arctic-antarctic or alpine-subalpine
distribution. All these features are also
present in T. thiopolizus.
Tetramelas thiopolizus is thus characterized
by the subsquamulose-areolate thallus con-
taining 6-O-methylarthothelin and usually
arthothelin, the presence of a yellow K+
purple-red pigment mainly concentrated in
the basal part of the hypothecium, the thick
proper excipulum that is brown in the outer
part and paler in the inner part, and the
large, smooth, 1-septate ascospores showing
Callispora-type thickenings when young and
sometimes a pseudoseptum in each cell when
overmature.
Among all Tetramelas species known, T.
thiopolizus is clearly distinguished by the
presence of the yellow K+ purple-red pig-
ment. The occurrence of yellow pigments
(eumitrin and secalonic acid A) was already
reported in that genus in, for example, T.
graminicola (Øvstedal) Kalb. Concerning
cinnamomeic acids, they are present in the
medulla of Buellia capitis-regum W. A. Weber
(Nordin 2000).
The main differences observed between
the type material of B. andicola, B. hypophana
and B. thiopoliza, also noted in the corre-
sponding original descriptions, lay in the
ascospore size and in the intensity of the
yellow colour of the thallus, that is in
the intensity of the spot test reactions with
C, K and KC. In our opinion, both charac-
ters are insufficient to separate taxa since
large ascospores typically show a broad range
of variation and since the concentration of
xanthones might vary in the same thallus.
Buellia thiopoliza has a yellowish thallus
and contains arthothelin and 6-O-
methylarthothelin whereas B. andicola is
greyish to ochraceous and contains only
6-O-methylarthothelin. Buellia hypophana
(greyish to ochraceous) has not been exam-
ined chemically due to the very scanty type
material. Thalline reactions with C, K and
KC are stronger in B. thiopoliza than in B.
andicola and B. hypophana. Concerning the
ascospores, they are on average somewhat
shorter in B. andicola and in B. hypophana
than in B. thiopoliza but not significantly
according to the statistical calculations: B.
andicola: (14–)17·2–21·1(–23) × (7–)7·5–
9(–10·5) m[
x
¯=19 × 8·2 m; SD = 2 × 0·7;
n=42]; B. hypophana: (14·5–)17–22·3(–26)
× (6·5–)8–8·4(–11) [
x
¯= 19·6 × 8·4 m; SD
= 2·7 × 0·9 m; n=44]; B. thiopoliza: (14–)
17·9–23·5(–27) × (6·5–)7·2–8·7(–10) m
[
x
¯= 20·7 × 8 m; 2·8 × 0·8 m; n=48].
The type material of B. reagens (LISU) was
not available for study. However, according
to the original description (Tavares 1947), B.
reagens “seems exceedingly well character-
ized by the chemical reactions”. These reac-
tions are the “K+ intensely (violet) red”
hypothecium and the “K+ yellowish and C+
intensely orange yellowish-grey thallus”. The
measurements given for the ascospores are
“15–17(–20) × 8–9 m”. Material studied
here and mentioned in the protologue
(LISU), as well as the specimen collected
close to the type locality (hb. v. d. Boom-
17327), have ascospores (16–)17·8–22·5
(–25) × (6·5–)7·5–9·3(11) m[
x
¯= 20·2 ×
8·4 m; SD = 2·4 × 0·9 m; n=55].
Two features distinguish the Andine speci-
mens (corresponding to B. andicola) from
all other specimens examined. They lack
arthothelin and the I+ medullar violet reac-
tion. However other Tetramelas taxa may or
may not contain arthothelin in addition to
6-O-methylarthothelin [e.g. T. chloroleucus
(Körb.) A. Nordin, T. concinnus (Th. Fr.)
Giralt and T. insignis (Nägeli ex Hepp) Kalb,
see Giralt et al. 2000 and Bungartz et al.
2004, 2007]. Moreover, xanthones, especi-
ally when present in traces, are difficult to
detect with routine thin-layer chromatog-
raphy (TLC). HPLC is a far more sensitive
method of detection and identification for
this category of compounds. As with other
chemical data given for Tetramelas species,
those included here for B. andicola are based
only on TLC analyses. Concerning the
medullar reaction with iodine, it has been
shown to be variable in other species of
Tetramelas, as, for example, in T. concinnus
420 THE LICHENOLOGIST Vol. 43

(cf. Scheidegger 1993; Bungartz et al. 2004
and Giralt et al. 2009).
Except for the differences mentioned, all
specimens studied here are congruent in the
following characters: the ecology (on mosses
in alpine/sub-alpine/montane habitats), the
thalline morphology, the anatomy and pig-
mentation of the proper excipulum, the mor-
phology and ontogeny of the ascospores, the
presence of the yellow K+ purple-red pig-
ment in the hypothecium, the large apical
cells of the paraphyses and the shape and size
of the conidia.
It should be noted that Poelt (1969) had
already tentatively proposed to synonymize
B. hypophana with B. thiopoliza. He probably
did so because of the very similar original
descriptions.
The genus Hypoflavia Marbach was segre-
gated from Buellia s. lat. by Marbach (2000).
According to this author, the main diagnostic
character of the genus Hypoflavia is the pres-
ence of a yellow pigment in the hypothecium
and the excipulum which reacts K+ yellow-
orange or violet. Additional generic diagnos-
tic features are the occurrence of norstictic
acid and the presence of subapical wall thick-
enings in the ascospores (Callispora-type).
The genus includes two species, the generic
type Hypoflavia velloziae (Kalb) Marbach
and H. andicola (Müll. Arg. ex Zahlbr.)
Marbach. The former is known from three
Brazilian localities including the type; the
latter from the type locality in Chile and one
additional locality in Venezuela (hb. Kalb
29314).
According to our investigations, the two
known specimens of Hypoflavia andicola
have a yellow pigment in the thallus and
apothecia which reacts K+ purple-red but
instead of norstictic acid, as described by
Marbach (2000), they contain 6-O-
methylarthothelin. They fit well with the
generic concept of Tetramelas and indeed
correspond well to T. thiopolizus.
Surprisingly, the known specimens of Hy-
poflavia velloziae lack the yellow pigment in
the hypothecium and the excipulum consid-
ered by Marbach (2000) as the main diag-
nostic character for the genus Hypoflavia.
The K+ yellow-orange reaction described by
Marbach (op. cit.) for these apothecial parts
of H. velloziae is the typical reaction given
by norstictic acid before the formation of red
crystals. Furthermore H. velloziae contains
norstictic acid also in the thallus, lacks xan-
thones, has slightly enlarged apical cells of
the paraphyses [(2–)3–4 m diam.] and fusi-
form conidia (6–9 × 1–1·5m), a character
hitherto unknown for this species and which,
according to Marbach (2000), is diagnostic
for the genus Gassicurtia. All characters
mentioned above do not accommodate H.
velloziae in the genus Tetramelas. Conse-
quently, only H. velloziae is retained in the
genus Hypoflavia, which now includes only
this species.
Ecology and distribution. Tetramelas thiopoli-
zus grows on mosses or plant debris on rocks
or earth, at medium to high altitudes in
the mountains of both hemispheres. In the
Southern Hemisphere, it is known from the
Andes of Chile (type locality) and Venezuela
(Marbach 2000: sub Hypoflavia andicola). In
the Northern Hemisphere, it occurs in the
mountains of the Iberian Peninsula (Rico
1992; Terrón-Alfonso et al. 2000; van den
Boom & Jansen 2002; van den Boom 2003;
Pérez-Ortega & Álvarez-Lafuente 2006; Paz-
Bermúdez & Giralt 2010: sub B. hypophana;
and Tavares 1947: sub B. reagens); Austria
(Türk & Hafellner 1993, 1999: sub B. hy-
pophana); France (type localities of Lecidea
thiopoliza and L. hypophana); Germany
(Wirth 1995a: sub. B. thiopoliza); and Italy
(Nimis & Martellos 2008: sub B. hypophana).
According to Nimis & Martellos (op. cit.)
the species occurs also in Siberia (Russian
Federation).
Tetramelas thiopolizus also grows on mosses
at altitudes of 1600–1900 m in Macaronesia
from where it has been reported as B. hy-
pophana from two localities in Madeira by
Kalb & Hafellner (1992) and from one local-
ity in Gran Canaria (Canary Islands) by
Hafellner (1995).
The records of B. hypophana by Etayo
(1990a, b) from the western Pyrenees
(Navarra, Spain) refer to Tetramelas chloroleucus
and those by Beauchamp et al. (2007) from
Switzerland (Valais) with filiform conidia,
2011 Tetramlas thiopolizus—Giralt & Clerc 421

probably to Buellia punctata f. muscicola Hepp
ex Koerb. em Arnold (cf. Poelt 1969: 141),
because of the small ascospores and the
muscicolous habitat.
Additional specimens examined.Portugal: Minho:
Castro Laboreiro, musgos do penedo no castelo, 1915,
Sampaio (487L, PO) (sub Buellia insignis var. geophila).
Beira Alta: Serra da Estrela-Sra. do Desterro, Cabeça da
Velha, sobre o granito musgoso, 950 m, 1946, Tavares
(LISU); Serra da Estrela-Sra. do Desterro, sobre os
blocos granı´ticos de um pinhal, sobre o granito mus-
goso, 800 m, 1945, Tavares (LISU-71); Serra da
Estrela-Entre as Lagoas Escurra e Comprida, sobre
Andreaea em granito, c. 1600 m, 1947, Tavares (LISU-
2223); Serra da Estrela, SW of Manteigas, NE of Lagoa
Comprida, Lagoa Redonda, N exposed steep granite,
1625 m, 1995, van den Boom (hb. v. d. Boom-17327)
[sub. B. reagens H. Magn. (= B. pulchella C. Tav. non
Tuck.)]. Trás-Os-Montes: N of Bragança, Montesinho,
path to storage lake with Pinus and granite outcrops,
small Quercus wood and Betula near source, sheltered S
exposed granite ± horizontal, 6°45·2#W; 41°56·5#N,
1200 m, 7 viii 1997, van den Boom (hb. v. d. Boom-
19535).— Spain: Extremadura: 41 km ENE of
Plasencia, N of Jarandilla, 1–3 km N of Guijo de Santa
Bárbara, along path E of river Jaranda, acid rock on SW
slope, 1400 m, 1989, van den Boom (hb. v. d. Boom-
8880) [sub B. hypophana].—Venezuela: Mérida: Liber-
tador, Pico Espejo, SE von Mérida, Loma Redonda,
8°35#N; 71°00#W, 4200 m, 1989, Kalb & Kalb 29314
(hb. Kalb) (sub B. andicola).
Specimens of Hypoflavia velloziae studied for compari-
son.Brazil: Bahia: Chapada diamantina, Serra do
Tombador, etwa 1 km vor der Stadt Morro do Chapeú,
1000 m, an Vellozia, 1980, Kalb (K. Kalb: Lichenes
Neotropici nº 363, holotype, hb. Kalb); zwischen Mundo
Novo und Morro Chapeú, 1000 m, 1980, Kalb 28758
(hb. Kalb). Minas Gerais: Serra do Espinhaço, Serra do
Caraça, umbegung des Klosters Caraça (Hauptsam-
melgebiet von Vainio), 1978, Kalb & Plöbst 28952 (hb.
Kalb); NW-Hangdes Berges Carapuça, Groutas do
P. Trombert, in feuchtem, dunklem Primärwals, 1978,
Kalb & Plöbst 28730 (hb. Kalb).
Phaeorrhiza sareptana (Tomin) H.
Mayrhofer & Poelt var. sphaerocarpa
(Th. Fr.) H. Mayrhofer & Poelt
Nova Hedwigia 30: 793 (1979) [1978].
New synonym:Buellia hypoleuca H. Magn., Bot. Not.
109: 150 (1956); type: Austria, Südtirol, Wintschgau,
trockene Felsen oberhalb Graun am Reschensee, on
earth, 1955, J. Poelt (M—holotype!).
For additional data on this taxon see
Magnusson (1956, sub B. hypoleuca),
Mayrhofer & Poelt (1978) and Mayrhofer &
Moberg (2002).
Key to all known species of Tetramelas
1 Lichenicolous, on foliose Physciaceae; thallus absent (endokapilic) ........2
Not lichenicolous or, if lichenicolous, thallus present ...............3
2(1) Ascospores 3-septate........T. pulverulentus (Anzi) A. Nordin & Tibell
Ascospores 1-septate............T. phaeophysciae A. Nordin & Tibell
3(1) Thallus sorediate; ascospores (1–)3-septate, 24–40 × 7–11·5 m; containing 6-
O-methylarthothelin and traces of the anthraquinones eumitrin and secalonic acid
A; on plant debris; known only from South Georgia ................
.........................T. graminicola (Øvstedal) Kalb
Thallus not sorediate; ascospores smaller; eumitrin and secalonic acid A always
lacking .......................................4
4(3) Hypothecium with a yellow pigment reacting K+ purple-red. Ascospores 1- septate,
(14–)17–22·5(–27) × (6·5–)7·4–9·1(–11) m, when overmature showing some-
times an additional pseudoseptum per cell; on mosses growing on rock or ground;
mountains of both hemispheres..... T. thiopolizus (Nyl.) Giralt & Clerc
Hypothecium without yellow pigment, K− .....................5
5(4) Thallus without xanthones C−, KC−, UV−, white and chalky containing only
atranorin (K+ yellow); ascospores 1-septate, 15–25 × 7–10 m; muscicolous and
saxicolous; mountains of northern Europe . . T. papillatus (Sommerf.) Kalb
Thallus always containing xanthones (C+ or KC+ or UV+ orange), not white and
chalky; atranorin present or not..........................6
422 THE LICHENOLOGIST Vol. 43

6(5) Apothecia with norstictic acid, K+ red (forming crystals); ascospores (1–)3- septate,
26–34 × 8–10 m; usually on calciferous ground, rarely lignicolous; arctic-
alpine ......................T. terricola (A. Nordin) Kalb
Apothecia without norstictic acid, K− (no red crystals) ..............7
7(6) Saxicolous ......................................8
Corticolous, lignicolous, terricolous, muscicolous or on plant debris .......9
8(7) Ascospores 1(–3)-septate, 13–21 × 6–10 m; thallus with arthothelin and often 6-O-
methylarthothelin; Northern Hemisphere . . . T. concinnus (Th. Fr.) Giralt
Ascospores (1–)3-septate, 17–28 × 8–11·5 m; thallus only with 6-O-
methylarthothelin; Southern Hemisphere (antarctic) ...............
........................T. granulosus (Darb.) A. Nordin
9(7) Corticolous or lignicolous .............................10
Terricolous, muscicolous or on plant debris ...................14
10(9) Ascospores 3-septate ................................ 11
Ascospores 1-septate, rarely 1(-3)-septate .................... 12
11(10)Ascospores 19–27·5 × 7·5–10·5m; thallus only with arthothelin; subalpine; North-
ern Hemisphere ........T. triphragmoides (Anzi) A. Nordin & Tibell
Ascospores 23–38 × 6–13 m; thallus only with 6-O-methylarthothelin; arctic-
(sub)alpine; Northern Hemisphere .... T. geophilus (Sommerf.) Norman
12(10)Ascospores 13–23 × 6–11 m; thallus with 6-O-methylarthothelin and rarely traces
of arthothelin; alpine-subalpine; Northern Hemisphere ..............
.......................T. chloroleucus (Körb.) A. Nordin
Ascospores larger, > 23 m............................13
13(12)Ascospores 1-septate, 27–31 × 10–11 m; thallus with 6-O-methylarthothelin,
atranorin and lobaric acid; high tropical mountains ................
...........................T. regiomontanus Marbach
Ascospores 1(-3) septate, 23–32 × 9–13 m; thallus with 6-O-methylarthothelin and
rarely traces of arthothelin; high European mountains...............
........................T. insignis (Nägeli ex Hepp) Kalb
14(9) Ascospores 3-septate ................................ 15
Ascospores 1-septate or 1(-3)-septate .......................16
15(14)Ascospores 26–34 × 8–10 m; apothecia containing norstictic acid (K+ red crys-
tals); thallus with atranorin, arthothelin and rarely 6-O-methylarthothelin;
arctic-alpine ..............................T. terricola
Ascospores 23–38 × 6–13 m; apothecia without norstictic acid; thallus containing
only 6-O-methylarthothelin; arctic-(sub)alpine ..........T. geophilus
16(14)Ascospores 13–25 × 5–7·5 m; thallus containing atranorin and 6-O-
methylarthothelin; known only from New Zealand . . T. confusus A. Nordin
Ascospores larger, > 25 × 7·5 m.........................17
17(16)Ascospores 1-septate, 27–31 × 10–11 m; thallus with 6-O-methylarthothelin, atra-
norin and lobaric acid; high tropical mountains .......T. regiomontanus
Ascospores 1(-3)-septate, 23–32 × 9–13 m; thallus with 6-O-methylarthothelin
and rarely arthothelin; high European mountains .......... T. insignis
The authors are indebted to the keepers of G, H, LISU,
MandPOandtoJ.Etayo,K.KalbandP.P.G.vanden
Boom for the loan of herbarium material used in this
study and to J. A. Elix for carrying out the HPLC
analyses and giving additional comments on the
chemistry. The first author is grateful to the Spanish
Government for funding of the project CGL2007-
66734-C03-02/BOS.
2011 Tetramlas thiopolizus—Giralt & Clerc 423

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