Lichenologist 32(2): 105–128 (2000)
doi:10.1006/lich.1999.0251
Available online at http://www.idealibrary.com on
NOTES ON SOME CORTICOLOUS AND
LIGNICOLOUS BUELLIA SPECIES FROM THE
IBERIAN PENINSULA
Mireia GIRALT*, Mercedes BARBERO* and John A. ELIX‡
Abstract: The morphology, anatomy, chemistry and ecology of some corticolous
and lignicolous species of Buellia from the lberian Peninsula that contain secondary
lichen compounds are discussed, including B. chloroleuca, B. disciformis, B. erubescens, B. hyperbolica and B. leptoclinoides. These species are compared with other
corticolous Buellia species that occur outside the study area, including B. arnoldii,
B. insignis, B. regularis and B. sanguinolenta. The misunderstood B. zahlbruckneri
J. Steiner and B. jorgei, are synonyms of B. erubescens, and the invalid taxon
B. zahlbruckneri sensu Schauer non Steiner and B. poeltii, are synonyms of
B. chloroleuca. New chemical data on several taxa are reported. A key to the species
and illustrations of their ascospores are provided. 2000 The British Lichen Society
Introduction
The present study is a further contribution towards extending the knowledge
of the Physciaceae in the lberian Peninsula. The aim of this investigation was
to reappraise all those corticolous and lignicolous Buellia species that contain
secondary lichen substances. Although the emphasis here is on the lberian
species, many of these taxa are also found elsewhere in Europe. Furthermore,
several species that do not occur in the study area have also been investigated
for comparative purposes.
The corticolous and lignicolous species that have vegetative propagules,
polyspored asci or triseptate ascospores, as well as secondary metabolites, are
not included in this paper but will be treated in future studies. Furthermore,
several of the specimens examined that have 8-spored asci and secondary
chemistry like the species discussed in this paper will be described as new in
a forthcoming paper.
It should be pointed out that this work should be regarded as a first step
towards a better understanding of these corticolous/lignicolous Buellia species.
It has been undertaken for a small part of the world and it is based on a
relatively small number of specimens. For this reason, we have not attempted
to examine all type specimens and until a world-wide taxonomic revision of
the genus Buellia is carried out many problems will remain unresolved.
*Department de Biologia Vegetal (Botànica), Facultat de Biologia, Universitat de Barcelona,
Diagonal 645, 08028 Barcelona, Spain.
‡Department of Chemistry, Faculty of Sciences, The Australian National University, Canberra
ACT 0200, Australia.
0024–2829/00/020105+24 r35.00/0
2000 The British Lichen Society
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In the present treatment all taxa are retained in the genus Buellia pending
further data to confirm whether the type species of the genus Buellia is B.
parasema de Not., as proposed by Kalb & Elix (1998), or B. aethalea (Ach.)
Th. Fr., as more recently proposed by Moberg et al. (1998).
Characters of the species
A comprehensive account of the comparative morphology and anatomy of the
Buellia species studied is not included. However, the taxonomic value of some
characters requires discussion.
According to our observations, the characters of greatest taxonomic value in
the genus Buellia are the development and structure of the ascospores, the
presence of hymenial oil droplets, the degree of development of the excipulum
and its reaction with K, and the presence of particular lichen compounds.
Other characters such as thalline and apothecial morphology and the degree of
swelling of the apical cells of the paraphyses are important features but of little
diagnostic significance.
The ascospores of the species treated in this paper are of the following types:
the Callispora-type or similar (compare Kalb 1986; Mayrhofer 1984;
Scheidegger 1993; Pusswald et al. 1994), with the inner ascospore wall
thickened subapically and at the septum (B. arnoldii, B. disciformis, B.
leptoclinoides, B. regularis and B. sanguinolenta). In addition, two species have
ascospores that are intermediate between the Callispora- and the Buellia-types
(B. chloroleuca and B. insignis). Finally, B. erubescens and B. hyperbolica possess
ascospores that are intermediate between the Physconia- and the Buellia-type
(see Scheidegger 1993), with the inner ascospore wall thickened at the
septum in the early stages of their ontogeny but lacking any wall thickening
when mature.
All the present species have ascospores that originate via a type A ontogeny:
in young and hyaline ascospores the septum is inserted before apical internal
wall-thickenings become distinct (cf. Giralt 1994; Giralt & Mayrhofer 1994,
1995).
Two types of ascospore ornamentation can easily be recognized with light
microscopy, at least after addition of 10% KOH: the weakly and the roughly
microrugulate ornamentation [type 1 and 2 of Nordin (1996), respectively].
The spore ornamentation is weak in B. leptoclinoides, rough in B. chloroleuca,
B. erubescens, B. hyperbolica and B. insignis and smooth in B. disciformis and in
the B. arnoldii-group.
The thallus morphology of all the species treated is rather variable and may
range from thin and continuous to thick and rimose. Well-developed thalli and
that of lignicolous specimens may often develop a granulose to verrucose
surface. The prothallus is especially visible when adjoining thalli of other
species.
The apothecia are lecideine in all the species treated except in B. hyperbolica
in which they very soon become biatorine. The excipulum is more or less well
developed in all the species except in B. hyperbolica where it is very thin. The
brownish pigmentation gives a yellow to yellow-brownish solution with K in
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107
B. chloroleuca, B. erubescens, B. hyperbolica and B. insignis (compare Scheidegger
1993 and Nordin 1996).
The hymenium is interspersed with numerous oil droplets (opaque even in
thin sections) in B. arnoldii, B. disciformis, B. leptoclinoides, B. regularis and
B. sanguinolenta, but oil droplets are absent in the other species treated.
The asci are 8-spored, clavate and of the typical Bacidia-type (Rambold
et al. 1994) in all the present species.
Spermogonia are rare and have been found only in B. chloroleuca, B.
disciformis and B. erubescens. The spermatia are always hyaline, bacilliform,
straight, single-celled and thin walled, of 4·5–9 m long and 0·8–1 m wide.
The spermatophores are branched, resembling the Type VI of Vobis &
Hawksworth (1981).
The species treated in this paper are characterized by the presence of various
secondary lichen compounds, which can be detected after the application of
reagents (spot tests) and include: atranorin, fumarprotocetraric acid, protocetraric acid, 6-O-methylarthothelin (=granulosin) and norstictic acid (with
connorstictic acid). Atranorin reacts K+yellow and it is present (at least in
traces) in all the species treated; fumarprotocetraric and protocetraric acids
react PD+orange-red and are present in B. hyperbolica; 6-O-methylarthothelin
reacts K+and C+yellowish and KC+orange and is present in B. chloroleuca
and B. insignis, whereas norstictic acid reacts K+red (with the formation
of red crystals under the microscope) and is present in B. erubescens and
B. sanguinolenta. All these colour reactions are usually strong and are visible
under the stereomicroscope.
Other secondary lichen compounds present in the species treated in this
paper (e.g. zeorin) that do not give spot test reactions or are present only in
traces (e.g. gyrophoric acid) must be detected by thin layer chromatography
(TLC) or high performance liquid chromatography (HPLC).
The following lichen compounds have been detected for the first time in the
genus Buellia: protocetraric, confumarprotocetraric, convirensic and hyposalazinic acids, and fulgoicin. Further, an unknown present in B. disciformis
(tentatively identified as norfulgoicin) and an unknown secalonic acid derivative present in B. erubescens [also detected in B. multispora and Gassicurtia
coccinea (Kalb & Elix 1998)] have yet to be fully characterized chemically.
For the most part, notes on the ecology and distribution of the species refer
only to the study area.
Materials and Methods
All the morphological descriptions are based on our personal examinations. Current mycological
terminology has been employed and generally follows that of Hawksworth et al. (1995). The
terminology used for the asci follows that of Rambold et al. (1994), while for the ascospore-types
the terminology of Kalb (1986), Mayrhofer (1982, 1984), Scheidegger (1993), Giralt (1994) and
Giralt & Mayrhofer (1994, 1995) is followed. The standardized methods of TLC (e.g. Culberson
& Ammann 1979; Culberson et al. 1981; Culberson & Johnson 1982) and HPLC (Feige et al.
1993) for the identification of lichen substances were employed.
As most of the species considered have previously been described in detail in other revisions,
including those of Sheard (1964), Schauer (1965) and Orange et al. (1992), the descriptions given
here emphasize characters of value for species identification but do not repeat features common
to all of the species.
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Key to the species
1
Thallus K+yellow-orange then red; containing norstictic acid . . 2
Thallus K+yellow, yellowish or yellowish brown, never K+yelloworange then red; norstictic acid absent . . . . . . . . . . . . . 3
2(1)
Hymenium with numerous oil droplets. Excipulum K. Ascospores
(21–)26–32(–35)(11–)12–13(–16) m, of the Callispora-type
. . . . . . . . . . . . . . . . . . . . . . Buellia sanguinolenta
Hymenium without numerous oil droplets. Excipulum K+yellow
(microscope slide). Ascospores (11–)13–16(–18)(6–)7–8·5
(–9·5) m, Physconia-type when young, then Buellia-type when
mature . . . . . . . . . . . Buellia erubescens (Chemotype I)
3(1)
Thallus PD+orange-red; fumarprotocetraric and protocetraric acids
present. Apothecia rapidly becoming biatorine, 0·2–0·3(–0·4) mm
diam. . . . . . . . . . . . . . . . . . . . . Buellia hyperbolica
Thallus PD; fumarprotocetraric and protocetraric acids absent.
Apothecia rarely becoming biatorine, significantly larger . . . . 4
4(3)
Hymenium with numerous oil droplets . . . . . . . . . . . . . . 5
Hymenium without numerous oil droplets . . . . . . . . . . . . . 7
5(4)
Ascospores 15–20(–23)(6·5–)8–9(–9·5) m, placodiolic acid
present . . . . . . . . . . . . . . . . . . Buellia leptoclinoides
Ascospores significantly larger, placodiolic acid absent . . . . . . 6
6(5)
Ascospores (23–)26–29(–31)(10–)12–14(–16·5) m, thallus containing only atranorin . . . . Buellia arnoldii (incl. B. regularis)
Ascospores (16–)19–25(–28)7·5–9(–11) m, thallus containing
atranorin, fulgidin and fulgoicin . . . . . . Buellia disciformis
7(4)
Thallus KC+and UV+orange, section C+orange (microscope
slide), containing 6-O-methylarthothelin. Ascospores intermediate
between the Callispora and the Buellia-type . . . . . . . . . . . 8
Thallus KCand UV, section C, 6-O-methylarthothelin
absent. Thallus K+yellow, containing atranorin and traces (not
always) of several homologues of norstictic acid. Ascospores
Physconia-type when young, then Buellia-type when mature, (11–)
13–16(–18)(6–)7–8·5(–9·5) m
. . . . . . . . . . . . . . . Buellia erubescens (Chemotype II)
8(7)
Ascospores (19–)21–25(–29)(8–)9–11(–12) m
. . . . . . . . . . . . . . . . . . . . . . . . . . Buellia insignis
Ascospores smaller, (13–)16–19(–22)(6–)7–9 m
. . . . . . . . . . . . . . . . . . . . . . . . Buellia chloroleuca
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F. 1. Ascospore ontogeny. A, Buellia choroleuca (Lichenes Alpium 192, W); B, B. insignis
(G—lectotype). Buellia-type ascospores showing tendencies to the Callispora-type (arrows).
Scale=10 m.
The Species
Buellia chloroleuca Körb.
Parerg. Lich.: 191 (1860); type: Polen: Schlesien, Sudeten, an Fichten, Körber (L!, M!—isotypes?).
New synonyms: Buellia poeltii T. Schauer, Mitt. Bot. München 5: 616 (1965); type: Germany:
Oberbayern, Lahnenwies-Graben prope Garmisch, in cortice Aceris pseudoplatani, 1470 m,
4 September 1963, Th. Schauer (M!—holotype).
(Fig. 1A)
Thallus whitish, whitish grey or cream, rarely thin and smooth to rimose,
usually moderately thick, warted, continuous and well developed, covering
large areas, not delimited by a blackish prothallus. Apothecia (0·3–)0·5–0·8
(–1) mm diam., abundant and confluent, adnate, rarely becoming sessile. Disc
initially plane, rapidly subconvex to strongly convex, epruinose. Proper
margin initially thick, becoming thinner and finally excluded. Excipulum
60–80 m wide, K+deep yellow-brownish (microscope slide). Hymenium
without oil droplets, 80–100 m high. Hypothecium interspersed with oil
droplets. Apical cells of the paraphyses strongly swollen, 4–6(–7) m wide,
brown pigmented. Ascospores (13–)16–19(–22)(6–)7–9 m, when young
showing tendencies to the Callispora-type, when mature of the Buellia-type,
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apices pointed, straight to slightly curved, roughly ornamented. Spermatia
bacilliform, straight, 4·5–5·5(–6)1 m.
Chemistry. Thallus UV+orange, K+and C+yellowish, KC+orange; thallus
section C+orange. Containing 6-O-methylarthothelin (=granulosin) (major),
arthothelin (minor) and atranorin (traces) by TLC and HPLC. Traces of
several additional unidentified substances have been detected in some of the
specimens analysed by TLC. Depending on the concentration of 6-Omethylarthothelin (with arthothelin), the spot tests mentioned above are
more or less clearly visible. According to Wirth (1988), Buellia poeltii (=B.
chloroleuca) contains placodiolic acid. This report is not in agreement with our
TLC and HPLC analyses.
The xanthone 6-O-methylarthothelin has previously been reported for this
species (as B. zahlbruckneri) by Huneck et al. (1992) and by Kalb & Elix
(1998). Within the genus Buellia, it is known also from B. granulosa (Leuckert
& Mathey 1975), B. insignis, B. melanochlora, B. mediospora and B. parasema
(Kalb & Elix 1998).
Taxonomic remarks. Schauer (1965) incorrectly neotypified Buellia zahlbruckneri J. Steiner and he chose as neotype the specimen Arnold: Lich. Exs.
1589 from M. The specimens of Arnold: Lich. Exs. 1589 (DUKE, FH, UPS)
investigated (invalid ‘isoneotypes’) belong to B. chloroleuca. Thus, the invalid
taxon B. zahlbruckneri in the sense of Schauer (1965) is a ‘synonym’ of
B. chloroleuca.
See also the taxonomic remarks under B. erubescens.
Discussion. Buellia chloroleuca is characterized by the presence of 6-Omethylarthothelin. Other distinguishing characters of this species include the
typically thick and warted thallus, which usually covers wide areas, the
abundant, confluent and adnate apothecia, the K+deep yellow-brownish
reaction of the rather well developed excipulum, the strongly swollen apical
cells of the paraphyses and the relatively small Buellia-type ascospores, which
show Callispora-type tendencies. Among the species treated, B. chloroleuca is
closely related to the corticolous/muscicolous B. insignis, which also contains
the xanthone 6-O-methylarthothelin and has similar morphological characters
(a rather thick and warted thallus, adnate and abundant apothecia, lack of
hymenial oil droplets, large apical cells of the paraphyses, etc). However, the
latter species is well separated by the larger ascospores (see comments under
B. insignis).
As indicated above (see Chemistry), the intensity of the spot test with UV,
K, C and KC may vary depending on the concentration of 6-Omethylarthothelin in the thallus. When these test reactions are doubful, B.
chloroleuca could be confused with the chemotype of B. erubescens, which lacks
or has low concentrations of norstictic acid. Apart from TLC analyses (which
give a rapid solution), the following morphological characters should be
carefully studied: the degree of development of thallus, the size of apothecia,
the thickness of the excipulum and its reaction with K, the size of the apical
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Iberian Buellia—Giralt et al.
111
cells of the paraphyses, the size of the ascospores and the ascospore-ontogeny
(see comments under this taxon).
Buellia poeltii was first described by Schauer (1965). The holotype and other
specimens examined give a K+pale yellow thalline reaction, which was
considered by Schauer (op. cit.) to be negative. For this reason, he compared
this species with other Buellia taxa that lack secondary lichen compounds,
such as Amandinea punctata (as B. punctata) and B. schaereri, rather than with
species with K+yellow thalli, such as B. chloroleuca or B. disciformis. Consequently B. poeltii was described as new and distinguished from A. punctata and
B. schaereri on the basis of its larger ascospores. Our morphological and
chemical studies on several isotypes and one additional specimen of B.
chloroleuca established that they differ from specimens of B. poeltii (holotype
included) only by their more granulose thalli, a character that we consider
of little taxonomic value. For this reason we synonymize B. poeltii with
B. chloroleuca.
The same thalline character separates the ‘isoneotypes’ and other specimens
of the invalid taxon B. zahlbruckneri sensu Schauer examined from B.
chloroleuca. But as Schauer’s neotypification of B. zahlbruckneri is totally
invalid no synonymy between B. chloroleuca and B. zahlbruckneri sensu
Schauer must be considered (more details are given under the taxonomic
remarks on B. erubescens).
Ecology and distribution. In the study area B. chloroleuca occurs at high
altitudes in the Pyrenees. Overall, the European distribution of B. chloroleuca
is poorly known due to confusion of this species with B. zahlbruckneri sensu
Schauer. Nevertheless, it seems to be a montane to alpine species, rather
common in the Alps where it occurs mainly on lignum. Many records of
B. zahlbruckneri growing on lignum may refer to this species. In the area
studied, only the record in Etayo (1989b) corresponds to B. chloroleuca. The
specimen reporded as B. zahlbruckneri in Boqueras (1993) could also belong
to B. chloroleuca but it is so severely parasitized by the lichenicolous fungus
Lichenodiplis lecanorae that its identity could not be confirmed. Other Iberian
records of B. zahlbruckneri checked refer to other Buellia taxa but not to
B. chloroleuca [e.g. to B. erubescens. Etayo (1989a) or to Buellia sp., Marcos
(1985)].
Exsiccatae examined: Anzi, Lich. exs. minus rari Ital. sup. 296, 297, 299 (M, as Buellia parasema
v. saprophila). Hepp, Fl. Eur. 150 (M, as Lecidea punctata v. saprophila). Rabenhorst, Lich. Eur.
729 (M, as Buellia parasema v. saprophila). Roumeguère, Lich. Gall. exs. 191 (M, as Lecidea
disciformis f. ecrustacea). Schaerer, Lich. Helv. exs. ed. I. 198 (M, as Lecidea punctata v. saprophila).
Vězda, Lich. sel. exs. 897 (W, as B. zahlbruckneri) and 1793 (as Buellia disciformis, BCC). Lichenes
Alpinum 192 (W, M, as B. zahlbruckneri).
Selected additional specimens examined: Spain:. Navarra: Larra, 1800 m, on dry twigs of Pinus
uncinata, J. Etayo (hb. Etayo).—Austria: Carinthia: Turracher Höhe ’supra lacum Scwarzsee,
1840–1900 m, ad lignum et corticem Pini cembrae’, 1985, A. Vězda (MUB-6999); Nockgebiet,
Huberalpe, östl. St. Lorenzen bei Ebene Reichenau, on lignum, 1984, S. Wagner (KL); Gailtaler
Alpen, Kerschdorfer Alm (östl. der Windischen Höhe), on lignum, 19 vi 1984, S. Wagner
(KL).—France: Alpes Maritimes: oberh. Casterine, westl. St. Dalmas de Tende, 1600 m, an
Larix, 1963, Th. Schauer (M).—Germany: Oberbayern: Wildsee am Krottenkopf bei Garmish,
1400 m, an Sorbus aucuparia, 1962, Th. Schauer (M); Sägertalbach bei Linderhof, 1400 m,
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an Fichtenstumpf, 1962, Th. Schauer (M).—Italy: Südtirol, Belluno, Pinus cembra-Äste mi
Valgrande nahe Schluderbach in den Ampezzaner Alpen, 1874, F. Arnold (M); Bozen, P.
cembra-Strunk mi Damersthale auf dem Shlem, 1867, F. Arnold (M); Trento, Dolomiten, Dürre
Fichten-Zweige bei Paneveggio, 1878, F. Arnold (M).—In Herbarium F. Arnold: locality not
indicated (M).
Buellia disciformis (Fr.) Mudd
Manual Brit. Lich.: 216 (1861)
Lecidea parasema de Not. var. disciformis Fr., Nov. Sched. Critic.: 9 (1826); type: Fries, Lich.
Suec. Exs. 215A (UPS!—lectotype).
(Fig. 2A)
Thallus whitish to whitish grey, varying from thin and smooth to rather thick
and cracked-areolate, continuous, often delimited by a blackish prothallus.
Apothecia (0·5–)0·7–1·1(1·3) mm diam., usually scattered, adnate to sessile.
Disc plane to subconvex, rarely strongly convex, epruinose. Proper margin
initially thick, entire to flexuose, prominent, becoming thinner, usually
persistent. Excipulum (50–)60–80(–100) m wide, K(microscope slide).
Hymenium with numerous oil droplets, 100–120 m high. Apical cells of
the paraphyses slightly swollen, 2·5–3·5(–4·5) m wide, brown pigmented.
Ascospores (16–)19–25(–28)7·5–9(–11) m, similar to the Callispora-type,
with inner wall clearly thickened subapically, but not or only slightly thickened
at septum, often curved and becoming triseptate, smooth. Spermatia elongate
bacilliform, straight, 5·5–7·5(–9)0·8–1 m.
Chemistry. Thallus K+yellow. Atranorin (major) and usually traces of
fulgidin (minor), fulgoicin (minor or trace), and of an additional unknown
lichen substance () (tentatively identified as norfulgoicin), by TLC and
HPLC. Fulgoicin is reported for the first time from the genus Buellia. Fulgidin
is also known from B. parasema de Not., the type species of the genus
proposed by Kalb & Elix (1998).
Discussion. The presence of atranorin, fulgidin and fulgoicin together with
the large apothecia, the hymenial oil droplets and the near Callispora-type
ascospores are diagnostic characters of B. disciformis. Morphologically B.
disciformis comes close to B. leptoclinoides but is clearly distinguished chemically. Further distinguishing features of B. leptoclinoides (absent in B. disciformis) include the thicker, often yellowish thallus due to the presence of
placodiolic acid, the more crowded and convex apothecia possessing large
crystals in the outer part of the excipulum, and the significantly smaller
ascospores. These taxa also exhibit ecological differences.
Buellia arnoldii is also similar morphologically but it contains only atranorin
and has larger ascospores (see also the discussion under that species).
Buellia disciformis is a relatively uniform species despite the fact that the size
and septation of the ascospores may vary. Whereas the average spore length in
the type material is 21 m, it can reach 25 m in other specimens examined.
The specimens that possess larger ascospores are also those that contain a
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Iberian Buellia—Giralt et al.
113
F. 2. Ascospore ontogeny. A, Buellia disciformis (Spain, Las Batuecas, 1989, Marcos, SALAF1577); B, B. leptoclinoides (Portugal, 14 km of Lagos, 1993, P. v. d. Boom, Boom 14670). Typical
Callispora-type ascospores. Scale =10 m.
higher number of triseptate ascospores. Several records of B. lauricassiae from
the study area refer to this morphotype of B. disciformis.
Buellia parasema de Not. resembles B. disciformis in having a hymenium
inspersed with oil drops and callisporoid ascospores with tendencies to
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become four-celled. This species is mainly distinguished from B. disciformis
by its alternative chemistry since it contains atranorin, as the major substance, and norstictic acid, dissectic acid, methyl -orsellinate, 6-Omethylarthothelin, brialmontin 1, brialmontin 2 and fulgidin, as minor
substances (cf. Kalb & Elix 1998).
Ecology and distribution: Buellia disciformis seems to be a rather common
species, widely distributed in Portugal and in northern Spain except for coastal
localities. It grows from low to middle altitudes, mainly on the smooth bark
of deciduous trees. In the Mediterranean area of the Iberian Peninsula, B.
disciformis is restricted to the montane forests of Fagus and has been reported
many times from the study area.
Additional specimens examined. Portugal: Algarve: Serra de Monchique, road Casais to
Marmelete near Rua Nova, 400 m, on Quercus suber, 1993, P.v.d. Boom, (hb. Boom 14621).
Alentejo: SSW of Évora, SE of Alvito, S side of Barragem Odivelas, 100 m, on Q. ilex, 1995, P.v.d.
Boom (hb. Boom 17185). Beira Litoral: 8 km SW of Oliveira do Hospital, Venda da Galizes to
Ponte das Tres Entradas, 350 m, on Pinus, 1993, P.v.d. Boom (hb. Boom 14505). Minho: Serra
do Gerez-Pedra Bela, on Betula, 1947, C. Tavares 2014, 2023 (LISU, UPS); Serra do
Gerez-Caldas, junto a Vacaria, on Betula, 1944, C. Tavares 82 (LISU); Serra do Gerez-Vacaria,
no viveiro, 450 m, on Fagus, 1948, C. Tavares 2541 (LISU). Tras os Montes: EN of Villa Real, N
of road to Murça, S of Escarao, 700 m, on Castanea, 1995, P.v.d. Boom (hb. Boom 16977); Villa
Real, EN side of Village, Parque forestal, 700 m, on Q. rubra, 1995, P.v.d. Boom (hb. Boom
16939).—Spain: Andalucía: Huelva, Galaroza, Sierra de Navahermosa, La Suerte, 680 m, on
Q. suber, 1992, S. Fos (VAB-8445). Aragón: 14 km WNW of Bielsa, Ordesa, NW of Parador,
1350 m, on Betula, 1992, P.v.d. Boom (hb. Boom 13599). Castilla-León: León, Puerto del Pando,
1300 m, on Q. robur and Q. petraea, J. Etayo (hb. Etayo 5676). Salamanca, Montemayor, on
Castanea, 1981, B. Marcos (SALAF-1360); Salamanca, Casillas de Flores, on Q. pyrenaica, 1982,
B. Marcos (SALAF-1368); Salamanca, Monsagro, on Castanea, 1984, B. Marcos (SALAF-1404);
Salamanca, Herguijuela de la Sierra, on Fagus sylvatica, 1982, B. Marcos (SALAF-1381);
Salamanca, Las Batuecas, La Alberca, on Ailanthus altissima, 1989, B. Marcos (SALAF-1577);
Palencia, Espigëte, Cardaño de Abajo, 1400 m, on Q. pyrenaica, 1990, A. Terrón & A. Ubeda
(LEB-205). Catalunya: Vallès Oriental, Montseny, Turó de l’Areny, 900 m, on F. sylvatica, 1978,
A. Gómez-Bolea (BCC-3340); Font de Passaret, 1200 m, on F. sylvatica, 1979, A. Gómez-Bolea
(BCC-3338); Turó de I’Home, on F. sylvatica, 1978, A. Gómez-Bolea (BCC-3337). Vall d’Aran
von Kursiva, Viella, Bosc de Conangles, on cortex, 1984, A. Gómez-Bolea (BCC-3075, 3322);
Artiga de Lin, 1500 m, on F. sylvatica, 1979, A. Gómez-Bolea (BCC-3339); Montsianès non
Kursiva, Serra dels Ports, La Sènia, Barranc del Retaule, 1000–1100 m, on F. sylvatica, 1987,
M. Boqueras (BCC-5866); ibid, 1986, V. Atienza (VAB-945). Galicia: La Coruña, Grañas,
Mañón, on Q. robur, 1988, A. Garcia (SANT-6388); Rebolás, Ortigueria, on Q. robur, 1988,
A. Garcia (SANT-6385); Ancares, on Betula, 1992, G. Paz Bermúdez et al. (SANT-8707).
Navarra: Zugarramundi, 209 m, on Q. robur, 1988, J. Etayo (hb. Etayo 4423); Goizueta, 200 m,
Q. robur, 1986, J. Etayo (hb. Etayo 4125).
Buellia erubescens Arnold
Verhandl. zool.-bot. Gesellsch. 25: 493 (1875); type: Austria, Tirol, im Walde zwischen Passthurm
and Kirchamwald, c. 1070–1274 m, on Alnus incana, viii 1873, F. Arnold (M!—holotype).
New synonyms: Buellia zahlbruckneri J. Steiner, Annal. Naturhist. Hofmuseums Wien 23: 122
(1909); type: Asia Minor, (Sandschack) districtus Trapenzunti: ‘in catena montium ad septentrionem fluminis Charshut inter vicos Fol et Eseli prope alpem Kisyl Ali-Jaila, 1850 m, cortice
Fagi et Aceris’, 16 vii 1907, Handel-Mazzetti 1040 (W!—lectotype, selected here, WU!isolectotype).
Buellia jorgei Samp., Bolet. Soc. Broterian. 2(2): 177 (1923); type: Portugal, Minho, Ponte do
Lima: Sá (na veiga), ix 1922, G. Sampaio (LISU!—syntype).
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Iberian Buellia—Giralt et al.
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F. 3. Ascospore ontogeny. A, Buellia erubescens (M—holotype); B, B. hyperbolica (Fl—holotype).
Buellia-type ascospores showing tendencies to the Physconia-type (arrows). Scale =10 m.
(Fig. 3A)
Thallus whitish to whitish grey, thin, smooth to rimose, continuous, usually
delimited by a black prothallus. Apothecia (0·3–)0·8·1(–1·5) mm diam., sessile, scattered. Disc usually plane, more rarely subconvex to convex, often
pruinose, pruina when present usually K+red crystals (microscope slide).
Proper margin thick, entire, prominent, becoming thinner with age, usually
persistent. Excipulum very well developed, expanding to 80–125 m below,
K+yellow (microscope slide). Hymenium without oil droplets, (80–)100–
110 m high. Apical cells of the paraphyses slightly swollen, 2–3·5(–4) m
wide, brown pigmented. Ascospores (11–)13–16(–18)(6–)7–8·5(–9·5) m,
some when yoúng of the Physconia-type, with inner wall slightly thickened at
septum, soon becoming of the Buellia-type, with walls uniformly thin, apices
rounded, straight, roughly ornamented. Spermatia bacilliform, 6–7·50·8–
1 m.
Chemistry. Chemotype I: Thallus K+yellow-orange then red, PDyelloworange; thallus section and often also epihymenium crystals K+deep yellow
followed by the formation of red crystals (microscope slide). Atranorin
(major), norstictic acid (major), connorstictic acid (trace), hyposalazinic acid
(trace),zeorin and unknown secalonic acid derivative (minor), by TLC and
HPLC.
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Chemotype II: Thallus K+yellow and PD+yellowish. Atranorin, traces of
norstictic and connorstictic acids () and traces of secalonic acid derivative
() by TLC.
Hyposalazinic acid is recorded here for the first time from the genus Buellia.
This substance was also detected, by TLC and HPLC, in a chemotype of the
saxicolous B. sequax, which lacks norstictic acid. In this chemotype, hyposalazinic acid co-occurs with hypostictic acid. Hypostictic acid has been
recorded previously from ‘Buellia’ lindingeri (Kalb & Hafellner 1992). Inoue
(1993) mentions an unidentified minor constituent in B. subfrigida and B.
frigida. According to the chromatograms published by Inoue (1993: 21) this
unidentified chemical could well be hypostictic acid.
Taxonomic remarks. As a consequence of Schauer’s treatment (1965: 623),
the synonymy of B. erubescens Arnold and B. zahlbruckneri J. Steiner has
remained in doubt. Some authors consider that these taxa are conspecific
(Orange et al. 1992) while others remained unconvinced (Santesson 1993).
Buellia zahlbruckneri was described by Steiner (1909) and after the Latin
diagnosis he included the following information: ‘An Buchen- und Ahornrinde
bei der Kisil Ali-Jaila, 1850 m, Handel-Mazzetti nr 1040’. Thus, the specimens
so labelled should be considered the type specimens of B. zahlbruckneri. We
have checked two specimens labelled as such by Steiner (1909), one stored in
W and the other one in WU. The former (W) corresponds to the chemotype
of B. erubescens that contains norstictic acid (thallus K+red) and the second
(WU) to the chemotype that possesses only traces (detected by TLC) of this
substance (thallus K+yellow). Given their analogous morphology, we believe
that B. zahlbruckneri J. Steiner is a synonym of B. erubescens Arnold.
In the original description of B. zahlbruckneri, Steiner (1909) discussed the
differences between the new species and B. disciformis but he did not compare
it with B. erubescens. Several specimens of B. erubescens in WU were identified
by Steiner as B. zahlbruckneri J. Steiner var. erubescens Arnold. This shows that
Steiner knew of B. erubescens Arnold but that he considered it to be a variety
(characterized by the K+red thallus reaction) of his new taxon (see
Zahlbruckner 1931: 431). When Schauer (1965) observed that the type
material of B. zahlbruckneri (probably the specimen from W) gave a K+red
thallus reaction, he thought that this specimen was a false type and concluded
that there was not type material of B. zahlbruckneri (‘Das erwähnte Exemplar
nr. 1040 muß falsch bestimmt sein, so daß für B. zahlbruckneri J. Steiner,
kein Typus existiert’). Consequently, Schauer (1965) incorrectly neotypified
B. zahlbruckneri and, on the basis of the discussion of Malme (1927: 258), he
chose as neotype the specimen Arnold: Lich. Exs 1589 from M.
As indicated before (see taxonomic remarks under B. chloroleuca), the
specimens (invalid ‘isoneotypes’) of Arnold: Lich. Exs. 1589 investigated
(DUKE, FH, UPS) belong to B. chloroleuca. Thus, the invalid taxon B.
zahlbruckneri in the sense of Schauer (1965) is a ‘synonym’ of B. chloroleuca.
Discussion. In the study area, B. erubescens is a rather uniform species,
normally well characterized by the K+red thallus, the large and often pruinose
apothecia, which always possess a well developed K+yellow excipulum, the
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117
lack of hymenial oil droplets and the small Buellia-type ascospores. Although
several of the specimens investigated lacked the K+red thallus reaction
(norstictic acid), traces of this substance could nearly always be detected by
TLC. Besides the diagnostic characters cited above, a further feature that can
assist in identifying specimens of B. erubescens that lack the K+red thallus
reaction is the K+strong yellow reaction of the excipulum (microscope
slide). A similar reaction can be observed in the excipulum of B. chloroleuca,
B. hyperbolica and B. insignis. However, these species are well separated from
B. erubescens by their alternative chemistry, larger ascospores and significantly
different ecology (see comments under those taxa).
In the study area, B. erubescens could be confused with an undescribed
Buellia species, a species that also contains norstictic acid, lacks hymenial oil
droplets and has small ascospores. However, the new species is well separated
by smaller apothecia with a poorly developed excipulum, larger ascospores
that show clear Callispora-type tendencies and alternative ecology.
Buellia sanguinolenta, a further species that contains norstictic acid, is readily
distinguished from B. erubescens by the presence of numerous oil droplets in
the hymenium and much larger Callispora-type ascospores (see also comments
under B. sanguinolenta).
No significant differences have been observed between the type of B. jorgei,
a taxon originally described by Sampaio (1923) from Portugal, and the
specimens of B. erubescens examined (including the holotype). For this reason
we synonymyze B. jorgei with B. erubescens.
Ecology and distribution. In the Atlantic regions of the Iberian Peninsula,
B. erubescens seems to be a common species that grows from low to middle
altitudes, usually on the smooth bark of deciduous trees. From these areas it
extends to the Pyrenees but does not reach the Mediterranean mountains.
Buellia erubescens has been reported many times from the study area. Indeed,
most records of B. jorgei also refer to this species. The Mediterranean
records of B. erubescens refer to the undescribed Buellia species mentioned
above.
Exsiccata examined. Merrill, Lich. exs. 165 (as B. parasema, W).
Additional specimens examined. Portugal: Beira Litoral: 12 km S of Oliveira do Hospital, Ponte
das Tres Entradas, 350 m, on Alnus, 1993, P. v.d. Boom (hb. Boom 14288); Serra da Lousa,
Quinta de Alfocheira, na Mata, on Tilia, 1943, C. Tavares (LISU); Serra do Caramulo, Paredes
do Guardao, próximo da Ponte dos Amores, on Prunus, 1942, C. Tavares (LISU). Minho: Serra
do Gerez-Pedra Bela, 750 m, on Betula, 1947, C. Tavares (LISU-2013, 2040, UPS); Serra do
Gerez, próximo da Bouça da Mó, on llex, 1948, C. Tavares (LISU-2688). Tras os Montes: Villa
Real, EN side of Village, Parque Forestal, 700 m, on Quercus rubra, 1938, P. v.d. Boom (hb. Boom
16938).—Spain: Castilla-León: Salamanca, Las Batuecas, on Q. rotundifolia, 1981, B. Marcos
(SALAF-1356); on Alnus, 1983, B. Marcos (SALAF-1355). Galicia: La Coruña, Taboada,
Monfero, on Q. robur, 1988, A. Garcia (SANT-6387); Sarandons, Abegondo, on Q. robur, 1989,
A. Garcia (SANT-4824); Trazo, on Q. robur, 1989, A. Garcia (SANT-4823); Mairame, Cercede,
on Q. robur, 1989, A. García (SANT-4822); Fene, on Robinia pseudoacacia, 1987, A. García
(SANT-5966); Igrexafreda, S. Sadurniño, on Q. robur, 1988, A. Garcia (SANT-4823); Presa de
Vilagudún, Ordes, 1989, A. Garcia (SALAF-6308); Caaveiro, Pontedeume, on Q. robur, 1986,
J. Alvarez & M.E. López de Silanes (SANT-2423, 2421); ibid. on Castanea and Alnus, 1986,
J. Alvarez & M.E. López de Silanes (SANT-2422); Santiago de Compostela, on Q. robur, 1982
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(SANT-18); Lugo, Busir, Guitriz, on Q. robur, 1988, A. Garcia (SANT-4823); Carretera de
Pambre, on Betula celtiberica, J. Etayo (hb. Etayo-5637); Pontevedra, Capital, on Q. robur 1987,
A. Garcia (SANT-7037); Lourizán, on C. sativa, Q. robur, 1982 (SANT-19, 20). Navarra:
Refugio de Belagua, 900–1000 m, on Fagus, 1985, J. Etayo (hb. Etayo-3708); Guerendiain,
Viscarret, 500 m, 1986, J. Etayo (hb. Etayo); Alcoz, 560 m, on Q. robur, 1986, J. Etayo (hb.
Etayo); Lizardoya, 800–100 m, 1985, J. Etayo (hb. Etayo 4313); Bertiz, 200 m, on Fagus, 1985,
J. Etayo (hb. Etayo 4341); Irati, 800–100 m, on Fagus, J. Etayo (hb. Etayo 3202); Oroquieta,
580 m, on Fagus, 1986 J. Etayo (hb. Etayo).
Buellia hyperbolica Bagl
Nuov. Giorn. Botan. Ital. 3: 266 (1871); type: Italy, Toscana: ‘Sulle vecchie ceppaie di castagno
al Mt. Pisano’, Beccari (FI!—holotype).
(Fig. 3B)
Thallus of whitish, whitish grey to yellowish green granules, continuous or
not, usually covering a wide area. Apothecia lecideine, rapidly becoming
biatorine, 0·2–0·3(–0·4) mm diam., adnate, abundant and confluent. Disc
plane at first, soon hemispherical. Proper margin very thin, and even when
juvenile almost totally excluded. Excipulum poorly developed, up to 20 m
wide, K+yellowish (microscope slide). Hymenium without oil drops, 70–90
(–100) m high. Hypothecium interspersed with abundant oil droplets,
100–120 m deep, K+yellow (microscope slide). Apical cells of the paraphyses strongly swollen (3·5–)4–5 m wide, pigmented dark brown. Ascospores
(19–)21–24(–29)(7·5)8–9·5(–11) m, of the Physconia-type when young,
with inner wall thickened at septum, Buellia-type when mature, with walls
uniformly thin; apices pointed, often slightly curved, roughly ornamented.
Spermatia not seen.
Chemistry. Thallus K+yellowish brown, PD+orange-red. Atranorin (traces),
fumarprotocetraric acid (major), protocetraric acid (minor), confumarprotocetraric acid (minor) and convirensic acid (traces), by TLC and HPLC.
Hitherto, no chemical data were available for this species. Furthermore, this is
the first record of protocetraric, convirensic and confumarprotocetraric acids
from the genus Buellia.
Discussion. The chemistry readily distinguishes B. hyperbolica from the
other species treated in the present paper. Other diagnostic characters of this
species are the granulose thallus, the small, biatorine apothecia showing a
poorly developed excipulum, the lack of hymenial oil droplets and the large
ascospores.
According to the various revisions of this genus (e.g. Imshaug 1951; Sheard
1964; Harris 1988; Scheidegger 1993), B. dialyta (Nyl.) Tuck. and B.
rubifaciens R.C. Harris, are the only species of the genus reported to contain
fumarprotocetraric acid. Buellia rubifaciens, a corticolous species from Florida,
differs from B. hyperbolica in having smaller ascospores, 13–16·56·5–7·5 m
(Harris 1988). Buellia dialyta, a corticolous species originally described from
California, also reported from several localities in the northeastern United
2000
Iberian Buellia—Giralt et al.
119
States (Imshaug 1951), would appear to be closely related if not conspecific
with B. hyperbolica, as both taxa have the same diagnostic characters.
Nevertheless we prefer to maintain the name B. hyperbolica until the type
specimen of B. dialyta is available for study.
According to Marbach (1999), B. dialyta and B. rubifaciens belong to the
new genus Chrismofulvea Marbach, mainly characterized by the presence of
small apothecia, fumarprotocetraric acid and Physconia-type ascospores. He
also includes in Chrismofulvea two additional species, C. omalia Marbach and
C. pinastri (Erichsen) Marbach. The former species is separated from Buellia
hyperbolica by its sorediate thallus and the latter by its hymenium inspersed
with oil droplets.
Ecology and distribution. Buellia hyperbolica is a corticolous/lignicolous
species that grows mainly on mature trunks and wood of Castanea and
Quercus. It seems to be locally abundant in relatively undisturbed Castanea and
Quercus forests of the Mediterranean and southern Atlantic regions of the
Iberian peninsula. Hitherto the European distribution of this species was
poorly known as it has been reported only from the type locality in Italy, from
Portugal, (Boom et al. 1990), and from Spain (Boom & Gómez-Bolea 1991).
An additional record in Alvarez & Carballal (1992) from Galicia (Sierra de
Caurel, Lugo) has not been verified. The species has also been collected in
Windsor Great Park (London), on an ancient Quercus, and in Tuscany (Italy),
on Castanea (Coppins pers. com.).
Additional specimens examined. Portugal: Minho: 32·5 km of Braga, Parque Natural de Gerês,
N of Gerês, on wood of Quercus, P. v.d. Boom (hb. Boom 6334, 6339).—Spain: Asturias: 35 km
E of Cangas de Onis, 10 km of Panes, 0·7 km S of Trescares, on Castanea, 250 m, 1989, P. v.d.
Boom (hb. Boom 8947); 28 km E of Cangas de Onis, 2 km E of Arenas de Cabrales, S of road to
Panes, 200 m, on wood of Castanea, 1989, P. v.d. Boom (hb. Boom 8918, BCC-5116). Cantabria:
Santander, Picos de Europa, 4 km W of Potes, Argüebanes, 500 m, on wood of Castanea, 1987,
P. v.d. Boom (hb. Boom 6066). Navarra: Roncesvalles, 1000 m, on Castanea wood, 1986, J. Etayo
(hb. Etayo 3631).
Buellia leptoclinoides (Nyl.) J. Steiner
Verhandl. zool.-bot. Gesellsch. Wien 57: 357 (1907); type: France, Pyrénées-orientales,
Port-Vendres, 6 vii 1887, Nylander (H-Nyl 10541—lectotype, not seen).
(Fig. 2B)
Thallus whitish, pale yellowish grey to ochraceous, thick, rimose to crackedareolate, continuous, sometimes delimited by a blackish prothallus. Apothecia
(0·3–)0·5–0·8(–1) mm diam., scattered to confluent, adnate to sessile.
Disc plane to convex, epruinose. Proper margin initially moderately
thick, becoming thinner, finally excluded. Excipulum (30–)40–60 m wide,
K(microscope slide). Hymenium with numerous oil droplets, 80–120 m
high. Apical cells of the paraphyses slightly swollen, 2·5–3·5 m wide,
dark brown pigmented. Ascospores 15–20(–23)(6·5–)8–9(–9·5) m, of
typical Callispora-type, with inner wall clearly thickened subapically and at
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the septum, sometimes slightly curved and becoming triseptate, weakly
ornamented. Spermatia 4·5–61 m, bacilliform, straight.
Chemistry. Thallus K+yellow. Atranorin (minor) with chloratranorin
(traces) and placodiolic acid (major), by TLC and HPLC.
Discussion. Buellia leptoclinoides is characterized by the thick, pale yellowishgrey to ochraceous, cracked-areolate thallus, which contains atranorin and
placodiolic acid, the abundant and often crowded apothecia with convex discs,
the hymenium with numerous oil droplets, and the typical Callispora-type
ascospores. Among the species treated, B. leptoclinoides is unique in containing
placodiolic acid. In the study area only B. arnoldii and B. disciformis, which
also possess hymenial oil droplets and Callispora- or near Callispora-type
ascospores, could be mistaken for B. leptoclinoides. However, both taxa are well
separated by the significantly larger ascospores, the alternative chemistry and
ecology. A further distinguishing feature of B. leptoclinoides, which is absent in
B. arnoldii and B. disciformis, is the presence of large crystal complexes in the
thallus and outer part of the excipulum (placodiolic acid?). The size of the
spermatia also distinguishes B. disciformis from B. leptoclinoides.
This is the first time that B. leptoclinoides has been reported as growing on
bark. For the most part the different substratum separates these collections
from the more typical, saxicolous specimens of B. leptoclinoides. An additional
feature that seems to distinguish both ecotypes has been observed in the spore
ornamentation, which is smooth in the saxicolous specimens (Scheidegger
1993; Nordin 1998) but is weak in those specimens growing on bark.
However, in our opinion, this character is of little taxonomic significance to
separate a new taxon.
According to the literature (e.g. Kalb 1986; Scheidegger 1993; Sheard
1992; Pusswald et al. 1994), B. leptoclinoides is clearly distinguished from all
other corticolous and lignicolous Buellia species previously included in the
genus Hafellia (see Kalb & Elix 1998; Marbach 1999) by its smaller
ascospores. Only B. bahiana (=Hafellia bahiana), has ascospores of similar size
(cf. Sheard 1992). However, the presence of norstictic acid makes this North
American species clearly distinct from B. leptoclinoides.
Buellia leptoclinoides differs from all other Buellia species previously included
in the genus Hafellia, by the presence of placodiolic acid.
Ecology and distribution. In the study area the corticolous specimens of
B. leptoclinoides have been collected only in coastal localities of central and
southern Portugal. It seems to be confined to the coast, occurring in warm
places subject to humid coastal winds as well as in the salt-spray belt. It grows
mainly on the bark of maritime trees such as Ceratonia and Ficus.
Specimens examined. Portugal: Algarve: N of Albufeira, 1 km N of Alte, 400 m, on Ceratonia,
1993, P.v.d. Boom (hb. Boom 14925); W of Lagos, road Vale de Boi to Barao de San Miquel,
25 m, on Prunus dulcis, 1993, P. v.d. Boom (hb. Boom 14571); 14 km W of Lagos, along road to
Salema, 50 m, on Ceratonia, 1993, P.v.d. Boom (hb. Boom 14670); Alvor, zwischen Lagos und
Portimao, on Ficus, 1988, O. Breuss (hb. Breuss 5288). Estremadura: 25 km W of Setubal, Aldiea
do Meco, 40 m, on Ficus, 1987, P.v.d. Boom (hb. Boom 6600).
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Iberian Buellia—Giralt et al.
121
F. 4. Ascospore ontogeny. A, Buellia arnoldii (Italy, Südtitol, Arnold, PRM—syntype); B, B.
regularis (Kalb 23182—isotype); C, B. sanguinolenta (M—holotype). Typical Callispora-type
ascospores. Scale =10 m.
Other corticolous Buellia species examined that do not occur in the
study area
Buellia arnoldii Servít
Vest. Král. Spol. Nauk. Tr. 2: 39 (1931); types: Austria: Tirol, auf dem dickerer Äste von Taxus
im Walde östl. von Plansee, Arnold (M!, PRM!—syntypes).—Germany: Oberbayern, Wetterstein,
dürre Zirbenäste auf dem Schachen bei Partenkirchen, Arnold (M!, PRM!—syntypes).—Italy:
Südtirol, auf Rinde abgestorbener Fichtenzweige im Latemarwald-Karerpass, Arnold (M!,
PRM!—syntypes).
(Fig. 4A)
Thallus whitish to whitish grey, thin, smooth to rimose, continuous.
Apothecia 0·4–0·8 mm diam, scattered, sessile. Disc flat to subconvex. Proper
margin thin, persistent. Excipulum 60–90 m wide, K(microscope slide).
Hymenium 100–150 m high, inspersed with numerous oil droplets. Apical
cells of the paraphyses slightly swollen, (2·5–)3–4 m wide, brown pigmented.
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Ascospores (23–)26–29(–31)(10–)12–14(–16·5), Callispora-type, with inner
ascospore wall thickened subapically and at the septum, apices pointed, often
curved, smooth. Spermatia not seen.
Chemistry. Thallus K+yellow. Atranorin, by TLC, according to our own
analyses and Orange et al. (1992). Due to the scanty material of B. arnoldii, no
type specimen has been analysed by TLC and no lectotype selected.
Discussion. Buellia arnoldii is characterized by the interspersed hymenium,
the large Callispora-type ascospores and the presenće of atranorin. Among the
other species studied, B. arnoldii could be mistaken only for B. sanguinolenta
or B. regularis, both of which have inspersed hymenia and Callispora-type
ascospores. Buellia sanguinolenta is distinguished from B. arnoldii by its
alternative chemistry as it contains norstictic and traces of connorstictic acids
in addition to atranorin (). Our investigations have indicated that in the
genus Buellia, specimens that differ only by the presence/absence of norstictic
acid should not be considered different taxa, but only different chemotypes.
Scheidegger (1993) and Nordin (1996) came to a similar conclusion where,
for example, norstictic acid was found in only one of the two chemotypes of
B. sequax and of B. alboatra and B. venusta, respectively. In the present paper,
no taxonomic significance has been given to the presence or absence of
norstictic acid in B. erubescens. According to Schauer (1965) and Orange et al.
(1992), an additional distinguishing feature between these two taxa can be
found in the size of the apical cells of the paraphyses, being 4–4·5 m wide in
B. arnoldii and only 2–3 m wide in B. sanguinolenta. However, according to
our observations, these cells measure (2·5–)3–4 m in the types of B. arnoldii
and 2·5–3·5(–4) m in that of B. sanguinolenta.
The recently described B. regularis (Kalb & Hafellner 1992) is closely
related if not conspecific with B. arnoldii. It is said to be distinguished by
(significantly?) shorter and wider ascospores (25–2913–16·5 m), that
furthermore, are generally strongly curved [viz. (23–)26–29(–31)(10–)
12–14(–16·5) m, often curved ascospores, in B. arnoldii].
In our opinion, the characters used for the delimitation of B. sanguinolenta
and B. regularis fall within the variability of B. arnoldii. Furthermore, according
to the literature, the three taxa show very similar ecological preferences (see
Ecology and distribution). Nevertheless, until a thorough revision of the
B. arnoldii-group is carried out, we refrain from placing B. sanguinolenta and
B. regularis into synonymy with B. arnoldii.
Additional studies on the secondary lichen substances and the spermatia
would probably help in the delimitation of the species of B. arnoldii-group.
Unfortunately, spermatia were not found in any of the specimens examined
(types included). Further, TLC analyses could not be carried out on the type
specimens of B. arnoldii or B. regularis because of the scanty type material.
Literature reports on the spermatia and the chemistry of B. arnoldii-group are
inconsistent. According to Harris (1977) and Sheard (1992), the spermatia of
B. arnoldii are elongate-ellipsoid (6·5–7·51 m), whereas Orange et al.
(1992) reported them to be narrowly ellipsoid to oblong (3–3·51 m). Only
Orange et al. (1992) have described the spermatia of B. sanguinolenta, these
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123
being oblong-ellipsoid (3–41–1·5 m). In the original description of B.
regularis (Kalb & Hafellner 1992) there is no mention of its spermatia. As far
as the secondary lichen substances are concerned, B. arnoldii and B. regularis
have been reported to contain atranorin (Orange et al. 1992; Kalb & Hafellner
1993) and B. sanguinolenta to contain norstictic acid (Orange et al. 1992). The
latter result is not in complete agreement with our TLC analyses, since we
have shown that the holotype of B. sanguinolenta possesses traces of atranorin
and connorstictic acid in addition to norstictic acid.
Another species present in the study area that could be mistaken for B.
arnoldii is B. disciformis. However, it is well separated by the better developed
thallus, larger apothecia, smaller ascospores and significantly different
chemistry and ecology (see also the discussion under B. disciformis).
Ecology and distribution. Buellia arnoldii seems to be an oceanic species that
is widespread in western Europe. It has also been reported from a few localities
scattered throughout the Alps (Servı́t & Nádvornik 1931; Schauer 1965),
from Sweden (Fritz 1998) and from Macaronesia (Breuss 1988). The species
also occurs in western Scotland (Coppins pers. comm.). It usually grows on
conifers, in the montane belt. The single record of B. arnoldii from the Iberian
Peninsula in Paz Bermudez (1993) refers to B. disciformis.
Additional specimens examined. Austria: Tirol: Salzburg, Baumleichen des Hochmoorgebiets der
Gerlosplatte, an Fichte, 1640 m, 1963, Th. Schauer (M).—France: Mittelfranken, Dürre
Fichtenäste im Walde des Affentals bei Eichstätt, 1873, F. Arnold (M).
Buellia insignis (Nägeli ex Hepp.) Th. Fr.
Nova Acta Reg. Soc. Scient. Upsal. 3(3): 327 (1861); type: Hepp, Fl. Eur. 39a (G!—lectotype,
selected here; G!, BM!, PRM!—isolectotypes).
(Fig. 1B)
Thallus whitish, grey, cream to ochraceous, usually thick and warted, rarely
smooth, continuous and well developed, not delimited by a blackish prothallus. Apothecia (0·5–)0·7–1(–1·3) mm diam., abundant and confluent, subimmersed to adnate, rarely becoming sessile. Disc initially plane to convex,
epruinose. Proper margin initially thick, rapidly becoming thinner and finally
excluded. Excipulum well developed, 70–100 m wide, K+yellow (microscope
slide). Hymenium without oil droplets, 100–130 m high. Hypothecium with
oil droplets. Apical cells of the paraphyses strongly swollen, 4–6(–7) m wide,
brown pigmented. Ascospores (19–)21–25(–29)(8–)9–11(–12) m, resembling the Callispora-type when young, Buellia-type when mature, apices
pointed, often curved, roughly ornamented. Spermatia not seen.
Chemistry. Thallus UV+orange, K+yellowish, KC+orange; thallus section
C+orange. Atranorin (traces), 6-O-methylarthothelin (major),arthothelin
(traces) and gyrophoric acid (traces), by TLC and HPLC. The clarity of the
spot tests are dependent upon the concentration of 6-O-methylarthothelin
(similar variation was also observed with B. chloroleuca). This is the first report
of 6-O-methylarthothelin, arthothelin and gyrophoric acid in B. insignis.
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The chemistry of B. insignis is close to that of several species of Pertusaria,
subgenus Pionospora, for example, P. bryontha and P. dispar (Archer 1993),
which also contain a combination of a chlorinated xanthone and an orcinol
tridepside (6-O-methylarthothelin and gyrophoric acid, respectively) as major
compounds.
Discussion. Buellia insignis is well characterized by its chemistry, the lack of
hymenial oil droplets and the large Buellia-type ascospores, which show
Callispora-type tendencies when young.
The description given in Sheard (1964) and Orange et al. (1992) is, at least
in part, based on one specimen that, in our opinion, does not correspond to B.
insignis. This specimen (Vězda: Lich. sel. exs. 621, BM) is readily distinguished
from B. insignis by possessing a thallus composed of subhemisphaerical
areolae, smaller apothecia with pruinose discs and hymenial oil droplets.
Imshaug (1951) considered that B. papillata (Sommerf.) Tuck. was
the correct name for B. insignis but according to Sheard (1964), Schauer
(1965), Wirth (1988) and Galloway et. al. (1998), these taxa are not
conspecific. This problem can be solved only by studying the type specimen of
B. papillata.
Ecology and distribution. Buellia insignis is an arctic-alpine species that occurs
on mosses, plant debris and more rarely on coniferous bark. It has not been
reported from the Iberian Peninsula. For its European distribution see Orange
et al. (1992) and Nimis (1993).
Exsiccatae examined. Fellman, Lichenes arctici colleti aestate 1863 201 (BM).
Selected specimens examined. Austria: ‘Ad terram et muscos in montibus altioribus et alpibus
prov. Sondriensis’ (BM).—Norway: Berleraag, 1664, Th. Fries (BM).—Sweden: Lule Lappmark, Kvikkjokk par., 3 km SSW of Kvikkjokk, SE slope of Nammatj, 500–525 m, Picea forest,
1977, B. J. Coppins & L. Tibell (BM).—Switzerland: Berner Oberland, Iffigenfall mi Simmebtal
bei Lenk, 1450 m. on Picea, 1964 Frey & Schauer (M).—‘In terra muscosa alpis Sujarrak
Lapponiae Lulensis’, 1871, J. Hellbom (BM).
Buellia regularis Kalb
Herzogia 9: 56 (1992); type: Portugal (Macaronesia), Madeira, an der Levada da Serra do Faial
entlang, etwas SW von Santo da Serra in Richtung Camacha. In letzten Resten eines alten, jedoch
stark gestörten, NW-exponierten Eucalyptus-Waldes. An Erica arborea, 750 m, 13 viii 1990, K. &
A. Kalb (hb. Kalb 23183—holotype, 23182!—isotype).
(Fig. 4B)
Thallus whitish grey, thin, smooth, continuous, often delimited by a black
prothallus. Apothecia 0·3–0·7 mm diam. Disc flat to subconvex. Proper
margin thin, persistent. Excipulum 60–80 m wide, K(microscope slide).
Hymenium 100–150 m high, interspersed with numerous oil droplets. Apical
cells of the paraphyses 2·5–3·5 m wide, brown pigmented. Ascospores 25–
2913–16·5 m, Callispora-type, with inner ascospore wall thickened subapically and at the septum; apices pointed, usually strongly curved, smooth.
Spermatia not seen.
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125
Chemistry. Thallus K+yellow. Atranorin by TLC (Kalb & Hafellner 1992).
Discussion. In our opinion, B. regularis is simply a form of B. arnoldii with
shorter, thicker and more strongly curved ascospores and which grows on
Erica arborea bark instead of on coniferous bark (including Juniperus). See also
comments under B. arnoldii.
Ecology and distribution. Buellia regularis is a submontane species originally
reported from Madeira where it grows on Erica arborea (Kalb & Hafellner
1992). It is also widespread in the Canary Islands, where it occurs mostly on
conifers, Erica and the twigs of Laurus (Hafellner 1995). It has not been
reported from the Iberian Peninsula. See also comments on the ecology and
distribution of B. arnoldii.
Buellia sanguinolenta T. Schauer
Mitt. Bot. München 5: 620 (1965); type: Austria, Niederösterreich, Gr. Urwald bei Lunz, in cortice
Abietis albae, 1100 m, 221962, Th. Schauer (M!—holotype).
(Fig. 4C)
Thallus thin, whitish, smooth, continuous or discontinuous and then
composed of granules. Apothecia 0·3–0·6 mm diam., scattered, sessile. Disc
plane to convex epruinose. Proper margin thin, becoming excluded. Excipulum
60–70 m wide, K(microscope slide). Hymenium up to 140 m high,
interspersed with numerous oil droplets. Apical cells of the paraphyses slightly
swollen, 2·5–3·5(–4) m wide, brownish pigmented. Ascospores (21–)26–32
(–35)(11–)12–13(–16) m, of the Callispora-type, with inner ascospore wall
thickened subapically and at the septum at all ontogenetic stages; apices
pointed, often curved, some spores becoming triseptate, smooth. Spermatia
not seen.
Chemistry. Thallus K+yellow-orange, at first, to finally red, PDyelloworange; thallus section K+deep yellow followed by the formation of red
crystals (microscope slide). Atranorin (traces), norstictic and connorstictic
acids by TLC.
Discussion. Buellia sanguinolenta is well characterized by the presence of
atranorin () and norstictic (with connorstictic) acid in the thallus, the
hymenium inspersed with numerous oil droplets and the large Callispora-type
ascospores. The presence of norstictic acid separates B. sanguinolenta from
B. arnoldii. In our opinion, B. sanguinolenta may be just one of the two
chemotypes of B. arnoldii, characterized by the occurrence of this lichen
substance (see also comments under B. arnoldii and B. regularis). Buellia
erubescens shows analogous chemical races.
Specimens of B. erubescens that contain norstictic acid are mainly distinguished from B. sanguinolenta by the lack of oil droplets in the hymenium
and smaller ascospores (see also comments under B. sanguinolenta).
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Ecology and distribution. According to Schauer (1965) and Orange et al.
(1992), B. sanguinolenta is an oceanic species, known from Scotland and from
three localities in the western Alps situated between 1100 and 1350 m
altitude. In Scotland it grows on smooth bark of Sorbus and in Austria on
Abies. The record of B. sanguinolenta from the Iberian Peninsula in Atienza
(1991) corresponds to the same undescribed Buellia species mentioned in the
discussion of B. erubescens (see above).
Additional specimen examined: Germany: Oberbayern, Garmisch, Lahnenwies-Graven,
1350 m, an Tanne, 1963, Schauer (GZU).
We are indebted to the directors and keepers of the following herbaria: BCC, BM, DUKE, FH,
FI, G, GZU, H, KL, L, LEB, LISU, M, MUB, PRM, SALAF, SANT, VAB, UPS, W, WU and
to P.v.d. Boom (Son), O. Breuss (Vienna), J. Etayo (Navarra), J. Hafellner (Graz), K. Kalb
(Neumarkt/OPf.), who kindly lent us their private collections. We wish to thank X. Llimona
(Barcelona) for critical revision of the manuscript. The projects PB 96–1115–C04–02
and PB97–0957 (Spanish Government) and the ‘Comissionat per a la Recerca’ (Catalan
Government), are acknowledged by the two first authors for financial support.
R
Alvarez, J. & Carballal, R. (1992) Líquenes y hongos liquenícolas interesantes de la Sierra de
Caurel (Lugo, Noroeste de España). Cryptogamie. Bryologie et Lichénologie 13: 359–369.
Archer, A. W. (1993) A chemical and morphological arrangement of the lichen genus. Pertusaria.
Bibliotheca Lichenologica 53: 1–17.
Atienza, V. (1991) Flora y Vegetación Liquénica Epifitica de las Comarcas de Els Ports y Baix Maestrat
(Castellón) y Territorios Próximos. Doctoral thesis. University of Valencia.
Boom, v. d. P. & Gómez-Bolea, A. (1991) Contribution to the lichen flora of Spain. Nova
Hedwigia 53: 497–505.
Boom, v. d. P., Aptroot, A. & Knaap, W. O. v. d. (1990) New and interesting lichen records from
Portugal. Nova Hedwigia 50: 463–472.
Boqueras, M. (1993) Flora i Vegetació dels Liquens Epifı́tics de les Terres Meridionals de Catalunya.
Doctoral thesis. University of Barcelona.
Breuss, O. (1988) Neue und bemerkenswerte Flechtenfunde aus Tenerife (Kanarische Inseln).
Linzer biologische Beiträge 20: 829–845.
Culberson, C. F. & Ammann, K. (1979) Standardmethode zur Dünnschichtchromatographie
von Flechtensubstanzen. Herzogia 5: 1–24.
Culberson, C. F. & Johnson, A. (1982) Substitution of methyl tert.-butyl ether for diethyl ether
in the standardized thin-layer chromatographic method for lichen products. Journal of
Chromatography 238: 483–487.
Culberson, C. F., Culberson, W. L. & Johnson, A. (1981) A standardized TLC analysis of
-orcinol depsidones. Bryologist 84: 16–29.
Etayo, J. (1989a) Liquenes Epifitos del Norte de Navarra. Doctoral thesis. University of Navarra.
Etayo, J. (1989b) Les lichens épiphytes du pin noir a Larra (Navarre, Espagne). Cryptogamie,
Bryologie et Lichénologie 10: 309–312.
Feige, G. B., Lumbsch, H. T., Huneck, S. & Elix J. A. (1993) The identification of lichen
substances by a standardized high-performance liquid chromatographic method. Journal of
Chromatography 646: 417–427.
Fritz, Ö. (1998) Buellia arnoldii påträffad in Halland. Graphis Scripta 9: 43–44.
Galloway, D. J., Sheard, J. W. & Elix, J. A. (1998) Additional lichen records from New Zealand
28. Buellia papillata (Sommerf.) Tuck. Australasian Lichenology 43: 6–8.
Giralt, M. (1994) Key to the corticolous and lignicolous species of the genus Rinodina in the
Iberian Peninsula and Balearic Islands. Bulletin de la Société Linnéenne de Provence 45:
317–326.
Giralt, M. & Mayrhofer, H. (1994) Four corticolous species of the genus Rinodina (lichenized
Ascomycetes, Physciaceae) containing atranorin in southern Europe and adjacent regions.
Nova Hedwigia 59: 129–142.
2000
Iberian Buellia—Giralt et al.
127
Giralt, M. & Mayrhofer, H. (1995) Some corticolous and lignicolous species of the genus
Rinodina (lichenized Ascomycetes, Physciaceae) lacking secondary lichen compounds and
vegetative propagules in Southern Europe and adjacent regions. Bibliotheca Lichenologica 57:
127–160.
Hafellner, J. (1995) Bemerkenswerte Funde von Flechten und lichenicole Pilzen auf makaronesischen Inseln III. Einige bisher auf den Kanarischen Inseln übersehene lecanorale Arten.
Linzer biologische Beiträge. 27: 489–505.
Harris, R. C. (1977) Lichens of the Straits Counties. Michigan: Ann Arbor.
Harris, R. C. (1988) Buellia in North America and central Florida or the virtues and rewards of
collections. Evansia 5: 37–45.
Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler, D. N. (1995) Ainsworth & Bisby’s
Dictionary of the Fungi. 8th. Edn. Wallingford, Surrey: C.A.B. International.
Huneck, S., John, V. & Elix, J. A. (1992) Zur Chemie einiger Strauch- und Krustenflechten aus
der Türkei. Herzogia 9: 173–179.
Imshaug, H. A. (1951) The Lichen-forming Species of the Genus Buellia in the United States and
Canada. Doctoral thesis. University of Michigan.
Inoue, M. (1993) Buellia subfrigida sp. nov. (Lichenes, Buelliaceae) from Lützow-Holm Bay Area
and Prince Olav Coast, East Antarctica. The asexual sorediate species forming a species pair
with B. frigida Darb. Nankyoku Shiryó (Antarctic Record) 37: 19–23.
Kalb, K. (1986) Lichenes Neotropici. Fasc. IX, Nr. 351–400: 1–16. Neumarkt.
Kalb, K. & Elix, J. A. (1998) The chemistry of some species of Buellia sensu lato (Lecanorales,
Lichenized Ascomycotina). Mycotaxon 68: 465–482.
Kalb, K. & Hafellner, J. (1992) Bemerkenswerte Flechten und lichenicole Pilze von der Insel
Madeira. Herzogia 9: 45–102.
Leuckert, C. & Mathey, A. (1975) Beiträge zur Chemotaxonomie einiger Xanthon-Haltiger Arten
der Flechtengattung Buellia. Herzogia 3: 461–488.
Malme, O. (1927) Lichenologiska notiser. Svenska botanisk tidskrift 21, H. 2: 251–259.
Marbach, B. (1999) Corticole und lignicole Arten der Flechtengattung Buellia sensu lato in den
Subtropen und Tropen. Bibliotheca Lichenologica (in press).
Marcos, B. (1985) Flora y Vegetación Liquénica Epifı́tica de las Sierras Meridionales Salmantinas.
Doctoral thesis. University of Salamanca.
Mayrhofer, H. (1982) Ascosporen und Evolution dér Flechtenfamilie Physciaceae. Journal of the
Hattori Botanical Laboratory 52: 313–321.
Mayrhofer, H. (1984) The saxicolous species of the genus Dimelaena, Rinodina and Rinodinella in
Australia. Nova Hedwigia 79: 511–536.
Moberg, R., Nordin, A. & Scheidegger, C. (1998) Proposal to change the listed type of the name
Buellia, nom. cons. (Physciaceae, lichenised Ascomycota). Taxon 48: 143.
Nimis, P. L. (1993) The Lichens of Italy. An Annotated Catalogue. Torino: Museo Regionale di
Scienze Naturali.
Nordin, A. (1996) Buellia species (Physciaceae) with pluriseptate spores in Nordern. Symbolae
Botanicae Upsalienses 31(3): 327–354.
Nordin, A. (1998) Ascospore characters in Physciaceae: an ultrastructural study. Symbolae
Botanicae Upsalienses 32(1): 195–208.
Orange, A., Coppins, B. J. & Scheidegger, C. (1992) Buellia de Not (1846). In The Lichen Flora
of Great Britain and Ireland (O. W. Purvis, B. J. Coppins, D. L. Hawksworth, P. W. James &
D. M. Moore, eds): 129–137. London: Natural History Museum Publications.
Paz Bermúdez, G. (1993) Líquenes Epífitos Sobre Abedul en Galicia. Doctoral thesis. University of
Santiago de Compostela.
Pusswald, W., Kantvilas, G. & Mayrhofer, H. (1994) Hafellia dissa and H. levieri (lichenised
Ascomycetes, Physciaceae), two corticolous and lignicolous species in Tasmania. Muelleria 8:
133–140.
Rambold, G., Mayrhofer, H. & Matzer, M. (1994) On the ascus types in the Physciaceae
(Lecanorales). Plant Systematics and Evolution 192: 31–40.
Sampaio, G. (1923) Novos materiais para a liquenologia portuguesa. Boletim da Sociedade
Broteriana 2: 161–179.
Santesson, R. (1993) The Lichens and Lichenicolous Fungi of Sweden and Norway. Lund:
SBT-förlaget.
128
THE LICHENOLOGIST
Vol. 32
Servít, M. & Nádvornik, J. (1931) Flechten aus der Cechoslovakei II. Karpatorussland und
Südostslovakei. Vestniku Královské Ceské Spolecnosti Nauk 2: 36–42.
Schauer, T. (1965) Die holz- und rindenbewohnenden Arten der Flechtengattung Buellia s.
str. im Nordalpenraum. Mitteilungen aus der Botanischen Staatssammlung München 5:
609–626.
Scheidegger, C. (1993) A revision of European saxicolous species of the genus Buellia de Not. and
formerly included genera. Lichenologist 25: 315–364.
Sheard, J. W. (1964) The genus Buellia de Notaris in the British Isles (excluding section Diploicia
(Massal.) Stiz.). Lichenologist 2: 225–262.
Sheard, J. W. (1992) The lichenized Ascomycete genus Hafellia in North America. Bryologist 95:
79–87.
Steiner, J. (1909) Lichenes. Annalen des naturhistorischen Hofmuseums, Wien 23: 107–123.
Vobis, G. & Hawksworth, D. L. (1981) Conidial lichen-forming fungi. In The Biology of Conidial
Fungi (G. T. Cole & B. Kendrick, eds): 245–273. New York: Academic Press.
Wirth, V. (1988) Flechtenflora. Bestimmung und Ökologische Kennzeichnung der Flechten Südwestdeutschlands und Angrenzender Gebiete. Stuttgart: Ulmer.
Zahlbruckner, A. (1931) Catalogus Lichenum Universalis. Bd. 7. Leipzig: Gebrüder Bornaträger.
Accepted for publication 20 November 1999