Phacothecium resurrected and the new genus Phacographa (Arthoniales) proposed

Abstract

Phacothecium Trevis. is resurrected from the synonymy of Arthonia resp. Opegrapha. Its holotype species is Phacothecium varium (Tul.) Trevis., a name replacing the commonly used heterotypic synonym Opegrapha physciaria (Nyl.) D.Hawksw. The new genus Phacographa Hafellner is described to accommodate Lecidea glaucomaria Nyl. (holotype) and some related species, including Leciographa zwackhii A.Massal. ex Zwackh. Furthermore, Phacographa protoparmeliae Hafellner is described as new to science, and the new combinations Phacographa glaucomaria (Nyl.) Hafellner and P. zwackhii (A.Massal. ex Zwackh) Hafellner are proposed. A lectotype and epitype is designated for Phacopsis varia Tul., a lectotype is chosen for Lecidea glaucomaria Nyl., and lectotype species are designated for the genera Celidiopsis A.Massal. (C. gyrolophii A.Massal.) and Spilodium A.Massal. (S. fuscopurpureum (Tul.) A.Massal.). The positions of Phacothecium and Phacographa among other opegraphoid genera, with special emphasis on the delimitation against other similar arthonialean genera with lichenicolous species, i.e. Opegrapha, Lecanographa and Plectocarpon, are discussed. The known species of opegraphoid lichenicolous fungi with widely exposed hymenia are keyed. Opegrapha anomea Nyl. and O. rotunda Hafellner could not be assigned to one of the two treated genera, nor O. phaeophysciae and O. sphaerophoricola.

Full text

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Diversity of Lichenology – Jubilee Volume.
A. Thell, M. R. D. Seaward & T. Feuerer (eds). Bibliotheca Lichenologica 100: 85-121.
J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, Berlin · Stuttgart, 2009.
Phacothecium resurrected and the new genus
Phacographa (Arthoniales) proposed
Josef HAFELLNER
Institut für Pflanzenwissenschaften, Karl-Franzens-Universität Graz, Holteigasse 6, A-8010
Graz, Austria; e-mail: josef.hafellner@uni-graz.at
Abstract: Phacothecium Trevis. is resurrected from the synonymy of Arthonia resp.
Opegrapha. Its holotype species is Phacothecium varium (Tul.) Trevis., a name replacing
the commonly used heterotypic synonym Opegrapha physciaria (Nyl.) D.Hawksw. The
new genus Phacographa Hafellner is described to accommodate Lecidea glaucomaria
Nyl. (holotype) and some related species, including Leciographa zwackhii A.Massal. ex
Zwackh. Furthermore, Phacographa protoparmeliae Hafellner is described as new to
science, and the new combinations Phacographa glaucomaria (Nyl.) Hafellner and P.
zwackhii (A.Massal. ex Zwackh) Hafellner are proposed. A lectotype and epitype is
designated for Phacopsis varia Tul., a lectotype is chosen for Lecidea glaucomaria Nyl.,
and lectotype species are designated for the genera Celidiopsis A.Massal. (C. gyrolophii
A.Massal.) and Spilodium A.Massal. (S. fuscopurpureum (Tul.) A.Massal.). The positions
of Phacothecium and Phacographa among other opegraphoid genera, with special
emphasis on the delimitation against other similar arthonialean genera with lichenicolous
species, i.e. Opegrapha, Lecanographa and Plectocarpon, are discussed. The known
species of opegraphoid lichenicolous fungi with widely exposed hymenia are keyed.
Opegrapha anomea Nyl. and O. rotunda Hafellner could not be assigned to one of the
two treated genera, nor O. phaeophysciae and O. sphaerophoricola.
Zusammenfassung: Phacothecium Trevis., lange Zeit in der Synonymie von Arthonia
resp. Opegrapha, wird wieder als Gattung anerkannt. Die Typusart ist Phacothecium
varium (Tul.) Trevis., ein Name, der das häufig benutzte heterotypische Synonym
Opegrapha physciaria (Nyl.) D.Hawksw. ersetzt. Die neue Gattung Phacographa
Hafellner wird beschrieben, um in ihr Lecidea glaucomaria Nyl. (holotype) und
Leciographa zwackhii A.Massal. ex Zwackh unterzubringen. Weiters wird Phacographa
protoparmeliae Hafellner neu beschrieben. Die neuen Kombinationen Phacographa
glaucomaria (Nyl.) Hafellner und Phacographa zwackhii (A.Massal. ex Zwackh)
Hafellner werden vorgeschlagen. Für Phacopsis varia Tul. wird ein Lectotypus und ein
Epitypus bestimmt. Für Lecidea glaucomaria Nyl wird ein Lectotypus ausgewählt.
Lectotypus-Arten werden bestimmt für die Gattungen Celidiopsis A.Massal. (C.
gyrolophii A.Massal.) und Spilodium A.Massal. (S. fuscopurpureum (Tul.) A.Massal.).
Die systematische Stellung von Phacothecium und Phacographa neben anderen
opegraphoiden Gattungen wird diskutiert. Spezielles Augenmerk wird auf die
Abgrenzung gegen ähnliche arthoniale Gattungen mit lichenicolen Arten gelegt, im
besonderen Opegrapha, Lecanographa, und Plectocarpon. Für die bisher bekannten
opegraphoiden lichenicolen Pilze mit weit entblößten Hymenien wird ein Schlüssel
vorgelegt. Opegrapha anomea Nyl. und O. rotunda Hafellner konnten wegen
Abweichungen in den Ausprägungen als wesentlich erachteter phänotypischer Merkmale

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keiner der beiden behandelten Gattungen zugeordnet werden, und auch nicht O.
phaeophysciae und O. sphaerophoricola.
Key words: Ascomycotina, Opegrapha, lichenicolous fungi, taxonomy, key
Introduction
In the past, separate genera have often been described for lichenicolous taxa
resulting in parallel sets of genera for lichens and lichenicolous fungi (e.g. Buellia
– Epilichen, Catillaria – Scutula) and on the other hand for fungi on higher plants
and lichenicolous taxa (e.g. Rosellinia – Roselliniella, Mycosphaerella
(Sphaerella) – Stigmidium (Pharcidia), Didymella – Didymellopsis). Although the
introduction of these genera was often based on formalistic arguments, later it
often turned out that the “parallel” taxa are justified, but for reasons (including
anatomical features, secondary chemistry, mitosporic states, sequence data) other
than originally stated.
A considerable number of lichenicolous fungi belong to the Arthoniales. As
arthonialean fungi are generally regarded as primarily lichenized, lichenicolous
members of the order could be called “secondarily delichenized fungi”. A number
of genera in the Arthoniales have also been proposed mainly, or only, because the
species live on lichens (e.g. Opegrapha – Leciographa, Arthonia – Conida), or
genera have been described at the same time not recognizing the true relationship
(e.g. Plectocarpon). In the second half of the 20th century, most of these generic
names were reduced to the synonymy of lichenized fungal genera, which often
have priority over their non-lichenized counterparts, but have to be reconsidered
in case they are suitable for better circumscribed off-splits. So, e.g. under
Arthonia, 6 (7) generic synonyms are listed, the type species of which is a
lichenicolous fungus, and under Opegrapha there are 4 (ERIKSSON &
HAWKSWORTH 1998, ROBERT et al. 2005) (Tables 1-2).
There is no doubt that the lichenicolous species of the larger arthonialean
genera Arthonia and Opegrapha, taken as a group would turn out to be
polyphyletic, as the lichenicolous species evolved from different species groups of
their lichenized core. This was for instance demonstrated for lichenicolous
Arthonia species (GRUBE & MATZER 1997).
Recently, several opegraphoid lichenicolous fungi have been called the
Opegrapha anomea-group (ERTZ et al. 2004, 2005). As one characteristic feature
for this species group, a breakdown of the stromatic layer obtecting the
hymenium, starting with the formation of deep clefts, was stressed. Besides O.
anomea Nyl. and O. physciaria (treated here under the heterotypic synonym
Phacothecium varium, see below), the previously described Opegrapha
plectocarpoidea Diederich, O. pulvinata Rehm ex Lojka, O. sphaerophoricola
Isbrand & Alstrup, O. zwackii A.Massal. ex Zwackh (to Phacographa, see below)
and O. staurothelicola Fink in J. Hedrick were listed as possibly belonging to this
group of species, whereas O. blakii Ertz & Diederich and later O. phaeophysciae
R.Sant., Diederich, Ertz & Christnach were added as new taxa. ERTZ et al. (2005)
discuss also which generic names would be available and mention Mycopegrapha
Vain. and Opegraphoidea Fink, both commonly listed among the synonyms of

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Opegrapha Ach. (Table 2). The oldest available name, however, which is based
on one of the species in discussion, Phacothecium Trevis., was not mentioned.
Table 1. Generic and subgeneric synonyms of Arthonia with lichenicolous type species
as listed by ERIKSSON & HAWKSWORTH (1998) and Mycobank database (ROBERT et al.
2005).
genus type species type designation
Celidiopsis A.Massal. (1856) C. gyrolophii A.Massal.1 lectotype, designated here!
Celidium Tul. (1852) C. fuscopurpureum Tul. lectotype, designated by
SANTESSON (1960)
Charcotia Hue (1915) C. rufidula Hue holotype
Conida A.Massal. (1856) C. clemens (Tul.) A.Massal. lectotype, designated by
CLEMENTS & SHEAR (1931)
Conida subgen. Conidella
Elenkin (1901)2
C. urceolata Elenkin holotype
Phacothecium Trevis.3 (1857) P. varium (Tul.) Trevis. holotype
Spilodium A.Massal. (1856) S. fuscopurpureum (Tul.)
A.Massal.4
lectotype, designated here!
1 C. insitiva (Körb.) A.Massal. is a second species listed with the protologue of the
genus.
2 Listed as genus by ERIKSSON & HAWKSWORTH (1998) and in Mycobank.
3 Listed by ERIKSSON & HAWKSWORTH (1998), in Mycobank recognized as
independent genus.
4 S. affine A.Massal. is a second species listed with the protologue of the genus.
Table 2. Generic synonyms of Opegrapha with lichenicolous type species as listed by
Eriksson & Hawksworth (1998) and Mycobank database (ROBERT et al. 2005).
genus type species type of type
Leciographa A.Massal. (1854) L. parasitica A.Massal. holotype
Lecoglyphis Clem. (1909) L. centrifuga (A.Massal.) Clem. holotype
Mycopegrapha Vain. (1921) M. hymeniicola Vain. holotype
Opegraphoidea Fink1 [in J.Hedrick]
(1933)
O. staurothelicola Fink1 [in J.Hedrick] holotype
1 Both the generic and species name are ascribed to Fink, and in the introductory
remarks HEDRICK (1933: 303) states that “...the descriptions were prepared by Dr.
Fink.” According to Article 46.4. of ICBN the author of the taxa is therefore Fink.
In the opinion of this author, the Opegrapha anomea-group in the sense of
ERTZ et al. (2005) is not monophyletic and therefore it is worth publishing a
grouping concept for some opegraphoid lichenicolous fungi developed in recent
years and which more suitably reflects the relationship of the species in
discussion. In the concept of this author, the Opegrapha anomea-group sensu
ERTZ et al. (2005) consists of at least three clearly distinct species groups, of
which two are treated as genera here.
Unfortunately both type species of the genera considered by ERTZ et al. (2005)
as possible generic names were not available for study. However, taking into
consideration the characters of the type species as published in the protologue,
Mycopegrapha hymeniicola Vain. can easily be incorporated in Opegrapha,

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whereas Opegraphoidea staurothelicola Fink in J.Hedrick needs to be carefully
studied again. This reinvestigation raises three possible results: 1. Opegraphoidea
is a synonym of Opegrapha, 2. Opegraphoidea is a synonym of Phacothecium,
and 3. Opegraphoidea is a generic name available for one of the other
opegraphoid species groups, but none of them will have a direct influence on the
results presented here.
Material and methods
Methods
Dried herbarium specimens have been examined. External morphology was
studied with a dissecting microscope (WILD M3, 6.4-40x). Anatomical studies of
the ascomata were carried out under the light microscope (LEICA DMRE, 100-
1000x). Sectioning was performed with a freezing microtome (LEITZ, sections of
12-15 μm) but squash preparations were also used, especially for ascus analysis.
Preparations were mounted in water. When necessary, contrasting was performed
by a pretreatment with lactic acid-cotton blue (MERCK 13741). Amyloid
reactions in hymenia were observed under various concentrations of Lugol's
reagent (MERCK 9261) with and without pretreatment with diluted KOH.
Conidiogenesis of pycnoconidia was studied in erythrosin B (ALDRICH 19,826-
9) in 10% ammonia. Sections and squash preparations were not pretreated with
KOH unless otherwise stated. Measurements refer to dimensions in tap water.
Secondary chemistry of non-crystallized apothecial pigments was tested according
to MEYER & PRINTZEN (2000) and the nomenclature there proposed is applied.
Abbreviations for institutional herbaria follow HOLMGREN et al. (1990) and
for author names those proposed by BRUMMITT & POWELL (1992).
Material used for comparative studies
Lecanographa grumulosa (Dufour) Egea & Torrente (all on thalli of Roccella
spec. unless otherwise stated)
Europe: Italy: Sicilia, Isole Pelagie, Linosa, Costa da Faraglione a Calla Mannarazzo, 2-
10 m, on Roccella phycopsis, 16.4.1992, J. Poelt (GZU). – Sicilia, Isole Pelagie,
Lampedusa, Presso Capo Grecale, c. 80 m, rocce calcaree verticali esposte a nord, auf
Roccella phycopsis, 11.4.1992, J. Poelt (GZU). – Sicilia, Isole Pelagie, Lampedusa, punta
occidentale dell’isola, tra Punta Parise, Capo Ponente e Capo Teresa, 80-100 m, auf
Roccella phycopsis, 13.4.1992, J. Poelt (GZU). – France: Korsika, Dept. Corse du Sud:
Punta de la Parata W von Ajaccio, c. 50 m; NE-exponierte Felsklippen an der Küste mit
niederem Buschwerk, auf schattigen Abbrüchen von Vulkanitschrofen, auf Roccella
phycopsis, 5.11.1993, H. Kammerer, H. Mayrhofer & E. Unger (GZU). – [Spain],
Balearen, Mallorca, Ca’n Picafort, Küstenbereich am östlichen Ortsrand, c. 5 m,
anstehende Kalk- und Konglomeratfelsen, auf Roccella phycopsis, 19.5.1985, J. Poelt et
al. (GZU). – Africa: Canary Islands: Lanzarote, sanft geneigte Hänge NE von Arrieta,
bei der Abzweigung nach Cueva de los Verdes, 29°09'30"N / 13°26'10"W, c. 60 m;
Malpais mit Sukkulentenbusch, auf niederen, porösen Lavablöcken, 8.4.1999, A.
Hafellner & J. Hafellner 47621 (herb. Hafellner). – Fuerteventura, E-seitiger Rücken des
Vulkans Arena, kurz N des Dorfes La Oliva, 28°37'50"N / 13°55'30"W, c. 280 m; altes,
durch Lesesteinmauern parzelliertes Malpais und dazwischen größere Schrofen, NW-
seitig an Steilflächen, 27.4.2000, A. Hafellner & J. Hafellner 53415 (herb. Hafellner). –

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La Gomera, S ober der Straße von Vallehermoso nach Hermigua, auf der Nordseite des
Roque Cano, 28°10'40"N/17°14'40"W; Vulkanit, auf geneigten Erdflächen in lückiger
Garrigue, c. 380 m, 11.2.1991, J. Hafellner 34401 & A. Hafellner (herb. Hafellner). - El
Hierro, E-exponierte Hänge SE unterhalb von Valverde, c. 400 m, 27°48'10"N/17°54'W;
Kleinio-Euphorbietum mit Vulkanit-blockwerk, in einem Überhang eines kleinen
Basaltschrofens, 10.2.1995, J. Hafellner 48299 (herb. Hafellner). – Madeira: Ponta de
S.Lourenco, 5 km E von Canical, SE-exponierte Vulkanitschrofen N ober der Ponta do
Buraco, c. 50 m, 32°44'30''N/16°41'W; an Vertikalflächen, auf Opegrapha spec. (th.),
13.2.1990, J. Hafellner 28211 & A. Hafellner (GZU). – Madeira: N-Küste, 4 km W Sao
Vicente an der Straße nach Seixal, N-exponierte Abbrüche ober der Straße, c. 20 m,
32°48'30''N/17°05'W; Vulkanit, auf Dirina massiliensis var. sorediata (th.), 21.2.1990, J.
Hafellner 27680 & A. Hafellner (GZU). – Madeira, N-Küste, 4 km W Sao Vicente an der
Straße nach Seixal, N-exponierte Abbrüche ober der Straße, c. 20 m,
32°48'30''N/17°05'W; Vulkanit, 21.2.1990, J. Hafellner 27680 & A. Hafellner (herb.
Hafellner). – Tunisia: Halbinsel Bon, SW of Kap Bon NW of El Haouaria, Ghar el
Kebir, Hänge um den antiken Sandsteinbruch, stark windbeeinflusste Gesteinsfluren, auf
Dirina massiliensis (th.), 18.4.1982, J. Poelt (GZU).
Lecanographa rinodinae (Vězda) R.Sant. (all on thalli of Phaeorrhiza nimbosa)
Austria: Salzburg, Hohe Tauern, Kapruner Tal, Schmiedingerkees oberhalb der
Krefelder Hütte, S-exponiert an Kalkschiefer über Moos, c. 2300 m, 6-7.9.1973, K. Kalb
138a (GZU). – Kärnten, Nationalpark Hohe Tauern, Glockner-Gruppe, NW-Grat des
Großen Margrötzen Kopfs W ober dem Hochtor, knapp SW unter dem Grat,
[47°05’10’’N / 12°50’05’’E], c. 2620 m, GF 8943/1; Kalkschiefer, über Moosen und
Pflanzenresten, 5.8.1996, J. Hafellner 38105 & H. Wittmann (herb. Hafellner). –
Norway: Tröndelag, Dovrefjell, Vistradal SE of Driva Kro, Gem. Oppdal, Hänge und
Felsabbrüche des Ryphuskollen über Ryphusseter, c. 1200 m, 4.9.1976, A. Buschardt &
J. Poelt (GZU). – Slovakia: Tatra Magna, montes Belanské Tatry, Ždiarska Vidla, c. 200
m, ad saxa calcarea, Aug.1971, A. Vězda (GZU).
Opegrapha centrifuga A.Massal. (type species of Lecoglyphis Clem.)
Austria: Kärnten, Gailtaler Alpen, unterste Abhänge des Dobratsch, am sog. Gailsteig,
zwischen Tscheltschnigkogel und Pungart, 600-800 m, auf Verrucaria spec. (th.),
Feb.1975, J. Poelt (GZU). – Kärnten, Karawanken, c. 7 km SSE von Villach,
Kanzianiberg S von Finkenstein, am W-Hang des Hügels, 46°33’10’’N / 13°52’25’’E, c.
700 m, GF 9449; Kalkschrofen, auf Verrucaria spec. (th.), 6.5.1992, J. Hafellner 41641
(herb. Hafellner). – Steiermark, Nördliche Kalkalpen, Hochschwab-Gruppe, Jassing NW
von Tragöß, am Eingang des Laminggrabens, 47°33’10’’N / 15°01’30’’E, c. 920 m, GF
8456/1; Fichtenwald, auf kleinen Kalkblöcken an der Böschung, auf Verrucaria spec.
(th.), 26.10.1996, J. Hafellner 38671 & I. Martínez (GZU). – Italy, [Friuli-Venezia
Giulia], Umgebung von Triest, bei Borgo Grotta Gigante, c. 200 m, Karstheiden und
Flaumeichengebüsch, an Kalkblöcken, auf Verrucaria spec. (th.), April 1985, P. L. Nimis,
J. Poelt & L. Poldini (GZU). – [Friuli-Venezia Giulia], Umgebung von Triest, Monte
Stena am Ostrand des Val Rosandra, c. 440 m, Karstheide und Kalkfelsen, auf Verrucaria
spec. (th.), April 1985, P. L. Nimis, J. Poelt & T. Wraber (GZU).
Opegrapha lamyi (O.J.Rich. ex Nyl.) Triebel
Europe: France, [Charente-Maritime] “Charente-Inférieure“, à Marennes, sur les frènes
[Fraxinus], on Lecanora spec. (th., apoth.), Oct.1879, O. J. Richard (GZU). – Spain:
Balearen, Mallorca, Coll de Sóller, Abhänge der Sierra de Alfabia über der Passhöhe, c.
500-550 m, auf Lecanora spec. (th., apoth.), 16.5.1985, J. Poelt (GZU). – North

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America, U.S.A.: Minnesota, Lake Co., 2 miles S of Kawishiwi Lake, 25 miles NW of
Tofte, low rock outcrops with pines, on bark of unnamed tree, on Lecanora spec. (th.,
apoth.), 29.6.1974, C. M. Wetmore 22305b (herb. Hafellner).
Opegrapha parasitica (A.Massal.) H.Olivier (type species of Leciographa
A.Massal.) (all on thalli of Aspicilia calcarea)
Europe: France: Dept. Bouches-du-Rhône, Montagne Ste. Victoire E von Aix-en-
Provence, W-Flanke N vom Refuge P. Cézanne, 500 m, Kalk, 16.5.1980, J. Hafellner
8336 (GZU). – Ibid., J. Hafellner 8401 (herb. Hafellner). – Dept. Bouches-du-Rhône,
Montagne Ste. Victoire E von Aix-en-Provence, Barrage de Bimont, am rechten Ufer des
Stausees, 370 m, Kalk, 16.5.1980, J. Hafellner 8421 (GZU). – Greece: Peloponnes,
Ostküste, Monembasia, NW-exp. Felswände auf einer kleinen Halbinsel etwa 5 km N
von Monembasia, c. 40 m, Kalk, 1.8.1976, H. Mayrhofer (GZU). – Dodekanes, Insel
Simi (Symi, Symä), N-exp. Kalkfelsen am Kloster Panormitis, c. 5 m, 7.9.1983, J. Poelt
(GZU).– Italy: Toscana, prov. Pisa, Monte Pisano, unterste W-exp. Hänge SE von S.
Giuliano Terme bei der Ortschaft Asciano, c. 20 m, Kalk, 28.10.1979, J. Hafellner 4643
& S. Titze (GZU, herb. Hafellner). – Calabria, prov. Cosenza, Golfo di Policastro, Isola di
Dino N von Scalea, c. 100 m, Kalk, 1.6.1979, H. Mayrhofer (GZU). – Ibid., J. Hafellner
41655 (herb. Hafellner). – Sardinia, prov. Nuoro, Monte Albo, NW-Abbrüche der Punta
Cupetti S von Cantoniera di S. Anna, c. 900 m, Kalk, 25.7.1985, P. L. Nimis & J. Poelt
(GZU). – Spain: Balearen, Mallorca, Bergrücken am NW-Rand von Paguera SE von
Andraitx, 200-270 m, Kalkfelsen, 14.5.1985, J. Poelt (GZU).
Opegrapha phaeophysciae R.Sant., Diederich, Ertz & Christnach
North America, U.S.A.: Minnesota, Pine County, 9 miles E of Askov and 13 miles ENE
of Sandstone, mixed hardwood, on Phaeophyscia hispidula (th.), 28.7.1983, J. P.
Schuster (herb. Hafellner 41654).
Opegrapha pulvinata Rehm ex Lojka (all on thalli of Dermatocarpon miniatum
unless otherwise stated)
Europe: Albania: Northern Albania, Kukës distr., near the village Bicaj c. 11 km S of
Kukës, 41°59'N / 20°25'E, c. 300 m; mixed forest with limestone outcrops, on calcareous
rocks, 4.6.2001, L. Kashta (GZU). – Greece: Peloponnes, Mistas bei Sparta,
Felsabbrüche im Klostergebiet, 12.4.1971, J. Poelt 10286 (GZU). – Austria: Tirol,
Osttirol, Hohe Tauern, Virgental, felsiger Rücken der Burg bei Obermauern, 1360-1400
m, auf Dermatocarpon spec. (th.), 1.9.1988, J. Poelt (GZU). – Salzburg, Nördliche
Kalkalpen, Hochkönig Massiv NE von Dienten, W-Hänge der Taghaube, am Steig ober
der Erich-Hütte, c. 1650 m, 47°24’10’’N / 13°02’50’’E, GF 8544/3; subalpine Weide
zwischen Pinus mugo-Beständen, auf Kalkblöcken, 29.8.1996, J. Hafellner 47356 (herb.
Hafellner). – Kärnten, Nationalpark Hohe Tauern, Glockner-Gruppe, Sattelalpe W von
Heiligenblut, c. 1 km SE der Bricciuskapelle, [47°03’01’’N / 12°48’10’’E], c. 1600 m,
GF 8942/2, auf Ca-haltigen Blöcken, 22.11.1987, J. Hafellner 20961 & M. Walther (herb.
Hafellner). – Kärnten, Hohe Tauern, [Goldberg-Gruppe], Mallnitz, E-exponierte Hänge
W des Bahnhofs, 46°59’N / 13°10’30’’E, 1180 m, GF 9045/1; auf einer Lesesteinmauer,
3.11.1986, J. Hafellner 15242 & G. Hofer (GZU), duplicates distributed in Santesson,
Fungi Lichenicoli exs. 114. – Kärnten, Metnitztal, Metnitz, Felswand am Fußsteig
zwischen Kirche und Straße, c. 860 m, GF 9051/1, auf basischem Silikat, auf
Dermatocarpon miniatum var. cirsodes (th.), 18.8.1988, S. Wagner (GZU). – Steiermark,
Murberge, 1.5 km NW von Teufenbach, 700 m NE von Pux, beim Puxer Loch,
47°08’17’’N / 14°20’57’’E, GF 8852/3, Marmorfelsen, 1.5.1994, J. Poelt, H. Pittoni &

91
W. Obermayer (GZU). – Steiermark: Seetaler Alpen, Zirbitzkogel-Massiv SW von
Judenburg, SE-exponierte Hänge des Rückens zwischen Speikkogel und Kreiskogel,
[47°05’30’’N / 14°33’35’’E], c. 1900 m, GF 8953/1; teilweise stark Ca-hältige
Amphibolite, auf Steil- und Neigungsflächen, 4.8.1990, J. Hafellner 26192 & W.
Obermayer (GZU). – Steiermark, Grazer Bergland, S-Hänge des Gamskogel W von
Kleinstübming, ca. 450 m, GF 8857, Dolomitschrofen im lichten Föhrenwald, auf
Dermatocarpon spec. (th.), 31.5.1981, J. Poelt (GZU). – Steiermark, Steirisches
Randgebirge, Grazer Bergland, Bergrücken NE vom Hörgasgraben, c. 3.5 km N von Stift
Rein, Kaschlsteig, 47°10’00’’N / 15°16’45’’E, c. 750 m, GF 8857/2; niedere
paläozoische Kalkschrofen im lichten Föhrenwald, auf Neigungsflächen, 24.12.2000, J.
Hafellner 53769 (GZU). – Italy, Trentino-Alto Adige, prov. Bolzano (Südtirol): Central
Alps, Ötztal Alps, Val Venosta (Vinschgau), hill S of Tárces (Tartsch) = Tartscher Bichl,
NE of Glorenza (Glurns), 46°40’35’’N / 10°33’40’’E, c. 1000 m; low outcrops of
siliceous schist in dry, steppe-like vegetation on slope exposed to the SW, on steep rock
faces, 3.9.2002, J. Hafellner 61292 (GZU). –Trentino-Alto Adige, prov. Bolzano
(Südtirol): [Central Alps, Ötztal Alps], Val Venosta (Vinschgau), Córzes (Kortsch) W of
Silandro (Schlanders), surroundings of St. Ägydius chapel N above the village,
46°38’00’’N / 10°45’45’’E, c. 900 m; outcrops of siliceous schist in dry, steppe-like
vegetation on slope exposed to the S, on steep rock faces, 3.9.2002, J. Hafellner 61252
(GZU). – [Trentino-Alto Adige], Südtirol, Vinschgau, Felsen am Eingang ins
Schlandrauntal bei Schlanders, 23.7.1976, J. Poelt (GZU). – Norway, Oppland, Vaga, W
von Vagamo, Abbrüche von Gneisfelsen, 5.9.1976, A. Buschardt & J. Poelt (GZU). –
Sweden, Södermanland, Sorunda par., Stora Wika, 58°56’N / 17°47’’E, on a seepage on
a grantitic rock, on Dermatocarpon intestiniforme (th.), 15.5.1945, R. Santesson (GZU) =
Santesson, Fungi Lichenicoli exs. 190. – Asia, Afghanistan: Prov. Kabul, Paghman
Gebirge, oberhalb des Ortes Paghman, c. 2540 m, Silikatfelsen, 10.4.1977, D. Podlech
57/3a (GZU, herb. Hafellner). – Prov. Kabul, Paghman Gebirge, oberhalb des Ortes
Paghman, c. 500 m unterhalb der Talgabelung Chap-Darrah und Rast-Darrah, 34°37'N /
68°56'E, c. 2540 m, Silikatfelsen und ergefüllte Spalten, 30.4.1977, D. Podlech 57/3
(GZU). – North America, Mexico: Sonora, 33 km SW of Moctezuma, S of the road to
Hermosillo, c. 950 m, 29°38'N/109°56'W; gentle N-facing slope above the stream, open
oak forest, on low NE-facing cliffs of basaltic rock, on Dermatocarpon americanum (th.),
10. 2.1993, J. Hafellner 37658 & A. Hafellner (GZU), duplicates distributed in
Santesson, Fungi Lichenicoli exs. 273. – Sonora, c. 27.5 km ENE of Ures, 620 m,
29°31'N/110°09'W; open shrubland on N-facing slope, on N-facing granitic outcrops, on
Dermatocarpon spec. (th.), 10.2.1993, J. Hafellner 37666 & A. Hafellner (GZU). –
U.S.A.: Utah, Weber Co., South Fork of Ogden River, Magpie Picnic area, 5500 ft., on
quarzite, on Dermatocarpon reticulatum (th.), 18.6.1956, L. Behling 143a (GZU). –
Arizona, Coconino Co., Grand Canyon National Park, slopes exposed to the N along
Grand View Trail below Grand View Point, c. 2000 m, 36°00'N/111°59'W; rocky slope
with scattered trees of Pinus edulis, on cliffs of calcareous sandstone, on Dermatocarpon
spec. (th.), 10.7.1994, J. Hafellner 36900 (GZU). – Arizona, Apache Co., Apache
National Forest, valley of Black River, surroundings of the bridge of road FR 25, c. 2100
m, 33°42'20"N/109°27'W; remnants of oak forest and cliffs above the riverbank, on cliffs
exposed to the S, on Dermatocarpon spec. (th.), 5.7.1994, J. Hafellner 36844 (GZU).
Opegrapha sphaerophoricola Isbrand & Alstrup (all on thalli of Sphaerophorus
globosus)
Europe, Spain: Asturias, Prov. Oviedo, Parque Nacional de Muniellos N von Cangas de
Narcea, 650 m, Wald im Talschluß mit Silikatblöcken; 4.9.1980, J. Hafellner 10217
(herb. Hafellner). – Africa, Canary Islands: La Gomera, Garajonay, in der Mulde N
unter dem Gipfel, c. 1360 m, 28°06'15"N/17°14'10"W; alter Erica-Wald, auf Borke von

92
Erica arborea, 18.2.1991, J. Hafellner 32815 & A. Hafellner (GZU). – La Gomera: Alto
de Cherelepin N ober der Straße von San Sebastian nach Valle Gran Rey, c. 1320 m,
28°06'50"N/17°15'30"W; Erica-Myrica-Buschwald, auf Borke von Erica arborea,
10.2.1991, J. Hafellner 32785 & A. Hafellner (herb. Hafellner). – North America,
U.S.A.: Oregon, Umgebung von Eugene, Hardesty Mountain, 26.5.1985, E. Kordesch
(herb. Hafellner 13383).
Plectocarpon lichenum (Sommerf.) D.Hawksw. (all on thalli of Lobaria
pulmonaria if not otherwise stated)
Europe, Albania: Central Albania, Tiranë distr.: on the mountain Dajt E above the town
Tiranë, between the restaurant “Panorama” and the saddle Qafa e Qershisë, 41°22'15"N /
19°55'20"E, c. 1100 m, deciduous forest over limestone on slope exp. to the W, on bark
of Acer opalus, 20.8.2007, J. Hafellner 71008 (together with M. Tretiach, L. Muggia, M.
Piccotto & J. Marka) (GZU). – Austria: Oberösterreich, Sengsengebirge, Feichtausee bei
Molln, an Acer pseudoplatanus, 9.5.1983, G. Gärtner (GZU). – Salzburg, Hohe Tauern,
Glockner Gruppe, Kapruner Tal, Kesselfall, NW-Hänge, 47°13’N / 12°43’45’’E, c. 1040
m, GF 8742/3; bergahornreicher Mischwald, auf Borke von Acer pseudoplatanus,
26.8.1996, J. Hafellner 47257 (herb. Hafellner). – Tirol, [Außerfern], an der Straße von
Plansee nach Griessen, c. 1000 m, 16.8.1953, A. Schröppel (GZU). – Tirol, Karwendel,
Tortal bei Hinterriß, Aug.1962, J. Poelt 1403 (GZU). –France: Dept. Puy-de-Dôme:
Auvergne, Monts Dore, Bois du Domais W von Super-Besse, c. 1300 m; Fagetum im W-
exponierten Talschluß, 31.7.1980, A. Bellemère & J. Hafellner 9537 (herb. Hafellner). –
Korsika, Dept. Haute-Corse: An der Straße von Vivario auf den Col de Vizzavona, c. 1
km NE von Tattone, c. 830 m; a) Kastanienhain in einer Weide, auf Astborke von
Castanea sativa b) an einer steinigen Straßenböschung; 3.11.1993, H. Mayrhofer & E.
Unger (GZU). – Italy: Basilicata, Prov. Potenza, NW-Abhänge der Coppola di Paola SE
von Rotonda, c. 1400 m; Kalkhangbuchenwald, 31.5.1979, J. Hafellner (herb. Hafellner).
– Ibid., J. Poelt (GZU). – Montenegro: Paß zwischen Andrijevica und Babljak, Anstieg
von der Paßhöhe zum Fuß des Kucki kom., c. 1650 m, Buchenwald, 18.7.1974, J.
Hafellner 586 (herb. Hafellner). – Spain: Asturias, Prov. Oviedo, Abhänge des Cordal de
la Mesa N unter dem Puerto de Ventana, c. 1340 m; Fagetum mit einzelnen
Silikatblöcken, 1.9.1980, J. Hafellner 10501 (GZU). – Asturias, Prov. Oviedo, Parque
Nacional de Muniellos N von Cangas de Narcea, 650 m, Wald im Talschluß mit
Silikatblöcken, 4.9.1980, J. Hafellner 10228 (herb. Hafellner). – United Kingdom:
Scotland, Westerness, Morvern, N of Glenmorvern Cottage, hazelwood, c. 10 m,
16.6.1992, B. J. Coppins, P. W. James & J. Poelt (GZU). – Africa, Tunisia: NW-
Tunesien, Les Chênes 7 km S von Ain Daraham, 730-760 m, Quercus faginea-Q. suber-
Wald, an Quercus, 11.4.1968, J. Poelt 5865 (GZU). – Canary Islands: La Gomera, La
Montañeta E ober El Cedro, am Höhenrücken SW vom Gipfel, 28°07'30"N / 17°12'W, c.
1020 m, Erica-Buschwald mit Stockausschlag von Laurus, auf Zweigen von Laurus
canariensis, 18.2.1991, J. Hafellner 33826 & A. Hafellner (herb. Hafellner). – Tenerife,
Bosque de la Esperanza, c. 4 km E unter Las Lagunetas, 28°25'30"N / 16°24'W, c. 1220
m, Kanarischer Kiefernwald mit Ilex canariensis-Unterwuchs, an dicken Ästen von Pinus
canariensis, 22.12.1994, J. Hafellner 33751 (GZU). – North America, Canada: British
Columbia, Wells Gray Park, Dawson Falls, forest of lower oroboreal subzone, in spray
zone of waterfall, 51°58’N / 120°07’W, c. 775 m, 25.8.1994, T. Goward & J. Poelt
(GZU). – U.S.A.: North Carolina, Alleghany Co., Blue Ridge Parkway, near Lowgap,
deciduous forest, 25.3.1970, W. Culberson & J. Poelt (GZU).

93
Pseudotryblidium neesii (Flot.) Rehm (all on bark of Abies alba)
Europe, Austria: Kärnten, Karnische Alpen, SE-Abhänge der Treßdorfer Höhe, c. 200 m
W der Straße auf das Naßfeld, unter der Forststraße c. 500 m N des Bodensees,
46°35’30’’N / 13°16’05’’E, c. 1040 m, 14.9.1994, M. Magnes 2061 & A. Wippel (GZU).
– Kärnten, Steiner Alpen, Vellacher Kotschna S von Eisenkappel, [46°23’45’’N /
14°34’10’’E], c. 1000 m, GF 9653/1, Mischwald über Dolomitschotter, 4.7.1990, J.
Hafellner 26853 & W. Petutschnig (GZU). – Steiermark, Nördliche Kalkalpen, Totes
Gebirge, Tauplitzalm-Gebiet, an der Straße von Bad Mitterndorf auf die Tauplitzalm, bei
der 1. Kehre, 47°34’35’’N / 13°58’10’’E, c. 1170 m, GF 8449/2; Buchen-Tannen-
Fichtenwald, 21.8.1988, J. Hafellner 50565, E. Schreiner & A. Hafellner (GZU). –
Niederösterreich, Nördliche Kalkalpen, Seetal SE von Lunz, kurz S hinter dem Mittersee,
[47°49’30’’N / 15°04’40’’E], c. 780 m; Tannen-Bergahornwald, 20.6.1981, J. Hafellner
9198 (GZU). – Steiermark, Steirisches Randgebirge, Koralpe, Reinisch Kogel NW von
Stainz, am rechten Ufer des Fallegg Baches, bei einer verfallenen Mühle S unter der
Sommereben, [46°56’25’’N / 15°09’55’’E], c. 940 m, GF 9056/4, Tannen-Fichtenwald;
22.5.1984, J. Hafellner 11281 (GZU). – Italy: Friuli - Venezia Giulia, Prov. Udine,
[Südalpen], Karnische Alpen: an der Straße vom Lago di Sauris auf den Paso del Pura,
NW von Ampezzo, [46°25’55’’N / 12°43’55’’E], c. 1320 m; Buchen-Tannen-
Fichtenwald, 26.7.1993, J. Hafellner 32610 (GZU).
Results
Phacothecium Trevis., Linnaea 27: 298 (1856).
Type: Phacothecium varium (Tul.) Trevis. (Phacopsis varia Tul.) (holotype)
Description: Ascomata black, discs irregular in shape, but mostly ± elongated.
Exciple carbonized, thin throughout, basally closed. Ascomatal development
starting as closed plectenchymatal globe, distinctly hemiangiocarpous. Hymenia
exposed by irregular rupture of the obtecting layer of excipular tissue, finally with
more or less broadly exposed discs, becoming lobed to divided with age.
Hymenium with hemiamyloid gel. Asci arthonialean, with stipe tapering towards
the base, with non-amyloid ascal wall, endoascus without a reactive apical ring,
dehiscence fissitunicate. Ascospores hyaline, with at least 3 septa, with rounded
ends, perispore thin, condensing with age to brown granules which constitute a
rugulose sculpture. So far monotypic, lichenicolous.
Discussion: The genus Phacothecium was introduced for a single species
described by Tulasne some years earlier, Phacopsis varia Tul. (TREVISAN 1856).
Phacothecium was commonly reduced to the synonymy under Arthonia Ach.,
however, as seen from to the phenotypic characters of the type species, it is
clearly closer to Opegrapha than to Arthonia.
ERTZ et al. (2004) argued that the type species Phacothecium varium (as
Opegrapha physciaria) may belong in the species group which later (ERTZ et al.
2005) was called the Opegrapha anomea-group. This concept is not followed
here. Although superficially similar, O. anomea is evidently not congeneric with
Phacothecium varium, an opinion which is supported by phenotypic characters
such as fine structure of the ascus wall and its biochemistry.

94
The species
Phacothecium varium (Tul.) Trevis., Linnaea 27: 298 (1856).
Bas.: Phacopsis varia Tul., Ann. Sci. Nat., Bot., sér. 3., 17: 125 (1852).
Syn.: Celidium varium (Tul.) A.Massal., Misc. Lich.: 44 [15] (1856). – Arthonia varia
(Tul.) Jatta, Syll. Lich. Ital.: 471 (1900) non (Ach.) Nyl. (1857) nec Opegrapha varia
Pers.
Type: “Crescit in Parmelia parietina...,…e Mauritania, ubi in truncis Cacti Opuntiae
creverant, a cl. Duriaeo relata sunt” (locality information from protologue) PC-, STR-.
Type specimen lost. Ann. Sci. Nat., Bot., sér. 3., 17: planche XIV, figs 1-3, 1852
(lectotype – designated here). Epitype: Canary Islands: Lanzarote, on the margin of
Malpais-plateau short SW of the village Guiguan (SW of Tinajo), 29°02’2’’N /
13°41’5’’W, 280 m, on branches of Ficus carica, on Xanthoria parietina, 9.4.1999, J.
Hafellner 47633 & A. Hafellner, distributed in Santesson, Fungi lichenicoli exs. no. 325
(GZU – epitype, designated here! Duplicates of epitype in herbaria receiving the
mentioned exsiccate, e.g. in UPS, M).
Note: No authentic material could be traced in PC or STR (the curators, in litt.).
As the lectotype designated above is demonstrably ambiguous (the depicted
details of hymenial elements would fit to a number of lichenicolous Opegrapha
species) and therefore cannot be critically identified for purposes of the precise
application of the name Phacopsis varia, an epitype is designated. No specimen
from Mauritania or Morocco was available, therefore a representative specimen
from nearby Lanzarote was chosen as epitype. The epitype specimen fits perfectly
with the detailed protologue.
Syn. Lecidea physciaria Nyl., Suppl. Lich. Paris: 8 (1897). – Leciographa physciaria
(Nyl.) H.Olivier, Bull. Acad. Int. Geogr. Bot. 16: 48 (1906). – Opegrapha physciaria
(Nyl.) D.Hawksw. & Coppins in Coppins et al., Lichenologist 24: 367 (1992).
Type: “France, Paris, Ormes de Carel, on Xanthoria parietina, 1827, Alph. de Brebisson
n. 814” (H-Nyl 11023 – lectotype!). Lectotype designated by ATIENZA (1992: 160).
Note: The lectotype specimen contains a second lichenicolous fungus, i.e. an
ascomycete with crowded perithecioid ascomata developed on the apothecial
discs. The sample is too scanty for a critical study.
Syn. Mycobilimbia xanthoricola Räsänen, Ann. Bot. Soc. Zool.-Bot. Fenn. „Vanamo“ 21,
Notul. Bot. 16: 2 (1946).
Type: Finland, Alandia, Geta, Östergeta, på vild ask vid Langnäs, on Xanthoria
parietina, 27. VIII. 1945, leg. L. Fagerström (H – lectotype!). Lectotype designated by
HAWKSWORTH & ATIENZA (1994: 49).
Icon.: TULASNE 1852: tab. XIV, figs 1-3 (drawings of portion of hymenium, ascospores,
conidiogenous cells with microconidia); ATIENZA 1992, figs 1H-N (drawings of asci in
various stages of development, ascospores, microconidia); this publication, Fig. 1
(longitudinal section of ascoma), Fig. 2 (asci).
Exs.: Santesson, Fungi lichenicoli exs. no. 325 sub Opegrapha physciaria (GZU);
Arnold, Lich. exs. 335a, b (as Celidium varium) (n. v., see Triebel & Scholz 2001);
Rabenhorst, Lich. Europ. exs. 785 (as Celidium varium) (n. v., see Triebel & Scholz
2001); Hafellner, Lichenicolous Biota no. adhuc ined.

95
Figs 1-2. Phacothecium varium, unstained, medium: water. Fig. 1. Ascoma in
longitudinal section (from Hafellner 46494). Fig. 2. Ascus, DIC (from Hafellner 47633).
Description (see also ATIENZA 1992): Ascomata black, ± round when young and
still closed, discs exposed by a breakdown of the obtecting layer, becoming lobed
to divided with age (?centrum oligolocular), resulting in dense clusters of 1-5
discs divided by sharp excipular margins. Exciple thin and basally closed, dark
brown throughout, in longitudinal section laterally 20-30 μm in diam., a radial
arrangement of the hyphae constituting the exciple not observable, no spectacular
reaction with K, only somewhat more blackish (“Atra-brown” sec. MEYER &
PRINTZEN 2000). Hypothecium hyaline to pale brownish, 30-50 μm high,
composed of hyphae with ± isodiametric cells (cells not distinctly elongated).
Hymenium hyaline, including the epihymenium 70-80 μm high, K–; hymenial gel
I+ blue then red, K/I+ blue; epihymenium brownish, K–; Asci broadly
subcylindrical to subclavate, tapering towards the base and distinctly stipitate,
8(-6-4) spored, 45-60 × 16-20 μm, ocular chamber reduced, wall I-, K/I-,

96
hemiamyloid apical ring structure lacking (Fig. 2). [Asci “...clavate, ..., 45-50 × 18
μm, ..., Opegrapha-type (sensu RENOBALES & BARRENO 1989) ...” sec. ATIENZA
(1992)]. Ascospores hyaline, 3-septate, oblong or slightly tapering towards the
lower end, (12-)14-16 × 5-7 μm. [Ascospore size 12.8-16 × c. 4 μm sec. TULASNE
(1852), 12-17 × 6 μm sec. Nylander’s annotations on the type specimen of
Lecidea physciaria, 10-16 × 5-7 μm sec. Räsänen’s annotations on the type
specimen of Mycobilimbia xanthoriae, (12-)15-16 × 6-7(-8) μm and erroneously
given as “3-celled” sec. ATIENZA (1992)]. Ascospores with persistently hyaline
proper wall, with distinct but thin perispore sheath, giving rise to a pale-brown
verruculose sculpture with age. Conidiomata frequently present in between groups
of ascomata, ellipsoid to spherical, unilocular, formation of microconidia
terminally on small conidiogenous cells incorporated in the condiomatal wall,
microconidia rod-shaped, 4-7 × 0.7-1 μm. [Pycnospore size mostly not longer
than 4.5 μm sec. TULASNE (1852), 6-7.5 × 1 μm sec. ATIENZA (1992)].
Hosts: Xanthoria parietina (th.) (1), X. calcicola (th.) (2), X. aureola (X.
ectaneoides) (th.) (3), X. parietina f. rutilans (th.) (4), X. spec. (th.) (5). Reports
on other hosts such as Caloplaca cirrochroa (LOJKA 1876), C. rosei (COLE &
HAWKSWORTH 2001), C. saxicola (JATTA 1882) and Lecanora dispersa, etc.
(ROUX 1977) are urgently in need of revision. In the literature cited below also
homotypic synonyms of the listed hosts may have been used.
Ecology: Phacothecium varium infests species of Xanthoria in a wide range of
habitats. When growing upon epiphytic X. parietina, the sites include roadside
trees (there also on canopy branches), groups of deciduous trees in rural
environments, but also open deciduous forests (e.g. ATIENZA 1992) and
Mediterranean shrub communities mixed with invasive Opuntia populations. It is
there where the lichenicolous fungus comes closest to its second host species the
epilithic Xanthoria calcicola.
Distribution: The species is widely distributed in the northern hemisphere from
the arid subtropical-Mediterranean zone northwards to the northern edge of the
temperate zone. The localities known so far are situated from sea level up the
lower montane vegetation belt. It is already known from numerous countries, but
various synonyms, namely Opegrapha physciaria or homotypic synonyms (A),
Celidium varium or homotypic synonyms (B), Mycobilimbia xanthoriicola (C),
may have been used. So far it has been recorded from the following European
countries: Albania (this publication), Austria (POETSCH & SCHIEDERMAYR 1872,
as (B) on (1), SIMMER 1900, as (B) without host, HAFELLNER 1994b, as (A) on
(1), NAVROTSKAYA et al. 1996, as (A) on (1), HAFELLNER & OBERMAYER 2001,
as (A) on (1)), Croatia (this publication), Czech Republic (KUTAK 1910, as (B) on
(4)), Finland (RÄSÄNEN 1946, as (C) on (1)), France (NYLANDER 1857, as Lecidea
glaucomaria [sic] on (1), OLIVIER 1884, as (B) on (1), NYLANDER 1897, as (A) on
(1), OLIVIER 1903, 1906, 1907, as (A) on (1), VOUAUX 1913, as (A) on (1),
BOULY DE LESDAIN 1914, as Celidium varians [sic] on (1), VOUAUX 1914, as (B)
on (1), LETTAU 1958, as (B) on (1), RONDON 1969, as (A, B) on (1), ROUX et al.
2006, as (A) on (1)), Germany (ARNOLD 1868, as (B) on (1), BAUSCH 1869, as
(B) on (1), REHM 1891, as (B) on (1), VOUAUX 1914, as (B) on (1), KEISSLER
1930, as (B) on (1), LETTAU 1958, as (B) on (1), TRIEBEL & SCHOLZ 2001, as (A)
on (1)), Ireland (HAWKSWORTH 1994, as (A) on (1)), Italy (JATTA 1882, 1889,

97
1909, all three as (B) on (1)), Poland (STEIN 1872, 1879, both as (B) on (1),
BOBERSKI 1886, as (B) on (1), LETTAU 1912b, as (B) on (1), VOUAUX 1914, as
(B) on (1), KEISSLER 1930, as (B) on (1)), Portugal (NAVROTSKAYA et al. 1996, as
(A) on (2)), Slovacia (LISICKÁ & LACKOVICOVÁ 1999, as (B) without host), Spain
(SANTESSON 1960, as (A) on (1), GIRALT & GOMEZ-BOLEA 1988, as (A) on (1),
ATIENZA 1992, as (A) on (1), NAVARRO-ROSINÉS et al. 1994, as (A) on (2),
CALATAYUD et al. 1995, as (A) on (1), GIRALT 1996, as (A) on (1), ATIENZA &
SEGARRA MORAGUES 1999, as (A) on (1), FOS & BARRENO 2002, as (A) on (1)),
Sweden (SANTESSON 1949, as (A) on (1), SANTESSON 1993, as (A) on (1)),
Ucraine (KONDRATYUK & KHODOSOVTSEV 1997, as (A) on (1)), United Kingdom
(HAWKSWORTH 1994, as (A) on (1), NAVROTSKAYA et al. 1996, as (A) on (1, 3),
HITCH 2006, 2007a, both as (A) on (1)), and the archipelagos Aegadian Islands
(NIMIS et al. 1994, as (A) on (1)), Balearic Islands (ETAYO 1996, as (A) on (1),
BOOM 1999, as (A) on (1)), Corfu (KEISSLER 1914, as (B) on (1)), Ischia (JATTA
1892, as (B) without host), and SICILY (Grillo et al. 2000, as (A) without host).
From Asia it has already been reported from Cyprus (LITTERSKI & MAYRHOFER
1998, 2000, both as (A) on (1)) and Israel (NAVROTSKAYA et al. 1996, as (A) on
(1)). In Africa it is so far known from Egypt (MÜLLER ARG. 1880, as (B) on (1),
STIZENBERGER 1890, as (B) on (1)), Mauritania (TULASNE 1852, as (A) on (1)),
Morocco (VOUAUX 1914, as (B) on (1)), and the archipelago Canary Islands
(HAFELLNER 1995, as (A) on (1), HAFELLNER 2005, as (A) on (1, 2)). In North
America it has been reported from northern Mexico (HAFELLNER et al. 2002, as
(A) on (1, 5)).
In need of revision are the records on unusual hosts from France (ROUX 1977,
as (B) on Lecanora dispersa and other lichens), Hungary (LOJKA 1876, as (B) on
Caloplaca cirrochroa), Italy (JATTA 1882, as (B) on Caloplaca saxicola), Ucraine
(KONDRATYUK & KHODOSOVTSEV 1997, as (A) on Xanthoria lobulata) and the
U.S.A. (COLE & HAWKSWORTH 2001, as (A) on Caloplaca rosei).
Discussion: The species was first described by TULASNE (1852) as Phacopsis
varia Tul. This epithet is not available in Opegrapha because of Opegrapha varia
Pers. Therefore for long time Opegrapha physciaria based on the heterotypic
synonym Lecidea physciaria Nyl. was used in the literature, whereas when
classified in Celidium, the name C. varium (Tul.) A.Massal. was applied to the
same lichenicolous fungus. In Phacothecium, there is no obstacle to using the
oldest legitimate name.
The species was treated in some detail by ATIENZA (1992 as Opegrapha
physciaria). KEISSLER (1930) treated the species as Celidium varium in the
subgenus Celidiopsis (A.Massal.) Sacc. The taxon Celidiopsis A.Massal. was
originally published at genus level to accommodate two lichenicolous fungi,
Celidiopsis gyrolophii A.Massal. (on Dictyonema glabratum as Gyrolophium
mauritianum) and Celidiopsis insitiva (Körb.) A.Massal. (MASSALONGO 1856).
No material of these two species was available for study, but it is worth
mentioning that when Massalongo combined Phacopsis varia Tul. into Celidium
he had some doubts of this placement by adding a question mark. Although to our
knowledge Celidiopsis so far has not been lectotypified (lectotype designated
below), the genus is treated as synonym of Arthonia in more recent outlines of
ascomycetes (e.g. ERIKSSON & HAWKSWORTH 1998) and systematic databanks
(e.g. Mycobank, checked 30 April 2008).

98
Species with asci that lack a hemiamyloid ring structure in the endoascal wall
layer are relatively rare in Opegrapha and closely related genera; they are more
common in Arthonia. Therefore, the ascus typology presented by TORRENTE &
EGEA (1989) is hardly applicable. The shape of the asci of Phacothecium varium
was perfectly drawn by ATIENZA (1992), with a relatively long stipe as
characteristic feature (compare Fig. 2 in this publication). They were called
“Opegrapha-type (sensu RENOBALES & BARRENO 1989) ...” by ATIENZA (1992),
however, an “Opegrapha-type” was not defined by RENOBALES & BARRENO
(1989) who pointed out the considerable variability of ascal characters within the
genus Opegrapha.
Further specimens seen: Europe: Albania: Southern Albania, Vlorë distr.: at the coast of
the Adriatic Sea NW of the town Vlorë, small island of the monastery “Fjetja e
Hyjlideses”, on the W coast, 40°31'00"N / 19°24'00"E, c. 10 m, fringe of evergreen
bushes, on bark of twigs of Pistacia lentiscus, (1), 21.8.2007, J. Hafellner 71009
(together with M. Tretiach, L. Muggia, M. Piccotto & J. Marka) (GZU), distributed in
Hafellner, Lichenicolous Biota no. adhuc ined. – Austria: Salzburg, Lungau, Taurachtal,
Weg zur Hofbauerhütte, c. 1220 m, GF 8847, auf Picea abies, (1), 29.5.1989, S. Roth
(GZU). – Steiermark: Zentralalpen, Murberge, kleiner Taleinschnitt SE unter dem Schloß
Rothenfels, c. 1.5 km SE von Oberwölz, 47°11’50’’N / 14°18’15’’E, c. 860 m, GF
8851/2; Gehölzstreifen am Bachufer, auf Borke von Salix spec., (1), 12.11.2000, J.
Hafellner 53233 (GZU). – Steiermark, Grazer Bergland, an der Straße NW vom Gasthof
Abraham im Stübinggraben, c. 680 m, an alten Apfelbäumen, (1), 27.1.1991, J. Poelt
(GZU). – Steiermark, Grazer Bergland, Auwald im Graben des Tulwitzbaches W
oberhalb von Leisenreith, c. 610 m, an Fraxinus excelsior, (1), together with Telogalla
olivieri, 28.1.1990, J. Poelt (GZU). – Croatia: Halbinsel Istrien, Inselgruppe “Due
Sorelli” [Uvala Valmascin] nahe Rovinj, (1), 20.5.1970, H. Wunder 5207 (GZU). –
France: Hérault, peupliers à Agde, ? du Grace, (1), July 1907, A. de Crozals (GZU). –
Italy: Isole Egadi, Marettimo, between Marettimo and Pizzo del Capraro, 26.3.1991, J.
Poelt (GZU). – Portugal: Algarve, zwischen Praia da Rocha und Praia do Vau, 10-20 m
von den Steilklippen entfernt, c. 30 m, (2), 11.6.1982, H. Wunder 1895 (GZU). – Spain:
Asturias, prov. Oviedo, Cabo de Peñas NE von Avilés, ordovicische Schiefer knapp über
Meeresniveau, (2), 3.9.1980, J. Hafellner 10658 (GZU). – Andalusia, Prov. Almeria,
Cabo de Gata, c. 5 km SW von San José, 20-100 m, Vulkanitblöcke, (2), 22.9.1980, J.
Hafellner 9684 (GZU). – Africa: Canary Islands: Lanzarote: am Rand des Malpais-
Plateaus kurz SW vom Dorf Guiguan (SW von Tinajo), 29°02'20"N / 13°41'55"W, c. 280
m; auf Ästen im Kroneninneren von Ficus carica, (1), 9.4.1999, A. Hafellner & J.
Hafellner 47637 (GZU). – Lanzarote: Guanapay E ober dem Ort Teguise, am NE-Rand
des Kraters, 29°03'30"N / 13°32'55"W, c. 450 m; N-Hänge an der Krater-Außenseite,
bodennah auf Ausbissen aus Vulkanitkonglomerat, (2), 9.4.1999, A. Hafellner & J.
Hafellner 47438 (GZU). – Lanzarote: Los Helechos N ober dem Dorf Maguez,
Bergrücken N des Kraters, 29°10'35"N / 13°29'40"W, c. 400 m, auf Basaltblöcken in
einer Triftweide, (2), 6.4.1999, A. Hafellner & J. Hafellner 47524 (GZU). – Lanzarote:
Peña de la Pequena NE ober dem Dorf Los Valles, El Agujero, im Gipfelbereich,
29°07'20"N / 13°29'50"W, c. 500 m; grobes Blockwerk und niedere Abbrüche aus Basalt
mit Sukkulentenbusch, S-exponiert an Neigungsflächen, (2), 8.4.1999, A. Hafellner & J.
Hafellner 47667, 47668 (GZU), distributed in Hafellner, Lichenicolous Biota no. adhuc
ined. – Lanzarote: Pico Naos N vom Dorf Femés, im Gipfelbereich, 28°55'30"N /
13°44'40"W, c. 400-415 m; grobes Blockwerk und niedere Abbrüche aus Basalt mit
Sukkulentenbusch, SW-exponiert an Neigungsflächen, (2), 7.4.1999, A. Hafellner & J.
Hafellner 47508 (herb. Hafellner). – Fuerteventura: nahe der E-Küste etwa halbwegs
zwischen Puerto del Rosario und Corralejo, sanft geneigte Hänge SE-seitig am Fuß des

99
Vulkans Roja, 28°37'40"N / 13°50'05"W, c. 50 m; Malpais, auf kleinen Lavablöcken, (2),
27.4.2000, A. Hafellner & J. Hafellner 53377 (GZU). – Ibid., (2), Hafellner 53383 (herb.
Hafellner). – Tenerife, Teno-Gebirge, NW unter Teno Alto am Steig hinab zur
Nordküste, hoch oberhalb von Las Casas (kurz E der Punta de Teno),
18°21’05’’N/16°54’20’’W, c. 400 m; Vulkanitschrofen im Sukkulentenbusch, auf
Steilflächen von Basaltfelsen, (2), 16.12.1998 (GZU).– Ibid., (2), 26.4.2005, J. Hafellner
64250 (GZU, herb. Hafellner). – Tenerife: Macizo de Teno, W des Ortes El Palmar an der
Straße nach Teno Alto, c. 600 m, 28°20'N/16°51'W, an freistehenden Ficus carica, (1),
15.2.1989, J. Hafellner 32860 & A. Hafellner (GZU). – La Gomera: sanft geneigte Hänge
W ober Arure, c. 860 m, 28°07'30"N/17°18'20"W; Kulturland, auf Zweigen von Pyrus
communis, (1), 12.2.1991, J. Hafellner 32819 & A. Hafellner (GZU). – North America:
Mexico: Baja California del Norte, Cerro Kenton and adjacent area W of Bahia de San
Quentin, 30°28’N/116°00’W, on low shrubs, (3), 15.2.1987, T. Nash & J. Poelt (GZU). –
Baja California Norte: peninsula SW of San Quintín, surroundings of Cerro Kenton,
small hills SW of the volcano, c. 50 m, 30°28'N/116°W; desert scrub, on branches of
small shrubs, (1), 24.2.1993, J. Hafellner 41099 & A. Hafellner (herb. Hafellner). – Baja
California Sur: coastal plain about 40 km S of Ciudad Constitution by the road (Hwy 1)
to La Paz, at turnoff to Puerto Cancun, c. 30 m, 24°43'N/111°33'W; desert scrub, on
branches of a small tree, (1), 17.2.1993, J. Hafellner 46494 (GZU).
Excluded species
Opegrapha anomea Nyl., Act. Soc. Linn. Bordeaux 21 (3. sér., 1): 399 (1857) aut
Bull. Soc. Bot. France 3: 552 (1857).
Syn.: Leciographa anomea (Nyl.) Branth & Rostr., Bot. Tidskr. 3: 253 (1869). –
Dactylospora anomea (Nyl.) Arnold, Flora 57: 109 (1874). – Mycobilimbia anomea
(Nyl.) Sacc. & D.Sacc., Sylloge Fung. 18: 185 (1906).
Type: [France]; ad abietes prope Mont Dore, Arvernia, on Pertusaria amara, leg. W.
Nylander (H-NYL 6581, lectotype!) Lectotype designated by ERTZ et al. (2004: 236). –
[France]; Mont Dore (M. du Capucin), ad abietes, on Pertusaria amara, leg. W. Nylander
(H-NYL 6580 – paratype!).
Note: Opegrapha anomea was published twice; both publications appeared in
1857 and in both the taxon is given as “n. sp.” It is impossible to decide which
publication appeared earlier and thus represents the protologue.
Heterotypic synonyms according to ERTZ et al. (2004):
Opegrapha quaternella Nyl., Flora 68: 449 (1885).
Leciographa pertusariae Vouaux in Pitard & Harmand, Bull. Soc. Bot. France 58, Mem.
22: 70 (1911).
Opegrapha wetmorei M.S.Cole & D.Hawksw., Mycotaxon 77: 322 (2001).
?Leciographa weissii Körb., Verh. Zool.-Bot. Ges. Wien 17: 707 (1867).
Icon.: ERTZ et al. 2004: 236, Fig. 1-4 (habitus)
Exs.: –
Description (diagnostic characters only, for a detailed description see ERTZ et al.
2004): Ascomata black, epruinose, roundish when young, obtecting stromatic
layer breaking up soon, resulting in irregular slits and splitting off of the covering
layer, finally with widely exposed discs. Exciple in longitudinal section dark

100
brown, K–. Hymenium hyaline, K–, I+ blue then red, K/I+ blue; epihymenium
brownish, K–; Asci broadly cylindrical to subclavate, rather abruptly pedicellate,
4(-8) spored, 35-40 × 8-13 μm, wall I–, K/I– except for an apical ring that is K/I+
blue. Ascospores hyaline, 3-septate, 17-26 × 6.5-9 μm, with distinct perispore
sheath, resulting in a brown verruculose sculpture with age.
Hosts: Pertusaria amara (th.) (1), P. albescens (th.) (2), P. epixantha (th.) (3), P.
exalbescens (th.) (4), P. velata (th.) (5), P. spec. (6), Ochrolechia androgyna (th.)
(7), O. trochophora (th.) (8), O. spec. (9) (in literature cited below partly under
their synonyms)
Ecology: Most of the host species are recorded only from North America (ERTZ et
al. 2004). Although Pertusaria amara in Europe is a very common crustose lichen
on bark, Opegrapha anomea at least in Europe is a rarely collected lichenicolous
fungus.
Distribution: The species is so far known from Europe, Africa and North
America. It has been reported under different synonyms (Opegrapha anomea or
homotypic synonyms (A), O. pertusariae or homotypic synonyms (B), O.
quaternella (C), O. wetmorei (D)) from Denmark (NYLANDER 1866, as (A)
without host, BRANTH & ROSTRUP 1869, as (A) on (6)), France (NYLANDER
1857a, b, as (A) on (1), ETAYO & DIEDERICH 1998, as (A) on (1, 2), ERTZ et al.
2004, as (A) on (1)), Norway (HOLIEN 2001, as (A) on (1)), the United Kingdom
(HITCH 2007b, as (A) on (1)), the Canary Islands (PITARD & HARMAND 1911,
RONDON 1969, both as (B) on (4)), Canada (ERTZ et al. 2004, as (A) on (7, 9)) and
the U.S.A. (NYLANDER 1885, as (C) on (5), COLE & HAWKSWORTH 2001, as (D)
on (8), HAFELLNER et al. 2002 as (B) on (1), ERTZ et al. 2004, as (A) on (1, 3, 6,
9)). Possibly also a record from Croatia (KÖRBER 1867, as Leciographa weissii on
(6) and/or (9)) belongs here.
Discussion: This species was treated in detail by ERTZ et al. (2004) who interpret
agglomerations of ascomata as a result of the centrifugal growth of single
ascomata.
Although the mode of opening of the ascomata in both Phacothecium varium
and Opegrapha anomea seem to happen in a similar way, i.e. starting with a slit-
like rupture followed by a mutual exposure of the hymenium, other characters
(e.g. ascus shape, details in ascus wall construction) indicate that the two species
are not very closely related to each other. Therefore we hesitate to transfer the
species to Phacothecium.
Further species that have been compared with Phacothecium varium, but also
have been found to be not congeneric, are Opegrapha phaeophysciae R.Sant. et
al. (ERTZ et al. 2005) and O. sphaerophoricola Isbrand & Alstrup (ISBRAND &
ALSTRUP 1992).
Together with the detailed treatment of O. anomea by ERTZ et al. (2004), a
further species, O. blakii Ertz & Diederich, was described. As no specimen of this
species has been seen, on basis of the figures published it is hesitatingly
transferred to Phacographa.

101
Phacographa Hafellner gen. nov.
MB 513173
Genus ad familiam Roccellaceae (sensu LUMBSCH & HUHNDORF 2007) pertinens.
Ascomata rotunda haud elongata, marginata, lecideoidea, persistenter unilocularia,
hymenio ab initio aperta aut partim residuis minutis excipuli obdecta. Asci late
subcylindrici, 4- ad 8-spori. Paries ascorum in strato interno anulo hemiamyloideo
instructus. Ascosporae phragmosporae, mature hyalinae. Opegrapha differt ascomatibus
hysterothecioideis. Lecanographa differt ascomatibus subhysterothecioideis hymeniis
albide pruinosis. Phacothecium differt ascomatibus tarde fissuris aperientibus, demum
confluentibus, ascis anulis hemiamyloideis destitutis. Species cognitae fungi lichenicoli
non lichenisati.
Type: Phacographa glaucomaria (Nyl.) Hafellner (Lecidea glaucomaria Nyl.)
(holotype)
Etymology: Phaco- from phakos [gr.] – lentil, shaped like the seed of a lentil, -
grapha from grapho [gr.] – write, in order to indicate its relationship with both
Phacothecium and Opegrapha
Description: Ascomata lecideoid, roundish, carbonized, centrum persistently
unilocular, i.e. one hymenium per ascoma, exposed from an early stage of
development, obtecting portions of plectenchyma crumbling off entirely, rarely
some remnants remaining for longer time on the disc, exposed parts of the disc
always with a dark brown tinge. Exciple laterally and basally well developed, dark
brown, somewhat more blackish with K. Hymenium penetrated by hemiamyloid
gel (I+ blue then orange, K/I+ blue). Asci arthonialean, broadly cylindrical to
subclavate, rather abruptly pedicellate, in the endoascal wall layer with
hemiamyloid apical ring. Ascospores with hyaline proper wall, with at least 3
septa, with rounded ends, surrounded by thin perisporal sheath, giving rise to a
verruculose sculpture with age. All know species lichenicolous.
Discussion: The genus Phacographa is introduced for a few lichenicolous
ascomycetes commonly treated as Opegrapha species.
Ascomata of Phacographa species are unilocular. Although this is sometimes
not clearly visible under the dissecting microscope in surface view, this is evident
in longitudinal sections; also old cup-shaped ascomata with eroded hymenia (such
are sometimes present on specimens beside healthy ones) show clearly that there
was only one hymenium developed in individual ascomata.
The ascomatal pigment is the same as in many other arthonialean fungi, called
“Pigment 1” by ERTZ et al. (2005), with “Atra-brown” (MEYER & PRINTZEN
(2000) as synonym, for which Opegrapha atra is the reference species.
Judging from the shape of ascomata, Phacographa is more similar to
Lecanactis (with apothecioid ascomata) than to Opegrapha (ascomata
hysterothecioid although sometimes short, as in O. parasitica (type of
Leciographa) and O. centrifuga (type of Lecoglyphis)); also Lecanographa was
compared but found to be sufficiently different by subhysterothecioid ascomata
with discs obtected by a white pruina. Lichenicolous growth is unknown in
Lecanactis but well documented in Opegrapha and Lecanographa. Phacothecium
differs by hemiangiocarpous ascomata, elongated ascomatal discs which are
developed by a retarded rupture of the obtecting plectenchyma, and asci lacking
hemiamyloid ring structures.

102
The species
Phacographa glaucomaria (Nyl.) Hafellner comb. nov.
MB 513174
Bas.: Lecidea glaucomaria Nyl., Bot. Notiser 1852: 177 (1852). – Opegrapha
glaucomaria (Nyl.) Källsten ex Hafellner, Bull. Soc. Linn. Provence 45: 227 (1994). –
Leciographa glaucomaria (Nyl.) A.L.Sm., Monogr. Brit. Lich., ed. 1, 2: 186 (1911).
Type: [Sweden]: Holmiae, on Lecanora rupicola, 1852, leg. W. Nylander (H-Nyl. 11019,
lectotype) Lectotype selected here! – Holmiae, on Lecanora rupicola, 1852, leg. W.
Nylander (H-Nyl. 11026, paratype!). – Prope Holmiam, on Lecanora rupicola, 1852, leg.
W. Nylander (H-Nyl. 11022 – paratype!).
Syn.: Dactylospora maculans Arnold, Verh. Zool.-Bot. Ges. Wien 37: 126, 150 (1887). –
Opegrapha maculans (Arnold) Hafellner ex Poelt & Türk, Herzogia 6(3-4): 453 (1984). –
Leciographa maculans (Arnold) Rehm, Rabenh. Krypt.-Fl., 2. Aufl. 1(3): 375 (1890).
Type: [Italy]:„…östlich über dem Rollepaß…“ (locality information from protologue) (n.
v.); Specimens from type locality however evidently not representing type material
distributed in Arnold, Lich. exs. 1250: [Italy, Trento], „an Felsklippen der Campiler
Schichten auf der Höhe ober dem Rollepass, Paneveggio in Südtirol“, auf Lecanora
rupicola coll., 21. VII. 1887, leg. F. Arnold (M, slide in GZU).
Icon.: NYLANDER 1852: tab. 1, fig. 10 (drawing of ascospores); this publication, Fig. 3
(longitudinal section of ascoma), Fig. 4 (ascus stained in K/I).
Exs.: Arnold, Lich. exs. 1250 sub Dactylospora maculans (M); Santesson, Fungi
Lichenicoli exs. 63 sub Opegrapha maculans (GZU); Hafellner, Lichenicolous Biota no.
adhuc ined.; Flagey, Lich. Algerienses 199 sub Dactylospora maculans (n. v.)
Description: Ascomata in dispersed groups on the thallus of the host, individual
ascomata in dense groups of mostly 5-25. Ascomata apothecioid, 0.4-0.6(-0.7)
mm diam., semi-immersed, with widely exposed, rough, dark brown disc and a
rough, often delicately fissured, black margin. Exciple in longitudinal section 40-
60 μm wide, dark brown. Hypothecium dark brown, 70-100 μm high, composed
of hyphae with distinctly elongated cells. Subhymenium 20-40 μm high,
composed of short-celled hyaline hyphae intermingled with some pale brown
ones. Hymenium hyaline, including the epihymenium 70-80 μm high, penetrated
by a hemiamyloid hymenial gel. Paraphysoids branched and anastomosing, 2-3
μm in diam., apically enlarged to pigmented tips measuring 4-6 μm in diam.
Epihymenial pigment brown, granular, deposited on the outside of the paraphysal
tips, K–. Asci broadly cylindrical to subclavate, shortly pedicellate, 8(-6-4)
spored, 70-80 × 22-28 μm, wall I–, K/I– except for an apical ring that is K/I+ blue
and fading in the adjacent parts of the endoascal wall (Fig. 4). Ascospores hyaline,
3-septate, 20-24(-26) × 6-7 μm, length-width-ratio ≥ 3, with thin perispore sheath,
giving rise to a pale to brown verruculose sculpture with age.
Hosts: Lecanora rupicola coll. (syn. L. sordida, L. glaucoma) (thallus and
apothecia) (1), L. rupicola ssp. sulphurata (thallus and apothecia) (2), L. bicincta
(thallus and apothecia) (3), L. spec. (4). In the literature cited below also one of
the synonyms may have been used.
Ecology: The species as circumscribed here is restricted to the Lecanora rupicola-
group. As for specimens of lichenicolous fungi, the hosts are often not very

103
critically determined; infraspecific taxa as defined by LEUCKERT & POELT (1989)
are mostly not mentioned on the labels. Even the upland species L. bicincta is
often not distinguished, which may be a more common host in higher altitudes as
seen from the data published so far. Records of the species upon Protoparmelia
badia have been listed under Phacographa protoparmeliae (see below).
Distribution: The species is widely distributed in the Northern Hemisphere, with
records from Europe, Asia, Africa and North America, from the arid subtropical-
Mediterranean zone northwards to the southern arctic zone. The localities known
so far are situated from lower montane up the alpine vegetation belt with an
altitudinal range of c. 680-2800 m. It is already known from several countries
under various synonyms (Opegrapha glaucomaria or homotypic synonyms (A),
Opegrapha maculans or homotypic synonyms (B)). So far it has been recorded
from the European countries Austria (SIMMER 1899, as (B) on (1), KEISSLER
1930, as (B) on (1), POELT & TÜRK 1984, as (A) on (3), HAFELLNER & SANCHO
1990, as (B) on (1), HAFELLNER & TÜRK 1995, as (A) on (1), BOOM et al. 1996,
as (A) on (3), HAFELLNER et al. 2004, as (A) on (3), France (OLIVIER 1903, 1906,
both as (A) on (1), COSTE 1993, as (A) on (1), ROUX et al. 2006, as (A) on (1)),
Greece (this publication), Ireland (KNOWLES 1929, as (A) on (1)), Italy
(BAGLIETTO 1871, as Leciographa parasitica on (1), ARNOLD 1887, 1889, both as
(B) on (1), KERNSTOCK 1895, as (B) on (1), ARNOLD 1896, as (B) without host,
KEISSLER 1930, as (B) on (1)), Macedonia (this publication), Portugal (SAMPAIO
1917, as (A) on (1)), Spain (HAFELLNER & SANCHO 1990, as (B) on (1)), SWEDEN
(Nylander 1852, as (A) on (1), SANTESSON 1986, as (B) on (1), SANTESSON 1993,
as (A) on (1), THOR 1993, as (A) on (1)), the United Kingdom (LEIGHTON 1879,
as (A) on (1), HITCH 1997, as (A) on (1)), and the island Corsica (HAFELLNER
1994a, as (A) on (1)). In Asia it has already been reported from Russia
(NYLANDER 1887, as (A) on undetermined lichen) and Turkey (HAFELLNER &
JOHN 2006, as (A) on (2)). In Africa it is so far known only from the archipelagos
Canary Islands (HAFELLNER 1995, as (A) on (1), HERNÁNDEZ PADRÓN et al. 2004,
as (A) on (1)) and Madeira (KALB & HAFELLNER 1992, as (B) on (1)). In North
America it has been reported from the United States (FINK 1935, as (A) on (4) pro
parte, HAFELLNER et al. 2002, as (A) on (1)) and Greenland (NYLANDER 1862, as
(A) without host, STIZENBERGER 1876, as (A) on (1)).
Discussion: Phacographa glaucomaria is a typical member in what the author
believes is a monophyletic species group, and it was therefore chosen as type
species. The closest relative is the newly described P. protoparmeliae, confined to
strains of Protoparmelia badia coll., which can be morphologically distinguished
by its more sessile mature ascomata and smaller (shorter) ascospores.
The species has nothing to do with Arthonia glaucomaria Nyl. 1856, a
heterotypic synonym of Arthonia varians (Dav.) Nyl., which also grows
preferably on Lecanora rupicola, but there in the hymenia. Several times Lecidea
glaucomaria Nyl. 1852 has been cited as the basionym of the Arthonia (e.g.
GALLOWAY 2007: 48), with the effect that the author citation for Arthonia
glaucomaria would be “(Nyl.) Nyl.”, which is definitely wrong. As far as the
linguistic root of the epithet is concerned, it is evident from the protologue
(NYLANDER 1852) that the author interpreted it as an adjective to be declined (and
not as or derived from a noun). For the host the name, Parmelia glaucoma was

104
Figs 3-4. Phacographa glaucomaria, medium: water. Fig. 3. Ascoma in longitudinal
section, unstained (from J. Poelt, 24.7.1994). Fig. 4. Young ascus, staining: K/I, (from J.
Poelt, 26.6.1983).
used and in the caption of the table the genitive is used (“Sporae Lecideae
glaucomariae”).
Further specimens seen: Europe: Austria: Salzburg: Lungau, [Niedere Tauern],
Schladminger Tauern, Weißpriachtal [N von Mauterndorf], Felsblöcke nördlich der
Lahnbrücke, 1290 m; GF 8747, (1), 8.9.1981, J. Poelt (GZU). – Kärnten, Nationalpark
Hohe Tauern, Ankogel-Gruppe, W Hänge des Auernig E ober Mallnitz, am Steig zum
Köpfl, 1200-1300 m, GF 9045/1, an teilweise Ca-hältigen Schrofen im lichten Lärchen-
Fichtenwald, (1), 25.10.1988, J. Hafellner 28619 & M. Walther (GZU). – Kärnten,
Nationalpark Hohe Tauern, Ankogel-Gruppe, Dösental E von Mallnitz, Geländerippe N
ober dem Arthur von Schmidt Haus, 2275-2400 m, GF 9045/2, Gneisblöcke, (1),
30.7.1989, J. Hafellner 22941 & R. Türk (GZU). – Kärnten, Hohe Tauern, Kreuzeck-
Gruppe, Schwarzsteinwände (Südwände des Schwarzsteins) E der Hochtristen, N der
Turgger Alm, [46°47’55’’N / 13°09’10’’E], 1950-2150 m, GF 9244/2, (3), 17.7.1978, J.
Hafellner 3889 (GZU). – Kärnten, Hohe Tauern, Kreuzeck-Gruppe, Hänge oberhalb der
Turgger Alm gegen die Schwarzstein Wände, [46°47’55’’N / 13°09’10’’E], 2000-2100

105
m, GF 9244/2, Grünschiefer, Überhangfläche, (3), 24.7.1994, J. Poelt (GZU). – Kärnten:
Hohe Tauern, Kreuzeck-Gruppe, Kalkschieferwände in den SE-Hängen der Sensenspitze
N der Turgger Alm, [46°47’40’’N / 13°08’50’’E], 1900-2100 m, GF 9244/2, (1),
17.7.1978, J. Poelt (GZU). – Steiermark, Niedere Tauern, Schladminger Tauern,
Krahbergzinken SE von Schladming, SW-Hänge knapp unter dem Gipfel, 47°21’20’’N /
13°44’30’’E, c. 2050 m, GF 8648/1; niedere Schrofen aus Glimmerschiefer (stellenweise
mit leichtem Kalkeinschlag) in steilen Rasen, auf Schrofen an Steilflächen, (3),
13.8.2001, J. Hafellner 56404 (GZU), distributed in Hafellner, Lichenicolous Biota no.
adhuc ined. – Steiermark, Niedere Tauern, Wölzer Tauern, W-Abhänge der
Gstemmerspitzen, [47°24’50’’N / 14°11’50’’E], c. 1900 m, GF 8551; Schrofen aus
Hornblendeschiefer, (1), 22.7.1985, J. Hafellner (GZU). – Ibid., (1), 20.7.1987, J.
Hafellner 16165 & L. Brako (GZU). – Steiermark, Gurktaler Alpen, Frauenalpe
(„Frauenalm“) S von Murau, S-exponierte Abbrüche zwischen Oberberg und dem Gipfel
der Frauenalpe, [47°04’10’’N / 14°08’40’’E], c. 1750 m, GF 8950/2; kalkhaltige
Schiefer, auf Steilflächen, (1), 26.6.1983, J. Poelt (GZU). – Ibid., (3), J. Hafellner 10359
(GZU).– France: [Bretagne, Dept. Finistère], Brest, 1868, Crouan (H-Nyl. 11020). –
Auvergne, Dept. Cantal, Orgues SE von Chadecol, E ober Blesle, c. 680 m; SW-
exponierte Basaltabbrüche, (1), 1.8.1980, A. Bellemère & J. Hafellner 9576 (GZU). –
Auvergne, Dept. Cantal, Monts du Cantal, Puy Mary, am SE-Grat gegen den Puy de
Peyre-Arse, c. 1600 m; Andesitblöcke, (1), 30.7.1980, A. Bellemère & J. Hafellner 9482
(GZU). – Korsika, Dept. Haute-Corse, Asco Tal, Forêt de Carrozzica, SW ober dem Ort
Asco, c. 940 m; lockerer Pinus nigra-Wald mit Felsblöcken, auf Rhyolith-Schrofen, (1),
2.11.1993, M. Krenn, H. Mayrhofer & E. Unger (GZU). – Dept. Haute-Corse, Monte
d'Oro - Massiv, Kuppe N über dem Col de Vizzavona, c. 1450 m; Granitblockwerk, auf
besonnten Neigungsflächen von Granitblöcken, (1), 3.11.1993, J. Hafellner 31590
(GZU). – Ibid., (1), B. Wieser (GZU). – Greece: Attika, kurz W unterhalb des
Gipfelgrates des Hymettos (Imittos) bei Athen, 800-900 m, N-seitige Abbrüche saurer
Silikate, (2), 14.4.1971, J. Poelt (GZU). – Italy: Trentino: Val di Moena S von Cavalese,
[46°15’40’’N / 11°27’15’’E], 1100-1200 m, an Silikatblöcken auf einer Weide, (1),
27.10.1984, J. Poelt (GZU). – Trento, „an Felsklippen der Campiler Schichten auf der
Höhe ober dem Rollepass, Paneveggio in Südtirol“, (1), 21.7.1887, F. Arnold (M, slide in
GZU) = Arnold, Lich. exs. 1250. – Emilia Romana, Prov. Reggio Emilia: Apennino,
Monte Ventasso S above the village Ramiseto, slopes exposed to N, SE above Lago
Calamone, 44°22’55’’N / 10°17’35’’E, c. 1450 m; scree of sandstone boulders, on
inclined rock faces of sandstone boulders, (1), 23.7.2001, J. Hafellner 58277 (GZU, herb.
Hafellner). – Calabria, Prov. Cosenza, Craticello, c. 1200 m, Felsgruppen in Macchie, (1),
11.7.1988, J. Poelt (GZU). – Mazedonia: Baba planina, N-Abhänge des Pelister,
Silikatblockwerk, (1), 13.7.1977, J. Poelt (GZU). – Spain: Prov. Madrid, Sierra de
Guadarrama: Aufstieg vom Puerto de los Cotos auf die Peñalara, SE-Flanke ober der
Laguna Grande, 2100-2200 m, Granitgneis, (1), 9.9.1980, J. Hafellner 10549 (GZU). –
Prov. Tarragona: Sierra de Prades, an der Straße vom Monasterio de Poblet nach Prades
kurz ober der Abzweigung nach Vilanova de Prades, c. 900 m, Quercetum pyrenaicae mit
Silikatblöcken, (1), 23.5.1983, J. Poelt (GZU). – Ibid., (1), J. Hafellner 17450, 17461
(GZU). – Sweden: Öland,, Böda par., Sjötorp, on grantic boulders of a stone wall by a
small road, (1), 13.8.1963, R. Santesson 16248 (GZU) = Santesson, Fungi Lichenicoli
exs. 63. – Asia: Turkey: Trabzon (61), Doğu Karadeniz Dağları, SE of Uzungöl, Balıklı
Göl, 40°32’N / 40°23’E, 2550 m, on siliceous rocks of granite, (2), 25.7.1997, V. John
10361 (GZU). – Africa: Madeira: c. 3.5 km W von Poiso ober der Straße auf den Pico
Areeiro, N ober der Achada Grande, Felsköpfe am Grat, c. 1600 m, 32°43'N/16°55'W;
Vulkanit, S-seitige Überhangsflächen, (1), 15.2.1990, J. Hafellner 27538 & A. Hafellner
(GZU). – Pico Areeiro, markante Felsrippe NE des Gipfels, c. 1800 m, 32°44'N/
16°55'20''W; NE-exponierte Abbrüche, auf Vulkanit, (1), 22.2.1990, J. Hafellner 27421
& A. Hafellner (herb. Hafellner). – Canary Islands: Gran Canaria: am Sattel SW unter

106
dem Pico de las Nieves, c. 1820 m, 27°57'30"N/15°34'30"W; Vulkanitkonglomeratblöcke
im lockeren Kiefernwald, auf N-seitigen Steilflächen großer Blöcke, (1), 2.3.1994, J.
Hafellner 32927 (GZU). – am Felsrücken kurz SE vom Roque Nublo c. 3 km oberhalb
von Ayacata, c. 1720 m, 27°58'N/15°37'W; Vulkanitkonglomeratschrofen, auf E-
exponierten Steilflächen, (1), 24.2.1994, J. Hafellner 32896 (herb. Hafellner). – North
America: U.S.A.: Colorado, Clear Creek Co., 2.4 km from Idaho Springs, up the Soda
Creek Road, on gneissic rock, (1), 7.9.1971, R. A. Anderson 8196 (GZU). – Arizona,
Apache Co., Apache National Forest, Mt. Baldy Wilderness, valley of the East Fork of
the Little Colorado River, between Phelp's cabin and the base of cliffs, c. 2800 m,
33°55'30"N/109°30'30"W; mixed conifer forest including large trees of Pseudotsuga
menziesii, on boulders of volcanic conglomerate at the edge of the forest, (1), 3.7.1994, J.
Hafellner 36639 (GZU).
Phacographa protoparmeliae Hafellner spec. nov.
MB 513175
Species differt a Phacographa glaucomaria ascomatibus plus elevatis, ascosporis
brevioribus. Habitat supra thallum Protoparmeliae badiae. Adhuc solum in Europa
inventa.
Type: Austria, Kärnten, Hohe Tauern, Kreuzeck-Gruppe, Kalkschieferwände in den SE-
Hängen der Sensenspitze N der Turgger Alm, [46°47’40’’N / 13°08’50’’E], c. 2000 m,
auf Amphibolit, (1), 17.7.1978, J. Hafellner 603 (GZU – holotypus).
Icon.: this publication, Fig. 5 (longitudinal section of ascoma), Fig. 6 (mature and
overmature ascospores).
Exs.: -
Description: Ascomata in dispersed groups on the thallus of the host, individual
ascomata in dense groups of mostly 3-10 or dispersed. Ascomata apothecioid,
semi-immersed only at the beginning, later ± sessile, with widely exposed, rough,
dark brown disc and a rough, often delicately fissured, black margin. Exciple in
longitudinal section 40-60 μm in diam., dark brown, basally fusing with a dark
brown hypothecium, 70-100 μm high, composed of hyphae with distinctly
elongated cells. Subhymenium 20-40 μm high, composed of short-celled hyaline
hyphae intermingled with some pale brown ones. Hymenium hyaline, including
the epihymenium 70-80 μm high, penetrated by a hemiamyloid hymenial gel.
Paraphysoids branched and anastomosing, 2-3 μm in diam., apically enlarged to
pigmented tips measuring 4-6 μm in diam. Epihymenial pigment middle brown,
granular, deposited on the outside of the paraphysal tips, K–. Asci broadly
cylindrical to subclavate, shortly pedicellate, 8(-6-4) spored, 70-80 × 22-28 μm,
wall I–, K/I– except for an apical ring that is K/I+ blue and fading in the adjacent
parts of the endoascal wall (Fig. 6). Ascospores hyaline, 3-septate, 18-21(-23) × 6-
7 μm, length : width ratio ≤ 3, with distinct perispore sheath, giving rise to a pale
to middle brown verruculose sculpture with age.
Hosts: Protoparmelia badia coll. (th.) (1)
Ecology: Although Protoparmelia badia is one of the commonest lichens on
siliceous rocks from the upper montane to the lower alpine belt in European
mountains, infections with Phacographa protoparmeliae are relatively rare.

107
Figs 5-6. Phacographa protoparmeliae (holotype), unstained, medium: water. Fig. 5.
Ascoma in longitudinal section. Fig. 6. Mature and overmature ascospores.
According to personal field observations, thalli growing on steep to subvertical
rock faces are more likely to become parasitized.
Distribution: The species is so far known only from Europe with localities in the
Alps and the mountain chains in Central Spain. It has earlier been recoded under
Opegrapha glaucomaria (A) or O. maculans (B) from Austria (HAFELLNER &
SANCHO 1990, as (B) on (1), BOOM et al. 1996, as (A) on (1), HAFELLNER 2002,
as (A) on (1), HAFELLNER et al. 2004, as (A) on (1), HAFELLNER & OBERMAYER
2007, as (A) on (1)), France (BOOM & BREUSS 2002, as (B) on (1)), Italy
(TRETIACH & HAFELLNER 2000, as (A) on (1)), and Spain (HAFELLNER &
SANCHO 1990, as (B) on (1)).
Discussion: The species is closely related to Phacographa glaucomaria, with
which it was confused in the past. Findings of Phacographa protoparmeliae have
therefore previously been published under the names Opegrapha maculans or O.
glaucomaria. Phacographa protoparmeliae is morphologically distinguished
from P. glaucomaria by its more sessile ascomata and shorter ascospores resulting
also in a different length : width ratio.
Further specimens seen (representing paratypes): Austria: Kärnten, Nationalpark Hohe
Tauern, Ankogel-Gruppe, Hänge E vom Hannover Haus gegen die Grauleitenspitze,
[47°02’45’’N / 13°13’30’’E], 2700-2850 m, GF 8945/3, auf Gneisblöcken, (1),
29.7.1989, R. Türk & J. Hafellner 32081 (GZU). – Kärnten, Hohe Tauern, Kreuzeck-
Gruppe, Hänge am Weg von der Emberger Alm zur Turgger Alm, 1800-2000 m, (1),
17.7.1978, J. Poelt (GZU). – Kärnten, Hohe Tauern, Reißeck-Gruppe, Gneishänge W
über dem Kleinen Mühldorfer See, S des Reißecks, ca. 2400 m, (1), 3.9.1974, J. Poelt
(GZU). – Kärnten, Steirisches Randgebirge, Koralpe, auf der Hanalpe über der
Weinebene, 1750-1850 m, isolierte Gneisfelsen (sog. Öfen), (1), 29.10.1985, J. Poelt &
G. Miehe (GZU). – Kärnten, Steirisches Randgebirge, Koralpe, c. 11.5 km NE über St.
Paul im Lavanttal, Krakaberg SE vom Koralpenhaus, S-seitig im Gipfelbereich,

108
46°46’50’’N / 14°58’15’’E, c. 2070 m, GF 9255/2; Blöcke und Ausbisse eines harten,
quarzreichen Schiefers, auf Neigungsflächen kleiner Felsausbisse, (1), 2.7.2007, J.
Hafellner 68494, L. Muggia & A. Hafellner (GZU). – Kärnten, Steirisches Randgebirge,
Koralpe E von Wolfsberg, W-Hänge des Bergrückens zwischen Hühnerstütze und
Seespitz, E ober dem Großen Kar, 46°47’45’’N / 14°58’40’’E, c. 1990 m, GF 9255/2;
bankige Glimmerschieferschrofen in alpinen Rasen, bodennah auf kleinen
Glimmerschieferblöcken, (1), 24.10.2004, J. Hafellner 70845 & A. Hafellner (herb.
Hafellner). – Steiermark, Niedere Tauern, Schladminger Tauern, E-Abhänge des
Bergrückens zwischen Gasselhöhe und Rippetegg SW von Schladming, am Steig von der
Gasselhöhehütte zum Mittersee, c. 1800 m, 47°21’10’’N / 13°35’50’’E, GF 8647/2;
Glimmerschieferblockwerk in Zwergstrauchheiden an der Waldgrenze, auf Steilflächen
von großen Schieferblöcken, (1), 24.8.2001, J. Hafellner 56892 & W. Obermayer (GZU,
herb. Hafellner). – Steiermark: Niedere Tauern, Wölzer Tauern, Rettlkirchspitze NW von
Oberwölz, im oberen Teil des NE-Grates SW ober der Neunkirchner Hütte, 47°15’45’’N
/ 14°08’05’’E, c. 2300 m, GF 8750/2; Gneisblockwerk und Fragmente alpiner
Vegetation, auf Steilflächen von Blöcken, (1), 24.8.2002, J. Hafellner 68167& J.
Miadlikowska (herb. Hafellner). – Steiermark: Niedere Tauern, Triebener Tauern,
Griesmoar Kogel SW von Wald am Schoberpaß, am N-Rücken halbwegs zwischen dem
Sattel zum Himmeleck und dem Gipfel, 47°25’15’’N / 14°36’10’’E, c. 1950 m, GF
8553/4; niedere Ausbisse aus Glimmerschiefer, stellenweise mit Ca-Spuren, in alpinen
Matten, auf Steilflächen, (1), 20.8.2002, J. Hafellner 58905 & J. Miadlikowska (GZU). –
Steiermark: Steirisches Randgebirge, Stubalpe, Größenberg S von Zeltweg, NW-Rücken
gegen Eppenstein, etwa halbwegs zwischen Wetterkopf und dem Gipfel, 47°05’15’’N /
14°47’35’’E, c. 2020 m, GF 8954/2; niedere Gneisausbisse in Zwergstrauchheiden, auf
Neigungsflächen kleiner Felsausbisse, (1), 15.8.2006, J. Hafellner 66588 (GZU). –
Steiermark: Steirisches Randgebirge, Stubalpe, Ameringkogel-Massiv E von Obdach, im
NE-exponierten Kar zwischen Ameringkogel und Größenberg, 47°04’30’’N /
14°48’15’’E, c. 2050-2100 m, GF 8954/2, Gneisschrofen, auf Gneisschrofen an
Neigungs- und Steilflächen, (1), 28.7.1990, J. Hafellner 28867 & W. Obermayer (GZU).
– Spain: Prov. Madrid, Sierra de Guadarrama: SE vom Puerto de los Cotos, 2150 m,
Felsköpfe an einem Geländegrat aus Gneis, (1), 8.9.1980, J. Hafellner 10924 (herb.
Hafellner).
Phacographa zwackhii (A.Massal. ex Zwackh) Hafellner comb. nov.
MB 513176
Bas.: Leciographa zwackhii A.Massal. ex Zwackh, Flora 45: 571 (1862) (ICBN 46.4.). –
Massalongo, Verh. Zool-Bot. Ges. Wien 10: 679 (1860) (nomen nudum, as “
Leucographa Zwackii”). – Syn.: Opegrapha zwackhii (A.Massal. ex Zwackh) Källsten ex
Hafellner, Herzogia 10: 15 (1994).
Type: [Germany]: “Parasitisch auf dem Thallus von Phlyctis argena an Buchen auf dem
Köhnigstuhle und Auerhahnkopfe“ leg. ?Zwackh (locality information from protologue,
type not seen).
Note: Although Zwackh ascribed the name of the new species to Massalongo,
there is no indication, that the validating description is also provided by
Massalongo. According to the code, the name of a new taxon must be attributed to
the author of the publication.
Icon.: Fig. 7 (longitudinal section of ascoma), Fig. 8 (ascus), Fig. 9 (ascus tip stained in
K/I).

109
Exs.: Arnold, Lich. exs. 253 (as Leciographa zwackhii) (GZU); Zwackh, Lich. exs. 353
(as Phlyctis argena with parasite) (n. v.).
Description: Ascomata scattered on the thallus of the host, embedded between
soredia of the host or breaking through the “cortical” plectenchyma of the non-
sorediate thallus margin, relatively small, 0.2-0.4(-0.5) mm in diam., with widely
exposed, rough, dark brown disc and a rough black margin. Exciple in
longitudinal section 20-30 μm in diam., dark brown. Hypothecium dark brown,
20-40 μm high, composed of hyphae with more or less isodiametric cells.
Subhymenium 20-30 μm high, composed of short-celled hyaline hyphae
intermingled with some pale brown ones. Hymenium hyaline, including the
epihymenium 70-80 μm high, penetrated by a hemiamyloid hymenial gel.
Paraphysoids branched and anastomosing, 2-3 μm in diam., apically enlarged to
pigmented tips measuring 4-6 μm in diam. Epihymenial pigment middle brown,
granular, deposited on the outside of the paraphysal tips, K-. Asci broadly
cylindrical to subclavate, shortly pedicellate, 8-(6-)spored, 60-70 × 18-25 μm,
wall I–, K/I– except for an apical ring that is K/I+ blue and fading in the adjacent
parts of the endoascal wall (Fig. 9). Ascospores hyaline, mostly 3- to 4-septate,
seldom some spores to 5-septate, slightly clavate, 18-22(-26) × 5-6(-7) μm, with
thin perispore sheath, giving rise to a pale to brown verruculose sculpture with
age.
Hosts: Phlyctis argena (th.) (1), Phlyctis agelaea (th.) (2)
Ecology: Although in Central Europe Phlyctis argena is one of the commonest
crustose lichens on bark, Phacographa zwackhii is a rarely collected lichenicolous
fungus. At least in Austria the localities are sites of old growth forests, thus P.
zwackhii might be a good indicator species of long ecological continuity in
montane mixed deciduous forests.
Distribution: The species is so far known only from Europe. It has been reported
under Opegrapha zwackii and its homotypic synonyms (all on (1) if not otherwise
stated) from Austria (HAFELLNER 1994b, 2002, TÜRK & BREUSS 1994), Estonia
(SUIJA 2005), France (RONDON 1969, on (2), ROUX 1982), Germany (ZWACKH
1862, BAUSCH 1869, LAHM 1884, SANDSTEDE 1889, REHM 1890, ERICHSEN 1906,
KEISSLER 1930, TRIEBEL & SCHOLZ 2001), Poland (STEIN 1879, EITNER 1901),
Sweden (ERICHSEN 1914, SANTESSON 1993), and the British Isles (GILBERT &
COPPINS 1992).
A record on an unusual host from Germany (LETTAU 1958, as Leciographa
zwackhii on Pertusaria amara) possibly refers to Opegrapha anomea.
Discussion: Phacographa zwackhii has distinctly smaller ascomata than the other
Phacographa species. Further diagnostic features are the relatively slim
ascospores and the frequent formation of a further septum resulting in partly 4-
septate ascospores (acc. to REHM 1890 rarely even 5-septate).
Pseudotryblidium neesii (Flot.) Rehm has sometimes been misidentified as
Opegrapha zwackhii (e.g. LETTAU 1912a) or both species have not been clearly
distinguished (compare also REHM 1890: 375). Pseudotryblidium neesii, a non-
lichenized, non-lichenicolous helotialean ascomycete is a specific inhabitant of
bark of Abies species. The ascomata often break through the spreading thalli of
corticolous crustose lichens, including Loxospora elatina and Phlyctis argena.

110
Further specimens seen (all on Phlyctis argena): Europe, Austria: Steiermark:
Nördliche Kalkalpen, Ennstaler Alpen, Gesäuse, Hartelsgraben c. 3 km W von Hieflau,
kurz vor der 2. Brücke, 47°35’15’’N / 14°42’25’’E, c. 660 m, GF 8454/1; Mischwald, auf
Borke von Fagus sylvatica, 20.5.1988, J. Hafellner 20249 & E. Schreiner (GZU). – Ibid.,
auf Borke von Acer pseudoplatanus, 20.5.1988, J. Hafellner 20201 & E. Schreiner
(GZU). – Steiermark, Niedere Tauern, Triebener Tauern S von Trieben, Großer
Grießstein, sanft geneigte Hänge am Nordfuß S ober dem Bildstock beim Gehöft Eberl,
47°24’35’’N / 14°33’05’’E, c. 1160 m, GF 8553/3; Fichtenforst mit einzelnen Bergahorn,
auf Borke von Acer pseudoplatanus, 15.8.2000, J. Hafellner 52208 (GZU). –
Niederösterreich, Österreichische Kalkalpen, Großer Urwald an den SE-Abhängen des
Dürrensteins, c. 1000-1150 m, Buchen-Tannen-Fichtenurwald, 19.6.1981, C.
Scheideggerno. 330 (GZU). – Niederösterreich, Rothwald c. 16 km W von Mariazell,
Kleiner Urwald, 980-1020 m, GF 8256, 29.7.1990, R. Türk & O. Breuß 7327 (herb.
Hafellner). – Niederösterreich: Nördliche Kalkalpen, Göller-Gruppe, Südhänge der
Weißmäuer E vom Lahnsattel, c. 1000 m, 47°46’30’’N / 15°31’30’’E, GF 8259/1;
Buchen-Tannen-Fichtenwald, an Stammborke von Acer pseudoplatanus, 22.5.1998, J.
Hafellner 50784 (herb. Hafellner). – Germany: Württemberg, Ellwangen, Wald bei
Rosenberg, an Abies alba, July 1862, Kemmler (GZU). – British Isles: Scotland, East
Perthshire, 6 km N of Blairgowrie, E side of River Ericht, Milton Wood, c. 125-165 m,
on Fraxinus excelsior, 1.10.1981, B. J. Coppins 8717 & R. K. Brinklow (GZU).
Figs 7-9. Phacographa zwackhii (Hafellner 20201), medium: water. Fig. 7. Ascoma in
longitudinal section, unstained. Fig. 8. Ascus, unstained, DIC. Fig. 9. Ascus, staining:
K/I.

111
Excluded species
“Opegrapha” rotunda Hafellner, Herzogia 10: 14 (1994).
Type: Austria: Steiermark, Nördliche Kalkalpen, Greith im Salzatal, E des Ortes an der
Straße von Gußwerk nach Weichselboden, 720 m, MTB 8257/3; am Rand einer
Mähwiese an freistehenden Fraxinus excelsior, auf Physconia distorta; 2. Juni 1987, J.
Hafellner 16001 (GZU – holotype!).
Icon.: Hafellner 1994b: 16 (drawing of paraphysoids, ascus and ascospores)
Exs.: –
Description: Ascomata lichenicolous, on the thallus of Physconia species,
frequently developed at the margin of the lobes, black, roundish, fully developed
c. 0.5-1 mm in diam., in external view similar to apothecia of a Dactylospora, but
with rough surface, with thin and somewhat prominent margins, with flat discs.
Exciple in longitudinal section dark brown, in the margin the hyphal cells radially
arranged. Hypothecium brown. Hymenium hyaline, penetrated by hemiamyloid
gel, I+ blue then orange, K/I+ blue, including the epihymenium c. 100 μm high.
Paraphysoids ramified and anastomosing, relatively short-celled, terminal cells
enlarged and with brown hyphal walls and embedded in brownish gel. Asci
arthonialean, (4-) to 8-spored, broadly cylindrical to somewhat clavate, 60-80 ×
15-20 μm large, dehiscence fissitunicate, no reaction with K/I. Ascospores
hyaline, 3-septate, somewhat clavate with the second cell being the broadest, 18-
22 × 5-6 μm large, with thin perispore sheath.
Hosts: Physconia distorta (thallus) (1)
Ecology: The species has rarely been found; therefore, its ecological needs are
still not well understood. Oceanic site conditions seem to be favourable.
Distribution: The species (all records as Opegrapha rotunda on Physconia
distorta) is so far known only from a few localities in Austria (HAFELLNER 1994b,
HAFELLNER 2003), Portugal (BOOM & ETAYO 2000) and the British Isles (HITCH
1995).
Discussion: As Opegrapha rotunda is characterized by its lecideoid ascomata, it
represents a further candidate for a member of the newly described genus.
However, we do not think that it belongs to Phacographa. Neither does it show
the typical dark red-brown colour of the apothecial discs nor do the asci react in a
comparable manner with Lugol’s as Phacographa species. Thus its position in
Phacographa would be relatively isolated and we do not see any progress in the
taxonomy of opegraphoid lichenicolous fungi with such a transfer.
Further specimens seen (all on Physconia distorta (th.)): Europe, Austria: Kärnten,
[Gailtaler Alpen], Goritschach W von Villach, [46°36’40’’N / 13°48’30’’E], Allee, an
Fraxinus excelsior, 9.2.1974, J. Poelt (GZU). – Kärnten, [Karawanken], Rosental, St.
Margarethen i. R., Niederdöfl, an der Bundesstraße c. 100 m vor der Abzweigung nach
Gupf, Berghof, neben der Feldkapelle in der Kurve, [46°32’20’’N / 14°27’05’’E], c. 700
m, GF 9452/4, an Aesculus hippocastanum, 17.9.1986, B. Auer 1471 (GZU). –
Steiermark: [Ennstaler Alpen], Gesäuse-Gebiet, Johnsbach, c. 0,5 km E des Gasthofs
Kölbl, [47°31’50’’N / 14°37’10’’E], c. 875 m, MTB 8453/4; an Fraxinus excelsior,
20.5.1988, J. Hafellner 20304 & E. Schreiner (herb. Hafellner). – Steiermark: Steirisches
Randgebirge, Fischbacher Alpen, „Oststeirisches Hügelland’’, bei Neudörfl S von

112
Birkfeld, am Ufer der Feistritz, [47°20’30’’N / 15°42’00’’E], c. 540 m, GF 8660/3,
Ufergehölzstreifen, an Fraxinus excelsior, 22.10.1989, J. Hafellner 22167 & A. Hafellner
(GZU).
Discussion
In advanced stages of development the crowded arrangement of ascomatal discs
in Phacothecium may recall to mind the composed ascomata of Plectocarpon and
Perigrapha. In all three genera early stages of ascoma development start with a
plectenchymatal body, in which the hymenia develop. Whereas in Plectocarpon
and Perigrapha development of hymenia happens from a plurilocular centrum, so
far this could not be clearly demonstrated in Phacothecium varium. Young
ascomata of P. varium seem to be always unilocular. Hemiangiocarpous
development of the hymenium with a relatively late exposure of the hymenium by
a slit-like to irregular rupture of the covering portions of the excipular envelope
are diagnostic. In longitudinal sections of older ascomata, a subdivision of the
hymenium by sterile portions is sometimes visible; however, this could also result
from secondary formation of hymenial lobes or lateral fusion of ascomata in
advanced stages of development. For Phacographa individually developing
ascomata with a persistently unilocular centrum and an apothecioid appearance
(hemiangiocarpous stage short and early degradation of the plectenchyma
obtecting the hymenium by crumbling) from an early stage of development on is a
diagnostic complex of characters.
Concerning its ascomatal characters, Phacographa is to some extent similar to
Lecanographa Egea & Torrente (type species: Lecanographa lyncea (Sm.) Egea
& Torrente). Lecanographa is one of several split-offs from Lecanactis (EGEA &
TORRENTE 1994), being now the most species rich genus in this clade. Under the
dissecting microscope Lecanographa species are easily recognized by the
subhysterothecioid shape of the ascomata in combination with the constantly
present whitish pruina, consisting of crystals insoluble in K and deposited on the
margin and the widely exposed hymenial surface, whereas apothecial discs of
Phacographa species are never pruinose. There is a certain tendency in
Lecanographa to lichenicolous growth, whereas the other crustose genera
Lecanactis s.str. (the Lecanactis abietina group), Cresponea (the former
Lecanactis premnea group), and Bactrospora appear to be strictly lichenized.
Lichenicolous behaviour in Lecanographa is either facultative as in L. grumulosa,
L. hemisphaerica (J.R.Laundon) Egea & Torrente and L. occidentalis Egea &
Torrente (EGEA et al. 1993, EGEA & TORRENTE 1994) or obligate as in L.
rinodinae. Lecanographa grumulosa is a frequent invader of Roccella thalli with
remarkable effects on their hosts (FEIGE et al. 1993) but is capable to grow also on
other Roccellaceae, Lecanographa hemisphaerica has been reported upon Dirina
massiliensis and the neotropical Lecanographa occidentalis can grow on
Roccellina species. The only obligately lichenicolous species so far known is
Lecanographa rinodinae (VĚZDA 1969), a rare species upon Phaeorrhiza nimbosa
with an arctic-alpine distribution pattern.
For distinguishing characters among the treated and some further genera see
also Table 3!

113
Key to lichenicolous opegraphoid fungi with widely exposed discs (compare
also the key provided by ERTZ et al. 2005: 7-8)
1 Ascomatal disc elongated, cleft-like at first, at least finally widely exposed 2
1* Ascomata persistently apothecioid, ascomatal disc roundish, widely exposed
from the beginning, a breakdown of the obtecting layer hardly observable,
ascomatal margin rounded ...............................................................................6
2 Ascomata subhysterothecioid, discs obtected by whitish pruina ..................... 3
2* of various shape, ascomatal disc dark brown to black, not conspicuously
pruinose ............................................................................................................4
3 Ascomata on thalli of Roccellaceae (Roccella, Dirina) .....................................
... Lecanographa grumulosa, L. hemisphaerica, or L. occidentalis (not treated)
3* Ascomata developed at the edge of areoles and apothecia of Phaeorrhiza
nimbosa .................................................. Lecanographa rinodinae (not treated)
4 Mature ascospores with heavily pigmented proper wall .....................................
............................................................Opegrapha pulvinata group (not treated)
4* Mature ascospores with hyaline proper wall, ascomata becoming lobed to
divided with age, discs oval to elongated, exposed by a breakdown of the
covering layer, ascomatal margin sharp............................................................ 5
5 Asci mostly 8-spored, without an apical hemiamyloid ring, on foliose
Teloschistaceae.................................................................Phacothecium varium
5* Asci mostly 4-spored, with a hemiamyloid ring apically in the endoascal wall
layer, on Pertusaria and Ochrolechia ................................. Opegrapha anomea
6 Ascomata solitary, with black disc, at the edge of Physconia thalli ...................
.........................................................................................“Opegrapha” rotunda
6* Ascomata dispersed or in dense groups, with dark brown discs, on crustose
lichens................................................................................................................7
7 Ascomata dispersed, 0.2-0.4 mm diam., developing on the thallus of Phlyctis
....................................................................................... Phacographa zwackhii
7* Ascomata larger, developing on saxicolous lichens .........................................8
8 Ascomata semi-immersed, hardly protruding over the thallus surface of the
host, ascospores 20-24(-26) × 6-7 μm, on species of the Lecanora rupicola
group ....................................................................... Phacographa glaucomaria
8* Ascomata semi-immersed only at the beginning, later sessile, ascospores 18-
21(-23) × 6-7 μm, on species of the Protoparmelia badia group .......................
............................................................................ Phacographa protoparmeliae

114
Table 3. Phenotypic characters of some opegraphoid arthonialean genera comprising
lichenicolous species.
character Opegrapha Lecanograph
a
Phacotheciu
m
Phacographa Plectocarpon
shape of
ascomata
elongated elongated to
oval
young
roundish, later
oval to
elongated
roundish elongated to
roundish
arrangement
of ascomata
solitary
dispersed
solitary
dispersed
agglomerated,
agglomeration
s dispersed
solitary,
dispersed or in
dense groups
composite,
enclosed in
stromatic
plectenchyma
centrum /
number of
ascomatal
discs per
centrum
unilocular / 1 unilocular / 1 uni- to
?oligolocular /
1–c..5
unilocular / 1 plurilocular /
n
shape of
ascomatal
discs
elongated,
sometimes
branched
elongated elongated ± round various,
punctiform to
elongated
colour of
ascomatal disc
black black black,
sometimes
with brownish
tinge
dark brown various
apothecial
pruina
frequently
absent, rarely
present, white
to greenish or
orange
whitish,
always present
absent absent mostly absent,
rarely present,
orange to red
interior ascal
wall and
reaction in K/I
with small
ocular
chamber, with
distinct
hemiamyloid
ring
with small
ocular
chamber, with
distinct
hemiamyloid
ring
ocular
chamber
reduced and
without
hemiamyloid
ring
with small
ocular
chamber, with
distinct
hemiamyloid
ring
with small
ocular
chamber, with
distinct
hemiamyloid
ring
septa of
ascospores
(1–)3–n 3–n 3 3–5 1–n
pigmentation
of proper
ascospore wall
lacking to
intense
lacking lacking lacking lacking
life habit lichenized or
lichenicolous
lichenized or
lichenicolous
obligately
lichenicolous
obligately
lichenicolous
obligately
lichenicolous
Acknowledgements
The author gratefully acknowledges the support by O. Vitikainen (Helsinki) who
arranged a loan of relevant type specimens as well as by W. Obermayer (Graz) for
help in preparing the illustrations and B. Coppins (Edinburgh), T. Feuerer
(Hamburg), M. Seaward (Bradford), and A. Thell (Lund) for critical reading.

115
References
ARNOLD, F. (1868): Die Lichenen des fränkischen Jura. Flora 51: 520-524.
ARNOLD, F. (1887): Lichenologische Ausflüge in Tirol XXIII. Predazzo und Paneveggio.
– Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 37: 81-150.
ARNOLD, F. (1889): Lichenologische Ausflüge in Tirol. XXIV. Finkenberg. Nachträge. –
Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 39: 249-266.
ARNOLD, F. (1896): Lichenologische Ausflüge in Tirol. XXVI. Pians. XXVII. Galtür.
XXVIII. Wolkenstein. XXIX. Plansee. Nachträge. – Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien 46: 101-143.
ATIENZA, V. (1992): Peridothelia oleae (Körber) D. Hawksw. and Opegrapha physciaria
(Nyl.) D. Hawksw. & Coppins, two poorly known west mediterranean fungal taxa. –
Anales del Jardín Botánico de Madrid 50: 159-162.
ATIENZA, V. & SEGARRA MORAGUES, J. (1999): Fragmenta chorologica occidentalia,
Lichenes, 7040-7082. – Anales del Jardín Botánico de Madrid 57: 148-151.
BAGLIETTO, F. (1871): Prospetto lichenologico della Toscana. – Nuovo Giornale
Botanico Italiano 3: 211-298.
BAUSCH, W. (1869): Übersicht der Flechten des Grossherzogthums Baden. –
Verhandlungen des Naturwissenschaftlichen Vereins Carlsruhe 4: I-XLII, 1-246.
BOBERSKI, L. (1886): Systematische Uebersicht der Flechten Galiziens.
Zusammengestellt auf Grundlage eigener und fremder Forschungen. – Verhandlungen
der Zoologisch-Botanischen Gesellschaft in Wien 36: 243-286.
BOOM, P. P. G. van den (1999): Some lichens and lichenicolous fungi from Majorca. –
Linzer Biologische Beiträge 31: 785-800.
BOOM, P. P. G. van den & BREUSS, O. (2002): Lichenen van het zomerkamp in de Cantal
(Frankrijk) zomer 1998. – Buxbaumiella 60: 3-16.
BOOM, P. P. G. van den, BREUSS, O., SPIER, L. & BRAND, A. M. (1996): Beitrag zur
Flechtenflora Kärntens. Ergebnisse der Feldtagung der bryologischen und
lichenologischen Arbeitsgruppe der KNNV in Weißbriach 1994. – Linzer Biologische
Beiträge 28: 619-654.
BOULY DE LESDAIN, M. (1914) : Recherches sur les lichens des environs de Dunkerque.
1er Supplément. P. Michel, Dunkerque, 189 pp.
BRANTH, J. S. DEICHMANN & ROSTRUP, E. (1869): Lichenes Daniae eller Danmarks
Laver. – Botanisk Tidsskrift 3: 127-284.
BRUMMITT, R. K. & POWELL, C. E. (1992): Authors of plant names. Royal Botanic
Gardens, Kew, 732 pp.
CALATAYUD, V., ATIENZA, V. & BARRENO, E. (1995): Lichenicolous fungi from the
Iberian peninsula and the Canary Islands 1. – Mycotaxon 55: 363-382.
COLE, M. S. & HAWKSWORTH, D. L. (2001): Lichenicolous fungi, mainly from the USA,
including Patriciomyces gen. nov. – Mycotaxon 77: 305-338.
COSTE, C. (1993) : Contribution a l’étude des champignons lichénisés ou lichénicoles de
la région Midi-Pyrénées. – Bulletin d'Informations de l'Association Française de
Lichénologie 18: 3-15.
EGEA, J. M. & TORRENTE, P. (1994): El género de hongos liquenizados Lecanactis
(Ascomycotina). – Bibliotheca Lichenologica 54: 1-205.
EGEA, J. M., TORRENTE, P. & MANRIQUE, E. (1993): The Lecanactis grumulosa group
(Opegraphaceae) in the Mediterranean region. – Plant Systematics and Evolution 187:
103-114.
EITNER, E. (1901): II. Nachtrag zur schlesischen Flechtenflora. – Jahresbericht der
Schlesischen Gesellschaft für Vaterländische Kultur 78: 5-27.
ERICHSEN, F. (1906): Beitrag zur Flechtenflora der Umgebung von Hamburg und
Holsteins. – Verhandlungen des Naturwissenschaftlichen Vereins Hamburg, 3. Folge,
13: 44-104.

116
ERICHSEN, F. (1914): Die Flechten von Kullen in Schweden. – Verhandlungen des
Naturwissenschaftlichen Vereins Hamburg, 3. Folge, 21: 25-94.
ERIKSSON, O. E. & HAWKSWORTH, D. L. (1998): Outline of the ascomycetes - 1998. –
Systema Ascomycetum 16: 83-296.
ERTZ, D., DIEDERICH, P. & MIADLIKOWSKA, J. (2004): The lichenicolous Opegrapha
species (Roccellaceae, Ascomycota) with 3-septate ascospores on Pertusaria and
Ochrolechia. – Botanical Journal of the Linnean Society 144: 235-241.
ERTZ, D., CHRISTNACH, C., WEDIN, M. & DIEDERICH, P. (2005): A world monograph of
the genus Plectocarpon (Roccellaceae, Arthoniales). – Bibliotheca Lichenologica 91:
1-155.
ETAYO, J. (1996) : Contribución al conocimiento de los líquenes y hongos liquenícolas de
Mallorca (Islas Baleares, España). – Bulletin de la Société Linnéenne de Provence 47:
111-121.
FEIGE, G. B., LUMBSCH, H. T. & MIES, B. (1993) : Morphological and chemical changes
in Roccella thalli infected by Lecanactis grumulosa (lichenized Ascomycetes,
Opegraphales). – Cryptogamic Botany 3: 101-107.
FINK, B. (1935): The Lichen Flora of the United States. (Completed for Publication by
Joyce Hedrick.) University of Michigan Press, Ann Arbor,
FOS, S. & BARRENO, E. (2002): Fragmenta chorologica occidentalia, Lichenes, 8106-
8141. – Anales del Jardín Botánico de Madrid 59: 310-311.
GALLOWAY, D. J. (2007): Flora of New Zealand: Lichens. (Revised second edition
including lichen-forming and lichenicolous fungi.) Volume 1: Abrothallus–
Pachyphiale. Manaaki Whenua Press, Lincoln, New Zealand, 1006 pp.
GILBERT, O. L. & COPPINS, B. J. (1992): The lichens of Caenlochan, Angus. –
Lichenologist 24: 143-163.
GIRALT, M. (1996): Líquens epífits i contaminació atmosfèrica a la plana i les serralades
litorals tarragonines. Institut d'Estudis Catalans, Secció de Ciències Biològiques,
Barcelona, 525 pp.
GIRALT, M. & GOMEZ-BOLEA, A. (1992): Alguns fongs liquenícoles trobats sobre
líquens epífits de Catalunya. – Actes del Simposi Internacional de Botánica Pius Font
i Quer, 1988 (Cryptogàmia) 1: 195-203.
GRILLO, M., CARNEMOLLA, G. & CARFI, M. G. (2000): I licheni della valle dell’Ippari
presso Vittoria e della zona archeologica di Camarina (Sicilia orientale). – Archivio
Geobotanico 6: 45-58.
GRUBE, M. & MATZER, M. (1997): Taxonomic concepts of lichenicolous Arthonia
species. – Bibliotheca Lichenologica 68: 1-17.
HAFELLNER, J. (1994a): Über Funde lichenicoler Pilze und Flechten auf Korsika
(Frankreich). – Bulletin de la Société Linnéenne de Provence 45 (Hommage
scientifique à G. Clauzade): 219-234.
HAFELLNER, J. (1994b): Beiträge zu einem Prodromus der lichenicolen Pilze Österreichs
und angrenzender Gebiete I. Einige neue oder seltene Arten. – Herzogia 10: 1-28.
HAFELLNER, J. (1995): Bemerkenswerte Funde von Flechten und lichenicolen Pilzen auf
makaronesischen Inseln II. Einige bisher übersehene arthoniale Arten. – Herzogia 11:
133-142.
HAFELLNER, J. (1998): Studien an lichenicolen Pilzen und Flechten X. Arthophacopsis,
eine neue Gattung lichenicoler Ascomyceten (Arthoniales). – Cryptogamie, Bryologie
et Lichénologie 19: 155-168.
HAFELLNER, J. (2002): Zur Diversität lichenisierter Pilze und ihrer Parasiten in den
Seckauer Tauern (Ostalpen, Niedere Tauern, Steiermark). – Mitteilungen der
Naturwissenschaftlichen Vereines für Steiermark 132: 83-137.
HAFELLNER, J. (2003): Ein Beitrag zur Flechtenflora der Fischbacher Alpen (Steiermark).
– Fritschiana 41: 21-40.

117
HAFELLNER, J. (2005): Bemerkenswerte Funde von Flechten und lichenicolen Pilzen auf
makaronesischen Inseln VII. Über einige bisher auf Lanzarote und Fuerteventura
übersehene Arten. – Fritschiana 50: 15-24.
HAFELLNER, J. & JOHN, V. (2006): Über Funde lichenicoler nicht-lichenisierter Pilze in
der Türkei, mit einer Synopsis der bisher im Land nachgewiesenen Taxa. – Herzogia
19: 155-176.
HAFELLNER, J. & OBERMAYER, W. (2001): Ein Beitrag zur Flechtenflora der Murberge
(Steiermark, Österreich). – Fritschiana 25: 19-32.
HAFELLNER, J. & OBERMAYER, W. (2007): Flechten und lichenicole Pilze im Gebiet der
Stubalpe (Österreich: Steiermark und Kärnten). – Mitteilungen der
Naturwissenschaftlichen Vereines für Steiermark 136: 5-59.
HAFELLNER, J. & SANCHO, L. G. (1990): Über einige lichenicole Pilze und Flechten aus
den Gebirgen Zentralspaniens und den Ostalpen. – Herzogia 8: 363-382.
HAFELLNER, J. & TÜRK, R. (1995): Über Funde lichenicoler Pilze und Flechten im
Nationalpark Hohe Tauern (Kärntner Anteil, Österreich). – Carinthia II 185/105: 599-
635.
HAFELLNER, J., TRIEBEL, D., RYAN, B. D., & NASH III, T. H. (2002): On lichenicolous
fungi from continental North America II. – Mycotaxon 84: 293-329.
HAFELLNER, J., KOCOURKOVÁ, J. & OBERMAYER, W. (2004): Records of lichenicolous
fungi from the northern Schladminger Tauern (Eastern Alps, Austria, Styria). –
Herzogia 17: 59-66.
HAUCK, M. (1996): Die Flechten Niedersachsens. Bestand, Ökologie, Gefährdung und
Naturschutz. – Naturschutz und Landschaftspflege in Niedersachsen 36: 1-208.
HAWKSWORTH, D. L. (1994): Notes on British lichenicolous fungi: VII. Lichenologist
26: 337–347.
HAWKSWORTH, D. L. (2003): The lichenicolous fungi of Great Britain and Ireland: an
overview and annotated checklist. – Lichenologist 35: 191-232.
HAWKSWORTH, D. L. & ATIENZA, V. (1994): The lichenicolous fungi described by Veli
Räsänen. – Acta Botanica Fennica 150: 47-77.
HERNÁNDEZ PADRÓN, C. E., SICILIA MARTÍN, D., PÉREZ VARGAS, I., PÉREZ DE PAZ, P.
L. & ETAYO SALAZAR, J. (2004): Líquenes y hongos liquenícolas. In: Beltrán Tejera,
E. (ed.). Hongos, líquenes y briófitos del Parque Nacional de la Caldera de
Taburiente. Organísmo Autónomo de Parque Nacionales. Ministerio de Medio
Ambiente, Madrid, pp. 233-350
HITCH, C. (ed.) (1995): New, rare or interesting British lichen records. – British Lichen
Society Bulletin 77: 37-42.
HITCH, C. (ed.) (1997): New, rare or interesting British lichen and lichenicolous fungus
records. – British Lichen Society Bulletin 80: 46-58.
HITCH, C. (ed.) (2006): New, rare and interesting lichens. – British Lichen Society
Bulletin 99: 34-46.
HITCH, C. (ed.) (2007a): New, rare and interesting lichens. – British Lichen Society
Bulletin 100: 72-90.
HITCH, C. (ed.) (2007b): New, rare and interesting lichens. – British Lichen Society
Bulletin 101: 66-86.
HOLIEN, H. (2001): Additions to the Norwegian flora of lichens and lichenicolous fungi
II - with some further distributional notes on Norwegian Caliciales. – Graphis Scripta
12: 51-58.
HOLMGREN, P. K., HOLMGREN, N. H. & BARNETT, L. C. (eds) (1990): Index
herbariorum. Part I. The herbaria of the world. 8th edition. Regnum Vegetetabile 120.
New York Botanical Garden for the International Association for Plant Taxonomy,
Bronx, New York, 693 pp.
ISBRAND, S. & ALSTRUP, V. (1992): Opegrapha sphaerophoricola sp. nov. – Bryologist
95: 233-234.

118
JATTA, A. (1882): Lichenum Italiae meridionalis manipulus quartus, quem collegit et
ordinavit. – Nuovo Giornale Botanico Italiano 14: 107-143.
JATTA, A. (1889): Monographia lichenum Italiae meridionalis. V. Vecchi, Trani, 261 pp.
JATTA, A. (1892): Licheni raccolti nell'isola d'Ischia fino all'Agosto del 1891. – Bollettino
della Società Botanica Italiano 1892: 206-211.
JATTA, A. (1909-1911): Lichenes. – Flora italica cryptogama 3: III-XXII, 1-958. – Rocca
S. Casciano: Tip. Cappelli. [Fasc. 1: 1-112 (1909); 2: 113-264 (1909); 3: 265-460
(1910); 4: 461-694 (1911); 695-776 (1911); 777-958, III-XXII (1911)]
KALB, K. & HAFELLNER, J. (1992): Bemerkenswerte Flechten und lichenicole Pilze von
der Insel Madeira. – Herzogia 9: 45-102.
KEISSLER, K. v. (1914): Fungi. – In: K. Rechinger, Beiträge zur Kryptogamenflora der
Insel Korfu nebst einigen Standorten von der albanischen Küste. – Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien 64: 143-149.
KEISSLER, K. v. (1930): Die Flechtenparasiten. – In: Dr. L. Rabenhorst’s Kryptogamen-
Flora von Deutschland, Österreich und der Schweiz, 2. Auflage, Band 8.
Akademische Verlagsgesellschaft, Leipzig, 712 pp.
KERNSTOCK, E. (1895): Lichenologische Beiträge. VI, [Nachträge] zu II, III. –
Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 44: 191-224.
KNOWLES, M. C. (1929): The lichens of Ireland. – Proceedings of the Royal Irish
Academy, Section B 38: 179-434.
KOCOURKOVÁ, J. (2000): Lichenicolous fungi of the Czech Republic (The first
commented checklist). – Acta Musei Nationalis Pragae. Sborník Nárosního Muzea V
Praze. Series B, Historia Naturalis 55: 59-169.
KONDRATYUK, S. Y. & KHODOSOVTSEV, A. Y. (1997): Novi dlja mikobioty Ukrainy
vydy lichenofilnich hrybiv. – Ukrainian Journal of Botany 54: 588-590.
KONDRATYUK, S. Y., NAVROTSKAYA, I. L., ZELENKO, S. D., WASSER, S. P. & NEVO, E.
(1996): The First Checklist of Lichen-forming and Lichenicolous Fungi of Israel. M.
G. Kholodny Institute of Botany & International Centre for Cryptogamic Plants and
Fungi, Haifa, 136 pp.
LAHM, G. (1882-1885): Zusammenstellung der in Westfalen beobachteten Flechten. –
Jahres-Bericht des Westfälischen Provinzial-Vereins für Wissen-schaft und Kunst 10:
117-172 (1882); 11: 98-153 (1883); 12: 126-149 (1884); 13: 57-85 (1885).
LEIGHTON, W. A. (1879): The Lichen-Flora of Great Britain, Ireland and the Channel
Islands. 3rd ed. Published by the author, Shrewsbury, 547 pp.
LETTAU, G. (1912a): Beiträge zur Lichenographie von Thüringen. – Hedwigia 52: 81-
264.
LETTAU, G. (1912b): Beiträge zur Lichenenflora von Ost- und Westpreussen. –In:
Caspary, R., Festschrift zum 50-jährigen Bestehen des Preußischen Botanischen
Vereins. Königsberg, pp. 17-91.
LETTAU, G. (1958): Flechten aus Mitteleuropa XIV. – Feddes Repertorium Specierum
Novarum Regni Vegetabilis 61: 105-171.
LEUCKERT, C. & POELT, J. (1989): Studien über die Lecanora rupicola-Gruppe in
Europa (Lecanoraceae). – Nova Hedwigia 49: 121-167.
LISICKÁ, E. & LACKOVICOVÁ, A. (1999): Predbezny zoznam lichenikolnych hub
Slovenska (A preliminary checklist of lichenicolous fungi of Slovakia). –In: A.
Leskovjanska (ed.). Zbornik Referatov, zo 7. zjazdu Slovenskej botanickej spolocnosti
pri SAV, Hrabusice-Poklesok. Spisska Nova Ves, pp. 201-203.
LITTERSKI, B. & MAYRHOFER, H. (1998): Catalogue of lichenized and lichenicolous
fungi of Cyprus. – Studia Geobotanica 16: 57-70.
LITTERSKI, B. & MAYRHOFER, H. (2000): Additional records to the lichen flora of
Cyprus. – Herzogia 14: 145-150.
LOJKA, H. (1876): Adatok Magyarhon zuzmó-virányához. II. – Mathematikai es
Termeszettudomanyi Kozlememyek 12: 89-129.

119
LUMBSCH, H. T. & HUHNDORF, S. M. (eds.) (2007): Outline of Ascomycota - 2007.
Myconet 13: 1-58.
MEYER, B. & PRINTZEN, C. (2000): Proposal for a standardized nomenclature and
characterization of insoluble lichen pigments. – Lichenologist 32: 571-583.
MÜLLER (Argoviensis), J. (1880): Les lichens d’Egypte. – Revue Mycologique 2: 38-44,
73-83.
NAVARRO-ROSINES, P., BOQUERAS, M. & LLIMONA, X., (1994): Primer catàleg dels
fongs liquenícoles de Catalunya i zones pròximes (NE de la Península Ibèrica). –
Buttletí de la Societat Catalana de Micologia 16-17: 165-204.
NAVROTSKAYA, I. L., KONDRATYUK, S. Y., WASSER, S. P., NEVO, E. & ZELENKO, S. D.
(1996): Lichens and lichenicolous fungi new for Israel and other countries. – Israel
Journal of Plant Sciences 44: 181-196.
NIMIS, P. L., POELT, J., TRETIACH, M., OTTONELLO, D., PUNTILLO, D. & VĚZDA, A.
(1994): Contributions to lichen floristics in Italy VII. – The lichens of Marettimo
(Egadi Islands, Sicily). – Bulletin de la Société Linnéenne de Provence 45 (Hommage
scientifique à G. Clauzade): 247-262.
NYLANDER, W. (1852): Observationes aliquot ad Synopsin Lichenum Holmiensium. –
Nya Botaniska Notiser 1852: 175-180.
NYLANDER, W., (1857a): Note sur les lichens recueillis en Auvergne pendant la session
extraordinaire de la société, en juillet 1856. – Bulletin de la Societé Botanique de
France 3: 548-552.
NYLANDER, W. (1857b): Prodromus Galliae et Algeriae, quem conscripsit. – Actes de la
Société Linnéenne de Bordeaux 21: 249-467.
NYLANDER, W. (1862): Ad lichenographiam Groenlandiae quaedam addenda. – Flora
45: 81-83.
NYLANDER, W. (1866): Addenda nova ad lichenographiam europaeam. Continuatio
altera. – Flora 49: 369-374.
NYLANDER, W. (1885): Arthoniae novae Americae borealis. – Flora 68: 447-449.
NYLANDER, W. (1887): Enumeratio lichenum Freti Behringii. – Bulletin de la Société
Linnéenne de Normandie, sér. 4, 1: 198-286.
NYLANDER, W. (1897): Supplément aux Lichens des Environs de Paris. P. Schmidt,
Paris, 20 pp.
OLIVIER, H. (1884): Flore analytique et dichotomique des lichens de l'Orne et
départements circonvoisins. Deuxième partie. F. Savy, Autheuil (Orne) & Paris,
Friedlander et Sohn, Berlin, pp. I-IV, 127-312.
OLIVIER, H. (1903): Exposé systématique et description des lichens de l'Ouest et du
Nord-Ouest de la France (Normandie, Bretagne, Anjou, Maine, Vendée). Deuxième
partie, fasc. 4. P. Klincksieck, Paris, pp. 271-426.
OLIVIER, H. (1906): Les principaux parasites de nos lichens francais. (Suite). – Bulletin
de l'Académie Internationale de Géographie Botanique 16: 42-48, 187-200, 253-264.
OLIVIER, H., (1907): Les principaux parasites de nos lichens francais. Premier
supplément. Bazoches-au-Houlme: chez l’autheur. P. Klincksieck, Paris.
PITARD, C.-J. & HARMAND, J. (1911): Contribution à l’étude de lichens des îles Canaries.
– Bulletin de la Société Botanique de France 58, Mémoires 22: 1-72.
POELT, J. & TÜRK, R. (1984): Die Flechten des Lungau - ein erstes Verzeichnis –
zugleich Bericht über die lichenologischen Ergebnisse der Lungau-Exkursion der
Bryologisch-Lichenologischen Arbeitsgemeinschaft für Mitteleuropa im September
1981. – Herzogia 6: 419-469.
POETSCH, J. S. & SCHIEDERMAYR, K. B. (1872): Systematische Aufzählung der im
Erzherzogthume Oesterreich ob der Enns bisher beobachteten samenlosen Pflanzen
(Kryptogamen). W. Braumüller, Wien, F. A. Brockhaus, Leipzig.
RÄSÄNEN, V. (1946): Lichenes novi. II. – Annales Botanici Societatis Zoologicae-
Botanicae Fennicae ‘Vanamo’ 21, Notulae Botanicae 16: 1-7.

120
REHM, H. (1887-1896): Ascomyceten: Hysteriaceen und Discomyceten. – Dr. L.
Rabenhorst’s Kryptogamen-Flora von Deutschland, Österreich und der Schweiz, 2.
Auflage, Band. 1, Abteilung. 3: E. Kummer, Leipzig, 1275 pp. [p. 1-64, 1887; 65-208,
1888; 209-336, 1889; 337-400, 1890; 401-608, 1891a; 609-720, 1892; 721-912, 1893;
913-1040, 1894; 1041-1104, 1895; 1105-1275, 1896.]
RENOBALES, G. & BARRENO, E. (1989): Estudio al microscopio óptico de los ascos en
algunas especies de Arthoniales y Opegraphales. – Anales del Jardín Botánico de
Madrid 46: 263-271.
ROBERT, V., STEGEHUIS G. & STALPERS, J. (2005): The MycoBank engine and related
databases. – http://www.mycobank.org (visited 30.4.2008)
RONDON, Y. (1969) : L’herbier des champignons parasites des lichens de l’abbé L.
Vouaux. – Revue Bryologique et Lichénologique 36: 737-745.
ROUX, C. (1977) : Champignons lichénisés ou lichénicoles intéressants pour la France
méridionale (II). – Bulletin du Muséum d'Histoire Naturelle de Marseille 37: 83-92.
ROUX, C. (1982) : Lichens observés lors de la 8e session extraordinaire de la Société
Botanique du Cente-Ouest en Provence occidentale. – Bulletin de la Société Botanique
du Centre-Ouest, Nouvelle Série 13: 210-228.
ROUX, C., COSTE, C., BRICAUD, O. & MASSON, D. (2006) : Catalogue des lichens et des
champignons lichénicoles de la région Languedoc–Roussillon (France méridionale). –
Bulletin de la Société Linnéenne de Provence 57: 85-200.
SAMPAIO, G. (1917) : Líquenes novos para a flora portuguesa (3.A série). – Brotéria,
Série Botânica 15: 125-145.
SANDSTEDE, H. (1889): Beiträge zu einer Lichenenflora des nordwestdeutschen
Tieflandes. – Abhandlungen herausgegeben vom Naturwissenschaftlichen Verein zu
Bremen 10: 439-480.
SANTESSON, R. (1949): Svampar som leva på lavar. – Svensk Botanisk Tidskrift 43: 141-
143.
SANTESSON, R. (1960): Lichenicolous fungi from northern Spain. – Svensk Botanisk
Tidskrift 54: 499-522.
SANTESSON, R. (1986): Fungi lichenicoli exsiccati. Fasc. 3 & 4 (Nos 51-100). –
Thunbergia 3: 1-18.
SANTESSON, R. (1993): The Lichens and Lichenicolous Fungi of Sweden and Norway.
SBT-förlaget, Lund.
SANTESSON, R. (2001): Fungi lichenicoli exsiccati. Fascicles 13 & 14 (Nos 301-350). –
Thunbergia 31: 1-18.
SIMMER, H. (1898): Erster Bericht über die Kryptogamenflora der Kreuzeckgruppe in
Kärnten. – Allgemeine Botanische Zeitung 4: 74-77, 99-100, 118-120, 141-144, 158-
159.
SIMMER, H. (1900): Simmer, Hans, Kryptogamen des Kreuzeckgebietes. – Botanisches
Centralblatt 81: 299-300.
STEIN, B. (1872): Proskau’s Flechten. – Verhandlungen des Botanischen Vereins der
Provinz Brandenburg 14: 51-57.
STEIN, B. (1879): Flechten. – In: Cohn, F. (ed.). Kryptogamen-Flora von Schlesien, Band
2, 2. Hälfte. J. U. Kern, Breslau, 400 pp.
STIZENBERGER, E. (1876): Index lichenum hyperboreorum. – Berichte über die
Thätigkeit der St. Gallischen Naturwissenschaftlichen Gesellschaft 1874-75: 189-245.
STIZENBERGER, E. (1890): Lichenaea africana. Fasc. prior. – Berichte über die Thätigkeit
der St. Gallischen Naturwissenschaftlichen Gesellschaft 1888-89: 105-149.
SUIJA, A. (2005): Lichenicolous fungi and lichens in Estonia I. Ascomycota. – Nova
Hedwigia 80: 247-267.
TEHLER, A. (1990): A new approach to the phylogeny of Euascomycetes with a cladistic
outline of Arthoniales focussing on Roccellaceae. – Canadian Journal of Botany 68:
2458-2492.

121
THOR, G. (1993): Lavfloran på Visingsö. – Graphis Scripta 5: 105-116.
TRETIACH, M. & HAFELLNER, J. (2000): Lichens and lichenicolous fungi of M. Fleons
(Carnic Alps, Italy). I. Epilithic and epigaeic species. – Herzogia 14: 99-111.
TORRENTE, P. & EGEA, J. M. (1989): La Familia Opegraphaceae en el area Mediterranea
de la peninsula Iberica y norte de Africa. – Bibliotheca Lichenologica 32: 1-282.
TREVISAN, V. (1856): Brigantiaea, novum lichenum genus. – Linnaea 28: 283-298.
TRIEBEL, D. & SCHOLZ, P. (2001): Lichenicolous fungi from Bavaria as represented in
the Botanische Staatssammlung München Sendtnera 7: 211-231.
TUCKERMAN, E. (1888): A Synopsis of the North American Lichens: Part II., comprising
the Lecideacei, and (in part) the Graphidacei. E. Anthony, New Bedford.
TÜRK, R. & BREUSS, O. (1994): Flechten aus Niederösterreich I. Steirisch-
niederösterreichische Kalkalpen. – Verhandlungen der Zoologisch-Botanischen
Gesellschaft in Österreich 131: 79-96.
VĚZDA, A. (1969): Beiträge zur Kenntnis der flechtenbewohnenden Pilze in der
Tschechoslowakei. II. – Zwei neue Arten: Opegrapha rinodinae sp. nov. und
Polycoccum galligenum sp. nov. Príspeveky k poznání lichenikolnich hub v
Ceskoslovensku. II. – Dva nové druhy: Opegrapha rinodinae sp. nov. a Polycoccum
galligenum sp. nov. – Ceská Mykologie 23: 104-109.
VĚZDA, A. (1970): Príspevek k poznání lichenikolnich hub v Ceskoslovensku III. Beitrag
zur Kenntnis der flechtenbewohnenden Pilze aus der Tschechoslowakei III. – Ceská
Mykologie 24: 220-229.
VOUAUX, L. (1913): Synopsis des champignons parasites de lichens. – Bulletin
Trimestriel de la Société Mycologique de France 29: 33-128, 399-446, 447-494.
VOUAUX, L. (1914): Synopsis des champignons parasites de lichens. – Bulletin
Trimestriel de la Société Mycologique de France 30: 135-198, 281-329.
ZHURBENKO, M. & SANTESSON, R. (1996): Lichenicolous fungi from the Russian Arctic.
– Herzogia 12: 147-161.
ZWACKH, W. (1862): Enumeratio lichenum florae heidelbergensis. – Flora 45: 467-474,
481-495, 504-512, 518-527, 530-537, 546-552, 561-572.

122