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Monographiae Botanicae 107
Anetta Wieczorek
The lichen genus Opegrapha s. l. in
Poland: morphological variability,
ecology, and distribution
Monographiae Botanicae 107
Official publication of the Polish Botanical Society
Anetta Wieczorek
The lichen genus Opegrapha s. l. in
Poland: morphological variability,
ecology, and distribution
Wrocław 2018
Editor-in-Chief of the series
Zygmunt Kącki, University of Wrocław, Poland
Honorary Editor-in-Chief
Krystyna Czyżewska, University of Łódź, Poland
Chairman of the Editorial Council
Jacek Herbich, University of Gdańsk, Poland
Editorial Council
Idoia Biurrun, University of the Basque Country, Spain
Gian Pietro Giusso del Galdo, University of Catania, Italy
Jan Holeksa, Adam Mickiewicz University in Poznań, Poland
Czesław Hołdyński, University of Warmia and Mazury in Olsztyn, Poland
Bogdan Jackowiak, Adam Mickiewicz University in Poznań, Poland
Zbigniew Mirek, W. Szafer Institute of Botany, Polish Academy of Sciences, Poland
Valentina Neshataeva, Komarov Botanical Institute of the Russian Academy of Sciences, Russian Federation
Marcin Nobis, Jagiellonian University, Poland
Arkadiusz Nowak, University of Opole, Poland
Vilém Pavlů, Crop Research Institute, Czech Republic
Agnieszka Anna Popiela, University of Szczecin, Poland
Lucyna Śliwa, W. Szafer Institute of Botany, Polish Academy of Sciences, Poland
Iveta Škodová, Slovak Academy of Sciences, Slovakia
David Zelený, National Taiwan University, Taiwan
Jan Żarnowiec, University of Bielsko-Biala, Poland
Editorial Secretary
Grzegorz Swacha, University of Wrocław, Poland
Managing/Production Editor
Piotr Otręba, Polish Botanical Society, Poland
Reviewers of the volume
Damien Ertz, Botanic Garden Meise, Belgium
Laszlo Lőkös, Hungarian Natural History Museum, Hungary
Lucyna Śliwa, W. Szafer Institute of Botany, Polish Academy of Sciences, Poland
Editorial office
University of Wrocław
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email: mb@pbsociety.org.pl
e-ISSN: 2392-2923 e-ISBN: 978-83-950171-0-0
p-ISSN: 0077-0655 p-ISBN: 978-83-950171-1-7
DOI: 10.5586/mb.2018.001
© The Author(s) 2018. This is an Open Access publication distributed under the terms of the
Creative Commons Attribution License, which permits redistribution, commercial and noncommercial, provided that the
original work is properly cited.
Citation: Wieczorek A. The lichen genus Opegrapha s. l. in Poland: morphological variability, ecology, and distribution. Wrocław:
Polish Botanical Society; 2018. (Monographiae Botanicae; vol 107). https://doi.org/10.5586/mb.2018.001
Published by Polish Botanical Society
Al. Ujazdowskie 4, 00-478 Warsaw, Poland
pbsociety.org.pl
Edited with financial assistance of the Polish Ministry of Science and Higher Education.
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Contents
1. Introduction
2. Research history of the genus
3. Material and methods
4. Results and discussion
4.1. Numerical analysis of the examined material
4.2. Morphological features
4.2.1. Thallus
4.2.2. Algae
4.2.3. Ascomata
4.2.4. Pycnidia
4.3. Ecology
4.4. Geographic distribution
4.5. Key to the species of Opegrapha s. l. found in Poland
4.6. Species characteristics
4.6.1. Alyxoria culmigena (Lib.) Ertz
4.6.2. Alyxoria mougeotii (A. Massal.) Ertz, Frisch & G. Thor
4.6.3. Alyxoria ochrocheila (Nyl.) Ertz & Tehler
4.6.4. Alyxoria varia (Pers.) Ertz & Tehler
4.6.5. Arthonia atra (Pers.) A. Schneid.
4.6.6. Arthonia calcarea (Turner ex Sm.) Ertz & Diederich
4.6.7. Gyrographa gyrocarpa (Flot.) Ertz & Tehler
4.6.8. Opegrapha dolomitica (Arnold) Clauzade & Cl. Roux ex Torrente
& Egea
4.6.9. Opegrapha geographicola (Arnold) Hafellner
4.6.10. Opegrapha lithyrga Ach.
4.6.11. Opegrapha niveoatra (Borrer) J. R. Laundon
4.6.12. Opegrapha saxicola Ach.
4.6.13. Opegrapha vermicellifera (Kunze) J. R. Laundon
4.6.14. Opegrapha vulgata (Ach.) Ach.
4.6.15. Phacographa glaucomaria (Nyl.) Hafellner
4.6.16. Pseudoschismatomma rufescens (Pers.) Ertz & Tehler
4.6.17. Zwackhia sorediifera (P. James) Ertz
4.6.18. Zwackhia viridis (Ach.) Poetsch & Schied.
5. Summary of results and conclusions
6. References
7. Appendix: list of examined herbarium specimens, literature records, and list of
surveyed localities
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About the author
Anetta Wieczorek; Department of Ecology and Environmental Protection, University of
Szczecin, Wąska 13, 71-415 Szczecin, Poland; email: anettaw@univ.szczecin.pl
Abstract
This monograph presents the results of research on the morphological and anatomical variability, ecology, and distribution of Opegrapha s. l. in Poland. The study is based on roughly
1,400 specimens from Polish and some European herbaria. Out of the 18 species of the genus
Opegrapha s. l. recorded from Poland, seven species belong to the genus Opegrapha Ach.,
four species now belong to the genus Alyxoria Ach. ex Gray, two species each are found in
the genera Arthonia Ach. and Zwackhia Körb, and one species belongs to each of the genera
Gyrographa Ertz & Tehler, Phacographa Hafellner, and Pseudoschismatomma Ertz & Tehler.
One of the species, Zwackhia sorediifera, has been reported from Poland for the first time.
Among the 18 species of the genus Opegrapha s. l., 10 are epiphytic, five epilithic, and two
lichenicolous. The first modern identification key for the species of Opegrapha s. l. in Poland is
presented. Numerous new regional records are provided that complement our knowledge of the
geographic distribution of some poorly known taxa, such as Alyxoria culmigena, A. mougeotii,
A. ochrocheila, Arthonia calcarea, Opegrapha dolomitica, O. geographicola, O. lithyrga, and
Phacographa glaucomaria. All species are characterized and discussed, and their diagnostic
characters illustrated. Geographic ranges of each species in Poland are presented on maps
based on revised herbarium materials.
Keywords
lichenized Ascomycota; Arthoniales; Opegraphaceae; Roccellaceae; species variability;
distribution maps; Poland
Acknowledgments
This work has been completed due to the kindness of many people. I am particularly grateful
to directors, curators, and private owners of herbaria for access to their collections. I would
like to sincerely thank Edyta Adamska (Toruń), Urszula Bielczyk (Cracow), Olga Biryukova
(Nizhny Novgorod, Russia), Krystyna Czyżewska (Łódź), Stanisław Cieśliński, Anna Łubek,
(Kielce), Wiesław Fałtynowicz (Wrocław), Paweł Czarnota (Poręba Wielka), Katarzyna Kolanko
(Białystok), Maria Kossowska (Wrocław), Robert Kościelniak (Cracow), Dariusz Kubiak
(Olsztyn), Martin Kukwa (Gdańsk), Aleey Melechin (Kirovsk, Russia), Jurga Motiejūnaitė
(Vilnius, Lithuania), Eugenia Muchnik (Moscow, Russia), Wiesław Mułenko (Lublin), Maria
Olech (Cracow), Žydrūnas Preikša (Vilnius, Lithuania), Ingrida Prigodina-Lukošiene (Vilnius,
Lithuania), Alfons Piteráns (Riga, Latvia), Lucyna Śliwa, Adam Flakus (Cracow), Mark Seaward
(Bradford, UK), Ulf Schiefelbein (Rostock, Germany), Katarzyna Szczepańska (Wrocław),
Andrei Tsurykau (Gomel, Belarus), Michał Węgrzyn (Cracow), and Waldemar Żukowski
(Poznań). I owe special thanks to Maciej Rogalski for valuable suggestions and corrections
to the manuscript. My sincere thanks are also due to Adam Zając and Agnieszka Popiela, for
their great kind-heartedness, continuous tracking of my research progress, and constructive
discussions. Besides, I am grateful to Magdalena Bihun, who spent many hours with me on
microscopic observations (SEM). The manuscript was partly translated into English by Sylwia
Ufnalska. I would also like to thank anonymous reviewers for helpful comments and valuable
suggestions on the manuscript.
Funding
This research was financially supported by the statutory funds of the Department of Ecology
and Environmental Protection, University of Szczecin.
Competing interests
No competing interests have been declared.
Wieczorek / The lichen genus Opegrapha s. l. in Poland
1. Introduction
Opegrapha s. l. is a species-rich genus of crustose lichens. Individuals are typically found
on the bark of various tree species or on rocks, decaying wood, and thalli of other lichens.
They are distributed in all climatic zones across the planet. Although it is difficult to assess
the number of species of this genus, the Index Fungorum contains about 600 species in the
genus, including all synonyms.
For most of the last century, a taxonomic concept was used for this genus in Europe, which
followed Redinger’s [1] monograph. However, individual species concepts in that monograph
did not deviate substantially from those presented previously by Acharius [2,3], Stizenberger
[4], or Nylander [5]. The widespread use of molecular methods resulted in a major change
for the systematics of this group. For example, Ertz et al. [6] drew attention to its polyphyly.
Moreover, molecular studies resulted in the introduction of a new family, the Lecanographaceae, which included the genera Alyxoria, Lecanographa, and Zwackhia [7,8]. In 2015,
several new genera, including Gyrographa and Pseudoschismatomma, were described in an
additional study by Ertz et al. [9].
In Poland, the genus Opegrapha has been studied in detail only by Nowak [10], whose
methods followed Redinger’s monograph [1]. In 1983, Nowak [10] recognized 35 species
and numerous intraspecific taxa on the basis of original observations. In 2003, Fałtynowicz
accepted 14 Opegrapha species (with three synonyms) in his lichen checklist of Poland [11].
In Poland, genus Opegrapha has recently been well recognized by Bielczyk [12] and Bielczyk
et al. [13] in the Carpathians.
The major objectives of this study were to: (i) determine the range of intraspecific variation
for selected morphological and anatomical characters using light microscopy (LM) and scanning electron microscopy (SEM); (ii) estimate taxonomic diversity of the genus Opegrapha
s. l. in Poland, with special emphasis on the O. vulgata group; (iii) construct a key for species
identification use in Poland; (iv) present distribution ranges for all species recorded from
Poland; and (v) analyze habitat preferences of individual species.
2. Research history of the genus
The genus Opegrapha first was described by Humboldt in 1793 [14] in the family Graphidaceae.
The original description contained three species, but these currently are regarded as synonyms
of Graphis scripta [15]. The first valid description of the genus Opegrapha was provided by
Acharius [16] in 1809. This generic name derived from his earlier work published in 1803
[17], where he first used the name Opegrapha vulgata (Ach.) Ach. The binomial name Lichen
vulgatus, from which he took this specific name, came from an earlier publication in 1798 [18].
On the basis of apothecium structure, Acharius [2] divided the genus Opegrapha into three
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
genera: Arthonia, Graphis, and Opegrapha. In total, the newly described genus Opegrapha
included 24 species [17], with five more species added in 1810 [2]. Santesson [15] suggested
that the name given by Acharius could be preserved, as it was also used by Massalongo [19],
Leighton [20], Körber [21,22], and Nylander [23,24]. Stizenberger’s [4] work, Über die Steinbewohnenden Opegrapha-Arten, took into account these earlier works and available materials
from contemporary researchers, and attempted to summarize the state of the taxonomy of
the group. In his treatment, Stizenberger included 15 species with numerous varieties and
forms. It is noteworthy that he was the first author to suggest the creation of the new family
Opegraphaceae [4]. However, his suggestion was not widely accepted by other botanists. In
1907, as a result of changes proposed by Zahlbruckner [25], the genus Opegrapha was placed
in the family Graphidaceae, a family that contained other systematically distinct genera,
including Graphis, Xylographa, and Encephalographa. That classification, with slight modifications, was accepted by most lichenologists [1,26]. Redinger [1] then presented a critical work
that provided the first detailed concept for the genus Opegrapha, which remained valid until
the 1970s. Redinger assigned 62 species to this genus that were divided into two sections:
Euopegrapha with 58 species, and Pleurothecium with four species.
For nearly 180 years, the genus Opegrapha was included in the family Graphidaceae. In
1970, Ozenda and Clauzade [27] argued that the family Opegraphaceae should be established, because of differences in the structure of the paraphyses. Important modifications
were proposed by Poelt [28], with his classification based primarily on the types of asci,
paraphyzes, and shapes of the ascospores, which allowed him to restore the family Opegraphaceae with the genus Opegrapha and placed in the order Arthoniales. In 1986, Eriksson
and Hawksworth [29] suggested that the families Opegraphaceae and Roccellaceae should
be moved to the order Opegraphales in the class Ascomycetes. The family Opegraphaceae,
as defined by Eriksoon and Hawksworth [29], was composed of 14 genera, including the
genus Opegrapha. Nearly 10 years later, Hawksworth et al. [30] modified the classification
of the genus Opegrapha, excluding it from the family Opegraphaceae. The genus Opegrapha
then was moved to the family Roccellaceae in the order Arthoniales, and this decision was
accepted by lichenologists [31–33].
In the late 1980s, Torrente and Egea [34] published results of their research on the family
Opegraphaceae in the Iberian Peninsula, and paid attention to the structure of some anatomical
parts that were not considered previously as diagnostic at the genus level within this group.
These primarily included ascus structure, with the apical apparatus and ascospore type. In
1998, Grube [33] reviewed the literature on the position and separation of genera within the
Arthoniales. Since then, the taxonomic concept for this group of lichens has been continuing to change. In the early twenty-first century, Ertz [35] published the first monograph
of the genus Opegrapha in the Palaeotropics. He characterized 45 species, including nine
new species. Among them, six species have a broad, pancontinental range and are found in
Europe as well.
Recently, Ertz et al. [6] used molecular sequence data to reveal that some genera of crustose
lichens, such as Enterographa and Opegrapha, were polyphyletic. Two species previously
belonging to the genus Opegrapha and occurring in Poland (O. calcarea and O. atra) were
thus placed in the family Arthoniaceae. However, Ertz and Tehler [7] once again restored
the family Opegraphaceae, and placed it in the order Arthoniales along with the families
Chrysothrixcaceae (with the genus Chrysothrix), Arthoniaceae (Arthonia, Arthothelium,
Cryptothecia, and Reichlingia), the newly created family Roccellographaceae (Dimidiographa,
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fulvophyton, and Roccellographa), and Roccellaceae (Chiodecton, Dendrographa, Dirina,
Enterographa, Erythrodecton, Lecanactis, Roccella, Roccellina, Schismatomma, Sigridea, and
Syncesia). In addition to the genus Opegrapha, the family Opegraphaceae also included the
genera Combea, Dictyographa, Dolichocarpus, Ingaderia, Paralecanographa, Paraingaderia,
Paraschismatomma, Pentagenella, Schizopelte, and Sparria. Moreover, the authors created a new
family, the Lecanographaceae, that contained the new genus Alyxoria (including among other
species classified earlier as Opegrapha culmigena, O. mougeotii, O. ochrolechia, O. varia; see
Diederich et al. [36]), the genus Zwackhia (with species classified earlier as Opegrapha viridis
and O. sorediifera and several others), and the numerously represented genus Lecanographa.
However, they noted that the systematic position of the family remained unclear.
In 2015, Ertz et al. [9] suggested that the new genera Gyrographa and Pseudoschismatomma
should be included with Dendrographa, Syncesia, Roccellina, Schismatomma, Lecanactis, Roccella,
Dirina, Sigridea, and Enterographa in the family Roccellaceae. Many of the species included
earlier in the family Roccellaceae are now part of the Opegraphaceae, Roccellographaceae,
and Lecanographaceae.
Tab. 1 summarizes the history of changes in systematic classification of the genus Opegrapha
s. l. representatives occurring in Poland.
3. Material and methods
This study is based on specimens from herbaria located in Poland (BDPA, KRA, KRAM,
KRAP, KTC, LBL, LOD, OLTC, POZ, TRN, UGDA, WA, WRSL) [37], the Institute of Biology,
University of Białystok, and abroad [KPABGI, OHHI, RSU (according to Index Herbariorum
Rossicum), BILAS, GSU, NNSU, and WI (according to Index Herbariorum)]. Materials from
private collections (herb. A. Flakus, herb. M. Kossowska, herb. M. Kukwa, herb. L. Lipnicki,
herb. E. Muchnik, herb. K. Szczepańska, herb. U. Schiefelbein, herb. M. Seaward), and
specimens collected during my field research from 1997 to 2014 were also included. My field
collections are housed in the Herbarium Stetinensis (SZUB). In total, nearly 1,400 specimens
(1,206 from Poland) were examined. The Polish material originated from all parts of the country.
Detailed data for each examined specimen were recorded. The analyzed morphological and
anatomical characters (Tab. 2, Fig. 1) were selected on the basis of the original observations
and published resources.
To analyze in detail some of the more subtle structures, such as paraphyzes, asci, and spores,
material from ascomata cross sections was squashed and stained in a drop of water containing
a solution of iodine in potassium iodide (IKI). To observe the color reaction of the walls and
apical structures of asci, 10% KOH and Lugol’s solution (KOH/IKI) were used. To observe
paraphyzes, 10% KOH alone was used. Lactophenol cotton blue allowed for visualization
of the apical nasse (the dome zone) in asci and ascospore cell walls. The color reaction in
thalli was examined using KOH (30% aqueous solution) (K), sodium hypochlorite (C), and
paraphenylenediamine (solution in 95% ethyl alcohol) (PD).
© The Author(s) 2018
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Tab. 1 History of systematic changes in the genus Opegrapha s. l. for the representatives that
occur in Poland.
Author
Family
Current genus name
Humboldt 1793
Graphidaceae
Opegrapha
Stizenberg 1862
Opegraphaceae
Opegrapha
Graphidaceae
Opegrapha
Poelt 1973
Opegraphaceae
Opegrapha
Eriksson and Hawksworth 1986
Zahlbruckner 1907
Opegraphaceae
Opegrapha
Hawksworth et al. 1995
Roccellaceae
Opegrapha
Hafellner 2009
Roccellaceae
Phacographa
Opegraphaceae
Opegrapha
Arthoniaceae
Arthonia
Lecanographaceae
Alyxoria, Zwackhia
Roccellaceae
Gyrographa,
Pseudoschismatomma
Ertz and Tehler 2011
Ertz et al. 2015
Tab. 2
Analyzed morphological and anatomical characters.
Thallus
Ascomata
Asci
Ascospores
Pycnidia
Conidia
White
Grey
White-grey
Brownish
Ascoma naked
Ascoma circular
Ascoma elliptic
Ascoma elongate
Ascoma unbranched
Ascoma branched
Ascoma convex
Ascoma sunken
Ascoma flat
ascoma length
Ascoma width
Disc convex (a) disc flat (b)
Disc concave (c)
Disc width
Excipulum type a
Excipulum type b
Excipulum type c
Excipulum type d
Excipulum type e
Excipulum continuous at
the base
Excipulum discontinuous
at the base
Excipulum edge thickness
Excipulum base thickness
Hypothecium height
Hymenium height
Pseudoepithecium height
Calcarea-type
Varia-type
Vulgata-type
Length
Width
Straight
Curved
Ellipsoid
Spindle-shaped
Needle-shaped
Central cell
enlarged
Cells uniform
Length
Width with
perispore
Number of septa
Perispore
thickness
Present (1)/
absent (0)
Sessile
Convex
Few (<10)
Numerous
(10–20)
Very numerous
(>20)
Black
Pruinose
Straight
Curved
Length
Width
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Fig. 1 Measurements of selected parts of the ascoma in Opegrapha s. l. (A) Ascoma cross
section: a – width; b – excipulum edge thickness; c – excipulum base thickness; d – disc
width; e – hypothecium height; f – hymenium height; g – pseudoepithecium height. (B)
Ascospore: a – length; b – width; c – perispore thickness. (C) Ascus: a – length; b – width.
Fig. 2 Excipulum types in Opegrapha s. l.
Fig. 3 Three grades of color reaction change for the inner
membrane and types of asci in Opegrapha s. l.
In this study, five types of excipulum were characterized, depending on their shape in
cross sections (see excipulum types in Fig. 2). In addition, I characterized three types of discs,
depending on their surface shape (Fig. 3): concave (A), flat (B), or convex (C). On the basis
of color reactions of the inner membrane, several types of asci were distinguished, following
Torrente and Egea [34].
The various structures of the ascomata were observed and measured under a light microscope (Nikon Eclipse E600 and Zeiss Stereo Lumar v12) (LM), and images were captured with
AioVision SE64Rel. 4.8 software. Additionally, a high-quality confocal laser scanning microscope (CLSM) and scanning electron microscope (Quanta 200 SEM) (SEM) were used.
Names of species used throughout the text are in accordance with valid taxonomic
classifications [7,9,36]. Nomenclature and author citations follow Index Fungorum. Only
basionyms and synonyms used by the authors of the collections are listed under the species.
Descriptions and discussions of species are based only on the material examined during the
course of this study.
Geographic elements were classified according to Wirth [38], Wirth et al. [39], Purvis et
al. [40], and Smith et al. [41].
Ecological characteristics of species of the genus Opegrapha s. l. are based on information
from labels of herbarium specimens and original field research.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
The geographic distribution of individual taxa in Poland was mapped on the ATPOL grid
square system [42]. All records marked on the maps were confirmed by relevant herbarium
materials. Localities of the studied specimens are listed alphabetically by the ATPOL grid
squares and named according to the regional division of Poland [43].
For each species, apart from the distribution map, the following data are given: (i) number
of localities (L); (ii) number of squares (10 × 10 km; S); (iii) diagram of the number of records
on particular substrate type, where the following abbreviations are used: Ab – Abies alba
(fir); Ac – Acer spp. (maple); Ae – Aesculus hippocastanum (horse chestnut); Al – Alnus spp.
(alder); Ap – Acer pseudoplatanus (sycamore); B – Betula spp. (birch); Cb – Carpinus betulus
(hornbeam); Co – Corylus avellana (hazel); Fa – Fagus sylvatica (beech); Fr – Fraxinus excelsior
(ash); l – limestone; Ma – Malus domestica (apple); Pa – Picea abies (spruce); Ps – Pinus spp.
(pine); Po – Populus spp. (poplar, etc.); Q – Quercus spp. (oak); r – rock; Rp – Robinia pseudacacia (false acacia); Sa – Salix spp. (willow, etc.); s – sandstone; So – Sorbus spp. (mountain
ash, etc.); T – Tilia spp. (linden); U – Ulmus spp. (elm); w – decaying wood.
4. Results and discussion
4.1. Numerical analysis of the examined material
On the basis of material from Poland obtained from 14 lichen herbaria and six private collections, 18 species were identified. According to the latest classifications [7,9,36], the species
belong to seven genera. Seven of these species are in the genus Opegrapha, two in the genus
Arthonia, four in the genus Alyxoria, two in the genus Zwackhia, and one species each in the
genera Gyrographa, Pseudoschismatomma, and Phacographa. This is the first time that the
species Zwackhia sorediifera has been reported from Poland. Two species represent nonlichenized lichenicolous fungi: Opegrapha geographicola and Phacographa glaucomaria. The largest
number of the studied specimens (322) originated from the Herbarium of Jan Kochanowski
at the University in Kielce, and slightly fewer (266) came from the lichen herbarium of the
W. Szafer Institute of Botany, Polish Academy of Sciences in Cracow (Fig. 4).
Among nearly 1,206 specimens studied from Poland, fewer than 7% were collected before
1945. In contrast, nearly 510 were collected between 1945 and 1980, and over 666 were collected since 1981 (Fig. 5).
The great majority of the examined specimens were from four species: Alyxoria varia
(334), Opegrapha vulgata (230), Zwackhia viridis (204), and Pseudoschismatomma rufescens
(180). Between 12 and 77 specimens were examined for six other species: O. niveoatra (77),
O. vermicellifera (52), Gyrographa gyrocarpa (47), Arthonia atra (34), O. dolomitica (24), and
Alyxoria mougeotii (12). The smallest number of specimens came from Alyxoria culmigena
and Zwackhia sorediifera (Fig. 6).
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Herb.Wieczorek
Herb. Szczepańska
Herb. Lipnicki
Herb. Kossowska
Herb. Flakus
Herb. Kukwa
WRS
WA
UGDA
TRN
POZ
KTC
LOD
LBL
KRAP
KRAM
KRA
HUMCS
GPN
BDPA
0
5
10
15
20
25
30
(%)
Fig. 4 Percent contribution of examined specimens from various herbaria.
Before 1946
1945–1980
After 1981
No data
0
100
200
300
400
Numbers of specimens
500
600
700
Fig. 5 Number of specimens collected before 1946, between 1945 and 1980, and since 1981.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Phacographa glaucomaria
Opegrapha saxicola
Opegrapha lithyrga
Opegrapha geographicola
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Alyxoria mougeotii
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha niveoatra
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
Alyxoria culmigena
0
50
100
150
200
250
300
Number of specimens
Fig. 6 Number of specimens of individual species of the genus Opegrapha s. l. in Poland.
4.2. Morphological features
4.2.1. Thallus
Members of the genus Opegrapha s. l. usually are uniformly crustose (Fig. 7) and are found
on the bark of various tree species, rocks, plant debris, and the thalli of other lichens. The
thallus of lichenized species in the genus Opegrapha s. l. is heterogeneous. Thalli of saxicolous
species in this genus may be epilithic (growing on the rock surface) or endolithic (penetrating
the rock, with a very thin outer layer), whereas epiphytic species always are epiphloeodal.
Thallus morphology varies widely. It can be uniform or composed of small aggregated nodules, sometimes powdery-granular. Areolate thallus (i.e., subdivided into segments, areoles)
usually is found near ascomata or in depressions of the substrate. In Gyrographa gyrocarpa, it
develops yellowish soralia that often are connected and form larger aggregations of soredia.
Sometimes, thalli are covered by delicate crevices or torn into pieces due to cracks in the
substrate, such as in a tree bark.
Thallus color is highly variable, consisting of various hues of gray, white, brown, and
green, which form lighter patches, clearly distinguished from the substrate, or only slightly
different from it (Fig. 8). In some species, thalli are bordered with a conspicuous prothallus
of various thickness.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 7 Thallus: (A) Opegrapha vermicellifera – thallus, SEM (leg.
W. Fałtynowicz, UGDA-L 2469); (B,C) O. dolomitica – endolithic thallus,
SEM (leg. J. Nowak, KRAM-L 12273); (D) O. vermicellifera – thallus
smooth, continuous, SEM (leg. A. Wieczorek, 1997, herb. Wieczorek);
(E) O. vulgata – thallus composed of small aggregated nodules, SEM (leg.
J. Nowak, KRAM-L 7289); (F) Arthonia atra – thallus crevices, LM (leg.
L. Lipnicki, 1989, herb. Lipnicki); (G) Gyrographa gyrocarpa – soredia,
SEM (leg. P. Czarnota, GPN 2145); (H) O. niveoatra – thallus divided
by bark cracks, LM (leg. S. Cieśliński & Z. Tobolewski, 1983, KTC); (I)
A. atra – thallus pale, LM (leg. A. Wieczorek, 2002, herb. Wieczorek);
(J) O. niveoatra – thallus dark, LM (leg. W. Fałtynowicz, UGDA-L 3435);
(K,L) Zwackhia viridis – prothallus, LM (leg. A. Wieczorek, 1998, herb.
Wieczorek). Scale bars: (A,B) 30 µm; (C,F,K,L) 1 mm; (D) 200 µm; (E)
0.5 µm; (G) 150 µm; (H) 1.5 mm; (I,J) 2 mm.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Opegrapha saxicola
Opegrapha lithyrga
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Alyxoria mougeotii
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha niveoatra
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
Alyxoria culmigena
0%
10%
20%
30%
White
40%
50%
60%
White-grey
70%
Brown
80%
90%
100%
Grey
Fig. 8 Variation in thallus color in selected species of the genus Opegrapha s. l. in Poland.
Fig. 9 Algae of the genus Trentepohlia in the thalli of Opegrapha s. l.: (A) short
filament in a hydrated thallus, SEM (leg. A. Wieczorek, 1998, herb. Wieczorek); (B)
single algal cells, SEM (leg. S. Cieśliński, 1990, KTC); (C) short filament, LM (leg.
A. Wieczorek, 1998, herb. Wieczorek). Scale bars: (A) 5 µm; (B) 4 µm; (C) 10 µm.
4.2.2. Algae
Thalli of lichenized species of the genus Opegrapha s. l. contain algae of the genus Trentepohlia
(Fig. 9). The algae form short filaments composed of several cells each that are linked together
loosely, particularly in the terminal parts. However, their shape can change depending on
water content of the thalli. The algal cells are orange because of a pigment that is a derivative
of β carotene and usually masks the green chlorophyll. Each algal cell contains one or several
chromatophores.
4.2.3. Ascomata
Species of the genus Opegrapha s. l. produce black ascomata classified as hysterothecia, varying in length from 0.1 mm to 2 mm. The ascomata do not (or very rarely) contain a slightly
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
whitish or yellowish pruinose. In most cases, ascomata are numerous, but rarely, thalli bear
only a few or no ascomata. In O. vermicellifera, ascomata are rare, but characteristic pycnidia
always are developed. An example of sterile, but easily identified species, is Gyrographa
gyrocarpa (due to the characteristic soralia, stained red with iodine). The other species are
unidentifiable when missing ascomata. The ascomata often are distributed evenly on the
thallus, but they may also form groups. They usually protrude above the thallus surface and
only rarely are they sessile or sunken.
The shape of the ascomata is highly variable and often differs within individual species.
They usually are circular, ellipsoid or elongate, more or less branched, rarely vermiform. They
are often fused. Only few species are characterized by low intraspecific variability in ascomata
shape, such as Arthonia calcarea. In the past, many taxonomists regarded the shape of the
ascomata as an important diagnostic feature. However, this feature is greatly modified under
the influence of changing environmental conditions and is of little importance for taxonomic
diagnosis in this genus (Fig. 10).
The ascomata disc is black, and rarely pruinose. Disc shape highly depends on the shape
of the ascomata: it may be narrow or wide, often depending on the ascomata’s maturity stage.
Usually well-developed discs are found in mature specimens, but they are sometimes difficult to distinguish in young specimens. The disc in the genus Opegrapha s. l. usually is the
widest in the central part of the ascomata and gradually tapers toward the apical parts. Most
species are concave or flat, very rarely convex, and the degree of convexity or concavity is
highly variable (Fig. 11). Disc’s area in SEM images is usually slightly granular. Its structure
is shaped by growing apically dilated paraphyzes.
The anatomical structure of the apothecia is an important diagnostic feature in the taxonomy of this genus. The ascomata of Opegrapha spp. are open (Fig. 12A) and composed of
four layers of various thickness: excipulum, hypothecium, hymenium, and pseudoepithecium
(Fig. 12B).
The excipulum (also known as the exciple) is the outermost layer of the ascomata (Fig. 13A).
In the genus Opegrapha s. l., the excipulum is mature throughout the period of the ascomata
development. The excipulum is brown-black to black; in some species, the color is continuous
at the base of ascoma (Fig. 13B), while in others it is discontinuous (Fig. 13C). The excipulum’s edge is well developed, the thickness of the edges in the upper part of the ascomata
varies and changes as the ascomata mature. In most cases during early ascomata maturation,
the excipulum forms strongly curved upward facing lips (Type A; Fig. 2). However, when
the ascomata grow, the excipulum is open (Types B, C, D, and E; Fig. 2). Most species of the
genus Opegrapha s. l. have the ascomata of Types B and C (Fig. 2), but in a few species (e.g.,
Alyxoria varia), widely opened apothecia develop (Types D and E) (Fig. 14). In these cases,
the edge and disc of the excipulum are black.
The hypothecium is the inner part of the ascomata that is located near the excipulum
(Fig. 15). It is formed by excipulum hyphae, which grow perpendicularly to the ascomata.
The hypothecium consists of lower parts of ascogenous hyphae. In species of the genus
Opegrapha s. l., this part of the ascomata is well defined because of the slightly darker color
of the adhering hymenium. Like hymenium, the hypothecium can be stained with iodine,
though sometimes the reaction is barely visible.
The hymenium is the part of the ascomata that contains asci and paraphyzes (Fig. 16A,B).
In species of the genus Opegrapha s. l., the hymenium is well developed, surrounded by the
excipulum, and adheres to the hypothecium. The hymenium is composed of crowded parallel
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 10 Morphology of ascomata in Opegrapha s. l., SEM: (A) circular,
convex, with a very narrow and concave disc (Arthonia atra, leg. Z. Tobolewski & S. Cieśliński, 1985, KTC); (B) circular, convex, with a wide and
concave disc (Alyxoria varia, leg. P. Czarnota, GPN 2875); (C) elongate,
convex, with a wide and flat disc (A. atra, leg. Z. Tobolewski, 1951, POZ);
(D) elliptic, convex, with a very narrow and concave disc (A. varia,
leg. P. Czarnota, GPN 2875); (E) branched, convex, with a narrow and
flat disc (Gyrographa gyrocarpa, leg. Z. Tobolewski, 1954, POZ); (F)
branched, convex, with a very narrow and convex disc (G. gyrocarpa,
leg. Z. Tobolewski, 1954, POZ); (G) elliptic, sessile, with a very narrow
and concave disc (Pseudoschismatomma rufescens, leg. Z. Tobolewski,
1955, POZ); (H) branched, convex, with a very narrow and concave
disc (Opegrapha vulgata, leg. J. Nowak, KRAM-L 13952); (I) young,
elliptic, convex, with a poorly developed, slightly concave disc (Alyxoria
mougeotii, leg. Z. Tobolewski, 1957, POZ); (J) circular, convex, with
a very wide and convex disc (Opegrapha dolomitica, leg. K. Toborowicz,
1975, KTC); (K,L) disc surface (A. varia, leg. P. Czarnota, GPN 2875).
Scale bars: (A,B) 50 µm; (C,E,G) 1 mm; (D) 0.5 mm; (F,H) 100 µm; (J)
200 µm; (K) 10 µm; (L) 4 µm.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Phacographa glaucomaria
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha saxicola
Opegrapha niveoatra
Opegrapha mougeotii
Opegrapha lithyrga
Opegrapha geographicola
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
Alyxoria culmigena
0%
10%
A - concave
20%
30%
B - flat
40%
50%
60%
70%
80%
90%
100%
C - convex
Fig. 11 Various types of discs in species of the genus Opegrapha s. l.
Fig. 12 Ascomata cross section: (A) Opegrapha vulgata, SEM (leg.
A. Wieczorek, 1997, herb. Wieczorek); (B) Pseudoschismatomma rufescens,
LM (leg. S. Cieśliński, 1990, KTC). Scale bars: (A) 20 µm; (B) 100 µm.
Fig. 13 Excipulum: (A) SEM image (Opegrapha vermicellifera, leg.
S. Cieśliński, 1992, KTC); (B) continuous at the base of ascoma, LM
(O. niveoatra, leg. W. Fałtynowicz, UGDA-L 3435); (C) discontinuous
at the base of ascoma, LM (Alyxoria varia, leg. L. Lipnicki, 1989, herb.
Lipnicki). Scale bars: (A) 20 µm; (B) 100 µm; (C) 50 µm.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Phacographa glaucomaria
Opegrapha saxicola
Opegrapha lithyrga
Opegrapha geographicola
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Alyxoria mougeotii
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha niveoatra
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
Alyxoria culmigena
0%
10%
20%
30%
A
40%
B
50%
C
60%
70%
D
80%
90%
100%
E
Fig. 14 Percent contribution of various excipulum types in individual species of
the genus Opegrapha s. l.
Fig. 15 Hypothecium: (A,B) Alyxoria varia, LM (leg. L. Lipnicki, 1989,
herb. Lipnicki); (C) ascogenous hyphae growing from the hypothecium,
SEM (Opegrapha vermicellifera, leg. S. Cieśliński, 1992, KTC). Scale bars:
(A,B) 100 µm; (C) 2 µm.
Fig. 16 Hymenium, SEM: (A) asci (Opegrapha vulgata, KRAM-L 14823);
(B) paraphyses (O. vulgata, KRAM-L 14823); (C) hyaline interhyphal
substance (O. vulgata, leg. D. Kubiak, 2001, OLTC). Scale bars: (A) 10 µm;
(B) 20 µm; (C) 3 µm.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
hyphae, which in the upper layer, near the excipulum, form lips curved toward the hymenium
center [34]. In species of the genus Opegrapha s. l., the hymenium is yellowish in color.
Paraphyzes are formed by sterile hyphae on the hypothecium. In the genus Opegrapha s. l.,
paraphyzes are persistent throughout the life span of the ascomata. At the first stages of the
ascomatal development, paraphyzes are connected to the lower part of the hypothecium. The
paraphyzes grow perpendicularly to the hypothecium, are crowded and are usually united with
asci by means of an interhyphal, hyaline substance (Fig. 16C). In mature ascomata, paraphyzes
are branched and distally clavate (club-shaped). Usually, they are branched above the asci.
The asci in the genus Opegrapha s. l. are cylindrical-clavate and clavate (Fig. 17A,B), usually
with a broader base. They are bitunicate, composed of an outer and inner membrane known
as the eoascus and endoascus, respectively (Fig. 17C). The eoascus is thin, covers the ascus,
and sometimes is amyloid and can be stained with KOH/IKI (Fig. 17D). The endoascus is
composed of two layers (inner and outer), which are distinguishable based on their different
sensitivity to double treatment with KOH/IKI. The inner layer can be stained blue more
intensely, while the outer layer remains unchanged or is stained only slightly. The staining
of the inner layer of the endoascus may be limited to the dome zone, be extended to half
the ascus height (Fig. 17E), or may extend to the whole length of the ascus. The variation in
staining and the structure of the apical apparatus are features that allowed Torrente and Egea
[34] to distinguish three types of ascus in the genus Opegrapha s. l. The first type, known as
the calcarea (Fig. 3), is characterized by a short stipe, with the inner layer of the endoascus
amyloid only in apical nasse, and a reduced ring with a large locular chamber. The variatype refers to clavate asci with the inner layer of the endoascus amyloid in the upper half
or slightly lower, with a short but wide chamber and no apical nasse. The vulgata-type has
cylindrical-clavate asci, with the inner layer of the endoascus amyloid nearly alongside its
whole length, with a small chamber and no apical nasse. At the apex of the ascus, the apical
apparatus is located (Fig. 17F,G).
Usually, the asci contain eight spores. During the course of spore growth, the outer membrane is damaged and the inner membrane expands until the ascus walls are destroyed and
the spores are released.
Ascospores in the genus Opegrapha s. l. are hyaline (besides pale brown in O. geographicola) and multicellular, with 4–15-septa. Their shape is variable, from elliptic to more or less
spindle-shaped (Fig. 18A–E), straight, or slightly curved. Ascospores vary in length from
11 µm to 61 µm, while they are 2 µm to 14 µm wide (including the outer covering, in the
widest part of the ascospore) in the hydrated state. Following dehydration, their width can
be reduced by up to 2 µm (Fig. 18F). The ascospores have one–two coverings: an inner one
known as the epispore, and the outer perispore, the presence or absence of the latter one is
an important diagnostic feature (Fig. 18G). The wall of the ascospore is hyaline, but after
long periods in a herbarium, it may turn light gray. The length and shape of the ascopores,
as well as the number of septa, have been used for species identification since Massalongo
[44]. Ascospores in the genus Opegrapha s. l. are autofluorescent [45] (Fig. 18H,I).
The pseudoepithecium is the upper most layer of the ascomata and is located above the
hymenium. This layer has been given various names. Korf [46] used the term pseudoepithecium to describe the undifferentiated material located above the hymenium. Muthappa
[47] believed that the epithecium is a layer composed of remnants of the hymenium, usually
consisting of damaged apical parts of paraphyzes. Eriksson [48] suggested using the term
pseudoepithecium for the layer present in all species having paraphyzes. Similarly, Hawksworth
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 17 Asci: (A) cylindrical-clavate, SEM (Zwackhia viridis, leg. A. Wieczorek,
1996, herb. Wieczorek); (B) clavate with mature ascospores, SEM (Pseudoschismatomma rufescens, leg. Z. Tobolewski, 1955, POZ); (C) two layers of the
endoascus, LM (Opegrapha vulgata, leg. J. Nowak, KRAM-L 16223); (D) amyloid
outer layer, LM (Arthonia calcarea, KRAM-L 26972); (E) outer layer of the
endoascus stained to half its length, LM (Alyxoria varia, leg. W. Fałtynowicz &
Z. Tobolewski, KRAM-L 34074); (F) apical apparatus, SEM (Arthonia calcarea,
KRAM-L 26972); (G) apical nasse, LM (A. calcarea, KRAM-L 26972). Scale
bars: (A) 30 µm; (B–E,G) 10 µm; (F) 5 µm.
[49] recommended the term pseudoepithecium, defining it as a shapeless granular covering,
which embeds the apical parts of paraphyzes. Considering the developmental forms of the
paraphyzes, and in light of the SEM observations in this study, it seems that there is no true
epithecium in the genus Opegrapha s. l. [34], and thus the use of the term pseudoepithecium
appears justified.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 18 Ascospores: (A) Opegrapha dolomitica, LM (leg. J. Nowak, KRAML 12349); (B) Alyxoria culmigena, SEM (leg. L. Lipnicki, 1989, herb. Lipnicki);
(C) Arthonia atra, SEM (leg. Z. Tobolewski, 1951, POZ); (D) Pseudoschismatomma rufescens, SEM (leg. S. Cieśliński, 1989, KTC); (E) Gyrographa
gyrocarpa, SEM (leg. J. Nowak, KRAM-L 17040); (F) dry, young ascospore
of A. atra, SEM (leg. P. Czarnota, GPN 231); (G) perispore and epispore of
O. dolomitica, LM (leg. J. Nowak, KRAM-L 12349); (H,I) autofluorescence
of O. dolomitica spores, CLSM (leg. J. Nowak, KRAM-L 12349). Scale bars:
(A,C,E,G) 10 µm; (B,D,F) 5 µm; (H) 20 µm; (I) 25 µm.
4.2.4. Pycnidia
The pycnidia in the genus Opegrapha are described in detail in my previous publication [50].
It is noteworthy that in species of this genus, the pycnidia are unilocular, i.e., characterized
by only by one locule [51] (Fig. 19A,B). The pycnidia are highly variable in shape and size,
even within species. Most of the analyzed specimens only had one ostiole (Fig. 19C–E),
although some pycnidia had two ostioles at the apex (Fig. 19F). The ostiole is nearly always
apical, but it also can develop along any part of the pycnidium. Ostioles usually are circular,
but sometimes they can be elongate or irregular. Ostiole size is also highly variable. The
pycnidia of the studied species are scattered on thalli or form clusters of various sizes. Most
of the analyzed species had prominent pycnidia (Fig. 19G–I, Fig. 20). For example, on one
specimen of a given species, some pycnidia were convex, while others were sunken in the
thallus. The pycnidia of the epilithic species occurred in more variable locations on the thallus
than those of species growing on the bark of trees.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 19 Pycnidia and conidia of the species of the genus Opegrapha s. l.: (A) O. vulgata,
SEM (leg. J. Nowak, KRAM-L 10343); (B) O. vermicellifera, LM (leg. W. Fałtynowicz, UGDAL 2785); (C) O. dolomitica, SEM (leg. J. Nowak, KRAM-L 29981; (D) O. saxicola, SEM (leg.
K. Szczepańska, 2004, herb. Szczepańska); (E) Pseudoschismatomma rufescens, SEM (leg.
Z. Tobolewski, 1953, POZ); (F) Alyxoria varia, SEM (leg. Z. Tobolewski, 1959, POZ); (G) O. vermicellifera, SEM (leg. S. Cieśliński, 1989, KTC); (H) O. niveoatra, SEM (leg. W. Fałtynowicz
& J. Miądlikowska, UGDA-L 3673); (I) Alyxoria varia, LM (leg. S. Cieśliński,1993, KTC); (J)
O. vulgata, SEM (leg. K. Czyżewska, LOD-L 520); (K) O. vermicellifera, SEM (leg. U. Bielczyk,
KRAM-L 44575); (L) O. vulgata, CLSM (leg. K. Glanc, KRAM-L 39101). Scale bars: (A)
500 µm; (B,C,E,I) 100 µm; (D,F–H) 50 µm; (J,L) 5 µm; (K) 2 µm.
The conidia produced by species of the genus Opegrapha s. l. are usually cylindrical or
clavate (Fig. 19J,K). The length of the conidia in the material that I analyzed varied between
species, even though length is one of the most important taxonomic characters. LM revealed
smooth and hyaline conidia, while under a scanning electron microscope, their delicate granular
ornamentation could be observed. Similar to ascospores, conidia in the genus Opegrapha s. l.
are autofluorescent [45] (Fig. 19L).
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Phacographa glaucomaria
Opegrapha saxicola
Opegrapha lithyrga
Opegrapha geographicola
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Alyxoria mougeotii
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha niveoatra
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
Alyxoria culmigena
0%
10%
20%
30%
Sessile
40%
50%
60%
70%
80%
90%
100%
Convex pycnidia
Fig. 20 Percentage of the contribution of sessile and convex pycnidia in the studied material.
4.3. Ecology
Species of the genus Opegrapha s. l. in Poland were shown to primarily colonize the bark of
various tree species (86% of records), but they also occur on rocks (10%), decaying wood (1%)
(Fig. 21), and sporadically on lichen thalli. The species composition of epiphytes and ephilithes
is presented in Fig. 22. Unfortunately, labels of the analyzed herbarium specimens did not
always contain complete information on habitat. Therefore, there generally was insufficient
information to estimate ecological requirements of individual species.
The analyzed taxa were recorded on the bark of 20 genera of trees. Most of them were
deciduous phorophytes, except for the conifers, such as Abies alba, Picea abies, and Pinus spp.
Representatives of Opegrapha s. l. are most frequently found on the smooth bark of Fagus
sylvatica and the cracked bark of Fraxinus excelsior, whilst they are found less frequently on
the bark of Carpinus betulus, Acer pseudoplatanus and other Acer spp., Alnus glutinosa, and
other phorophytes (Fig. 23).
The bark of Fagus sylvatica was most often colonized by Opegrapha niveoatra and O. vulgata,
while the bark of Fraxinus excelsior was colonized by Pseudoschismatomma rufescens, and
the bark of Carpinus betulus by O. niveoatra.
Nearly all of the epiphytic species are moderately acidophilic. They prefer slightly acid to
neutral sites, with a range of pH from 4.9 to 7.0 [38]. They grow on the bark of various tree
species, usually in open woodlands and on moderately moist sites, including oak-lindenhornbeam forest, acidophilus and eutrophic beech forests, acidophilus and thermophilous
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Rock; 114 (10%)
Other unidentified;
38 (3%)
Wood; 13 (1%)
Bark of various
trees; 1,041 (86%)
Fig. 21 Numeric and percentage of the contribution of ecological groups in the
genus Opegrapha s. l.
Opegrapha saxicola
Epilithic
Opegrapha lithyrga
Opegrapha dolomitica
Gyrographa gyrocarpa
Arthonia calcarea
Alyxoria mougeotii
Zwackhia viridis
Zwackhia sorediifera
Pseudoschismatomma rufescens
Epiphytic
Opegrapha vulgata
Opegrapha vermicellifera
Opegrapha niveoatra
Arthonia atra
Alyxoria varia
Alyxoria ochrocheila
(%)
Alyxoria culmigena
0
5
10
15
20
25
30
35
Fig. 22 Percentage contribution of specimens of epiphytic and epilithic species.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Ulmus spp.
Tilia spp.
Sorbus spp.
Salix spp.
Robinia pseudacacia
Quercus spp.
Populus spp.
Pinus spp.
Picea abies
Malus domestica
Fraxinus excelsior
Fagus sylvatica
Corylus avellana
Carpinus betulus
Betula spp.
Alnus spp.
Aesculus hippocastanum
Acer spp.
Acer pseudoplatanus
Abies alba
0
50
100
150
Number of specimens
200
250
Fig. 23 Number of specimens of epiphytic species of Opegrapha s. l. on various phorophytes.
oak forests, streamside alder-ash forest, and black alder bog forest. They are found in plant
communities of the Opegraphetum rufescentis Almb. 1948 ex Klem. 1955 and Opegraphetum
vermicelliferae Almborn 1948 [38] association.
The most frequently recorded epiphytic species from Poland include Alyxoria varia
(311 herbarium specimens), Opegrapha vulgata (226), and Pseudoschismatomma rufescens
(164). More research is needed on rare species (Alyxoria culmigena, O. niveoatra, or O. vermicellifera) to determine their distribution and ecological preferences.
All of the recorded epiphytic lichen species, except for Alyxoria culmigena and A. ochrocheila
(extremely rare species), are on the “Red list of threatened lichens in Poland” [52]. One of
them, O. vermicellifera, is regarded as an indicator of lowland primeval forests [53].
There are six epilithic lichens of the genus Opegrapha s. l. found in Poland: Alyxoria mougeotii,
Arthonia calcarea, Gyrographa gyrocarpa, Opegrapha dolomitica, O. lithyrga, and O. saxicola.
They occur primarily in uplands and mountains, and grow mostly on limestones, but occasionally also on sandstones and other rocks (Fig. 24), shaded and moist sites, or sunlit and
moderately dry sites. Species found on natural limestones are highly specific (e.g., A. calcarea,
O. dolomitica, and O. saxicola). In contrast, Gyrographa gyrocarpa prefers sandstones. The
other species, O. mougeotii and O. lithyrga, are as frequent on both limestone and sandstone.
As epilithic lichens, they usually are major components of epilithic moss-lichen communities.
For example, Wirth et al. [39] listed O. gyrocarpa and O. lithyrga as components of mountain
lichen grasslands of the alliance Enterographetum zonatae (Fegel 1939) Wirth 1972.
The most frequently recorded epilithic species are Gyrographa gyrocarpa (both sterile and
fertile specimens were noted) and Opegrapha dolomitica.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Sandstone
Rock (granite)
Limestone
0
10
20
30
40
50
60 (%)
Fig. 24 Percentage contribution of epilithic species on various rocks types.
Three of the epilithic lichen species of this group recorded from Poland (Arthonia calcarea,
Gyrographa gyrocarpa, and Opegrapha dolomitica) are threatened in Poland [52]. One of them,
Alyxoria mougeotii, is on the red list of the Beskid Sądecki Mountains [54].
4.4. Geographic distribution
Opegrapha s. l. is a genus with a wide, pancontinental distribution, as it is known from all
continents except Antarctica. Most of the species recorded from Poland are widespread in
the country.
Epilithic species are distributed mostly in the southern part of Poland (Fig. 25A). All of
them are typical mountain species, found in both the subalpine and montane zones. The
Fig. 25 Distribution of (A) epilithic and (B) epiphytic species of Opegrapha s. l. in Poland.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 26 The number of species of the genus Opegrapha s. l.
recorded in individual 10 × 10-km grids, which are based on
the ATPOL grid square system in Poland.
distribution of epiphytic species is associated with extensive natural woodlands. They were
recorded both in lowlands and in mountains (Fig. 25B).
The highest species diversity was recorded in southern Poland, in the West Beskids, Tatras,
and Bieszczady Mountains. In the north, high diversity occurs in the Pojezierze Mazurskie
(Masurian Lake District) and Nizina Północnopodlaska (North Podlachian Lowland)
(Fig. 26). The higher number of species in these regions results from the presence of the old,
well-preserved forests and more favorable ecological conditions [55,56], but this may be
a consequence of more intensive lichen surveys conducted in the area.
4.5. Key to the species of Opegrapha s. l. found in Poland
1
1*
2
2*
3
3*
4
4*
5
5*
6
Ascospores up to three-septate
Ascospores 4–15-septate
On rock
On tree bark, wood, plant debris
Growing directly on rock
Lichenicolous
Thallus C−, KC−
Thallus C+ reddish, KC+ reddish
Asci calcarea-type
Asci varia-type
Ascospores length <22 µm
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2
11
3
8
4
7
5
Gyrographa gyrocarpa (4.6.7)
Arthonia calcarea (4.6.6)
6
O. saxicola (4.6.12)
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6*
7
7*
8
8*
9
9*
10
10*
11
11*
12
12*
13
13*
16
16*
17
17*
18
18*
19
19*
Ascospores length >22 µm
O. dolomitica (4.6.8)
Hypothecium pale
O. geographicola (4.6.9)
Hypothecium dark, grey to grey-brown
Phacographa glaucomaria (4.6.15)
Asci calcarea-type
Arthonia atra (4.6.5)
Asci varia-type
9
Conidia 12–19 µm long, two–three-septate
Alyxoria ochrocheila (4.6.3)
Conidia 4–7 µm long
10
Ascospores 3–4 µm wide, perispore absent
Pseudoschismatomma rufescens (4.6.16)
Ascospores 4–7 µm wide, perispore present
Alyxoria culmigena (4.6.1)
On rock
12
On tree bark, wood, plant debris
13
Ascospore cells of similar size, asci vulgata-type
O. lithyrga (4.6.10)
Middle cell of ascospore enlarged, asci varia-type
Alyxoria mougeotii (4.6.2)
Ascospores four–six-septate
16
Ascospores 6–15-septate, fusiform
19
Middle cell of ascospore enlarged
Alyxoria varia (4.6.4)
Ascospore cells of similar size
17
Conidia straight, pycnidia whitish pruinose
O. vermicellifera (4.6.13)
Conidia straight or curved, pycnidia epruinose
18
Conidia 4–9 µm long
O. niveoatra (4.6.11)
Conidia 9–15 µm long
O. vulgata (4.6.14)
Perispore 1.5–2.0 µm thick
Zwackhia viridis (4.6.18)
Perispore 0.5–0.9 µm thick
Zwackhia sorediifera (4.6.17)
4.6. Species characteristics
4.6.1. Alyxoria culmigena (Lib.) Ertz
in Diederich, Ertz, Eichler, Cezanne, Boom, Fischer, Killmann, van den Broeck & Sérusiau, Bull. Soc. Nat.
luemb. 113: 105 (2012).
Opegrapha culmigena Lib., Pl. crypt. Arduenna, fasc. (Liège) 1: No. 15 (1830).
= Opegrapha betulina Sm., in Smith & Sowerby, English Botany. 32: Tab. 2281 (1811).
= Opegrapha herbarum Mont., in Guillemin, Arch. Bot. (Forlì) 2: 302 (1833).
Thallus thin, smooth or delicately wrinkled, grey-whitish, sometimes with yellow-green
tint. Ascomata numerous, solitary or in small groups, short or elongate, unbranched, rarely
branched, 0.3–2 × 0.1–0.3 mm. Discs narrow, concave, black, not pruinose. Excipulum thick,
elevated, permanent, black, dark brown, K−, continuous below the hypothecium, 35–45 µm
thick laterally and 50–90 µm at the base. Hypothecium 40–60 µm high, I+ slightly stained
blue. Hymenium 50–70 µm high, hyaline, I+ reddish. Pseudoepithecium up to 25 µm high,
I+ pale blue. Asci elongate, cylindrical or elongate-ellipsoid, 40–60 × 15–18 µm, varia-type.
Ascospores hyaline, spindle-shaped, with blunt ends, three-septate, 18–26 × (4)5–7 µm, with
perispore of 0.7−1.2 µm thick. Pycnidia scattered, sunken, visible as very small, blackish dots.
Conidia straight, 4–7 × 0.5–1 µm (Fig. 27). Chemistry: thalli K−, C−, PD−.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 27 Alyxoria culmigena (Lib.) Ertz: (A,B) thallus and ascomata, LM
(leg. A. Wieczorek, 1997, herb. Wieczorek); (C–F) thallus and ascomata, SEM
(leg. L. Lipnicki, 1989, herb. Lipnicki); (G) ascus with ascospores, LM (leg.
A. Wieczorek, 1997, herb. Wieczorek); (H) ascospore, SEM (leg. L. Lipnicki,
1989, herb. Lipnicki); (I,J) ascospores, LM (leg. A. Wieczorek, 1997, herb.
Wieczorek); (K) pycnidia, SEM (leg. A. Wieczorek, 1997, herb. Wieczorek);
(L) conidia, LM (leg. A. Wieczorek, 1997, herb. Wieczorek). Scale bars: (A)
1 mm; (B–D,K) 100 µm; (E,F) 250 µm; (G) 20 µm; (H,J) 10 µm; (L) 5 µm.
Discussion. According to Nowak [10], Alyxoria culmigena (= O. herbarum) is very closely
related to Arthonia atra, and sometimes they can be difficult to distinguish. My observations
show that Alyxoria culmigena has slightly longer and straighter ascospores and conidia. Nowak
[10] also reported that this species, in comparison to Arthonia atra, has wider ascospores, but
I could not confirm that report. Other similar of epiphytic taxa with three-septate ascospores
are Alyxoria ochrocheila and Pseudoschismatomma rufescens. Alyxoria culmigena differs from
A. ochrocheila by much shorter, unicellular conidia and a lack of pruina, and it differs from
P. rufescens by having wider ascospores and the presence of a perispore.
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Because of the small number of specimens, it is difficult to assess variability within this
species. Nowak [10] distinguished two varieties: var. herbarum known from France, Germany,
and Belgium, and var. betulina (Oliv.) J. Nowak reported from Poland. The latter is supposed
to have two forms: for. betulina and for. conferta (Erichs.) J. Nowak. The specimens that
I examined produced mostly unbranched (or occasionally branched) ascomata making it
difficult to distinguish among the different forms. The population from the Puszcza Barlinecka
(Barlinecka Forest) had some features of for. betulina that typically has unbranched ascomata,
but branched ascomata also were found on the thallus.
According to the latter cited authors, these features justify the creation of a separate genus.
Separating the genus Alyxoria also was confirmed by results of molecular studies [7] and in
my anatomical research.
Nomenclatural notes. This species is one of the first taxa recognized in the genus Opegrapha. It was described by Smith in 1811 [57] as O. betulina. Under the name O. herbarum, it
was reported by Montagne in 1833 [58]. Both names (O. betulina and O. herbarum) of this
species have been widely accepted by lichenologists [59,60]. The species was included in the
genus Opegrapha until recently. Currently, with O. ochrocheila, O. varia, O. subelevata Nyl.,
and O. bicolor R. C. Harris & Lendemer, it is placed in the genus Alyxoria. This generic name
was first used by Gray [61] to describe A. diaphora Gray and A. notha (Ach.) Gray, and it
was based on the generic name Alyxorina, which first was used by Acharius [2]. That name,
however, was not accepted, and contemporary lichenologists returned its initial generic
name, Opegrapha [62–64]. In all Alyxoria species, varia-type asci are produced, which are
characterized by a short, wide ocular chamber without an apical nasse, and staining to about
half of the inner layer of the endoascus [34]. According to the cited authors [34], these features
justify the creation of a separate genus. Separating the genes Alyxaria was also confirmed by
results of molecular studies [7] and in my anatomical research.
Distribution outside Europe. Species with a wide geographic range, found in Africa (Morocco: [65]), South and North America (Canada [66] and Alaska [67]), Asia (Korea [68] and
Taiwan [69]), and in Australia [70].
Habitat and distribution in Europe. Alyxoria culmigena is found on smooth bark of these
deciduous trees: Carpinus betulus, Fagus sylvatica, Acer spp., Fraxinus excelsior, and Quercus
spp. Occasionally in Belarus, it was recorded from the bark of Pinus spp., but rarely from
decaying wood or bark of tree trunks.
In Europe it is known from, e.g., Austria [71], Estonia [72], Ireland [73], Italy [74], Portugal
(Madeira [75]), and Slovenia [76].
Habitat and distribution in Poland. In Poland, it grows on the bark of Populus spp. and Fagus
sylvatica, where it is probably the rarest species of the genus Opegrapha s. l., first reported by
Ohlert [77] and Lettau [78]. It was later reported from the Pobrzeże Kaszubskie (Kashubian
Coastland) [79], Pobrzeże Szczecińskie (Szczecin Coastland) [80], Pogórze RożnowskoCiężkowickie (Rożnowsko-Ciężkowickie Foothills) [81], Pojezierze Ełckie (Ełk Lakeland)
[82], Pojezierze Kaszubskie (Kashubian Lake District) [83], Pomorze Gdańskie (Gdańsk
Pomerania) [84], Równina Drawska (Drawno Plain) [85], Równina Słupska (Słupsk Plain)
[86], and Wyżyna Lubelska (Lublin Upland) [87].
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 28 Distribution of Alyxoria culmigena in Poland.
Only two herbarium specimens were available to study from two localities in northwestern
Poland (Fig. 28). The records are published here for the first time.
4.6.2. Alyxoria mougeotii (A. Massal.) Ertz, Frisch & G. Thor
in Frisch, Thor, Ertz & Grube, Taxon 63(4): 727–744 (2014).
Opegrapha mougeotii A. Massal., Memor. Lich.: 103 (1853).
Thallus very thin, poorly developed, continuous or cracked, granulose, whitish, sometimes
with grey-brown tint. Ascomata numerous, solitary or in small groups, unbranched or
branched, 0.5–2.5 × 0.2–0.4 mm. Discs narrow, very rarely slightly widened, concave or flat,
black, not pruinose. Excipulum well defined, permanent, black, naked, dull or sparsely dotted
with granules of thallus, continuous below the hypothecium, 30–50 µm thick laterally and
40–70 µm at the base. Hypothecium 20–40 µm high. Hymenium 45–70 µm high, I+ reddish.
Pseudoepithecium 1.2–1.5 µm high. Asci club-shaped, mature 55–70 × 15–20 µm, young
40–55 × 13–15 µm, inner layer of endoascus amyloid in upper part, ring visible around
a small ocular chamber (varia-type). Ascospores hyaline, spindle-shaped, straight or slightly
curved, five–eight-septate, 19–30 × 4–8 µm, with enlarged middle cell, pointed distal cells,
and with perispore of 0.8–2 µm thick. Pycnidia sunken in thallus, conidia straight, 4–6 ×
1 µm (Fig. 29). Chemistry: thalli K−, C−, PD−.
Discussion. Alyxoria mougeotii sometimes resembles O. saxicola in the ascomata shape
and thallus habit, but these two species differ in ascospore size and structure. The major
distinguishing feature between the species is that A. mougeotii has an enlarged middle cell
in the ascospore.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 29 Alyxoria mougeotii (A. Massal.) Ertz, Frisch & G. Thor: (A,B)
thallus and ascomata, LM (leg. Z. Tobolewski, 1954, POZ); (C–F)
thallus and ascomata, SEM [(C,D) leg. J. Nowak, KRAM-L 4132; (E,F)
leg. J. Nowak, KRAM-L 4128)]; (G) ascus with ascospores, LM (leg.
A. Wieczorek, 1997, herb. Wieczorek); (H,I) ascospores, LM (leg.
A. Wieczorek, 1985, herb. Wieczorek); (J,K) pycnidium, SEM (leg.
Z. Tobolewski, 1954 Aug 6, POZ); (L) conidia, LM (leg. Z. Tobolewski,
1954, POZ). Scale bars: (A) 200 µm; (B–D,F,K) 100 µm; (E,J) 50 µm;
(G–I) 10 µm; (L) 5 µm.
Alyxoria mougeotii has highly variable sizes and shapes of the ascomata, depending on
environmental conditions. Their ascomata usually are ellipsoid [var. lutescens (Ach.) Redgr.],
rarely circular, or strongly elongate and branched (var. mougeotii) [10]. In most of the examined
ascomata, the disc was narrow and flat or slightly convex. They generally had an excipulum
of type B, but only occasionally type C. The white thallus was sessile, and convex pycnidia
were observed (Fig. 30).
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
100
C
D
90
C
80
B
C
B
C
B
70
60
%
B
50
40
A
30
B
A
A
B
A
20
10
0
A
1
2
3
4
5
6
7
Fig. 30 Variation in selected morphological and anatomical features of Alxyoria mougeotii. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata: A – unbranched,
B – branched; 3 – disc type: A – concave, B – flat, C – convex; 4 – thallus color: A – white,
B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia:
A – few, B – numerous, C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
Distribution outside Europe. Alxyoria mougeotii is known only from a few localities outside
of Europe, including Morocco [65] and Tunisia [88] in North Africa.
Habitat and distribution in Europe. Epilithic types are found typically growing on limestone, rarely on sandstone, and in shaded and moist sites. Its distribution has been studied
insufficiently. Reported from Austria [71], Estonia [89], Germany [90,91], Great Britain [92],
Hungary [93], Ireland [94], Malta [95], the Netherlands [96], Norway [97], Portugal (Madeira,
Azores [75,98]), Spain [99], Sweden [97], and Ukraine [100].
Habitat and distribution in Poland. Like other parts of Europe, in Poland Alxyoria mougeotii
grows on limestone, but less often on sandstone or slate. It grows primarily on vertical or
hanging sites that dry and shaded, and occasionally in slightly moist sites. Sometimes, it can
be found in partly sunny sites.
Its first records were from the Wąwóz Szopczański [Sobczański] (Szopczański Gorge)
(in the Pieniny Mountains) [101]. Later, it was reported from several other localities in the
southern part of the country, including the Beskid Sądecki mountain range [102,103], Dolina
Dolnego Sanu (Lower San River Valley) [104], Świętokrzyskie Mountains [105], Łańcuch
Tatrzański macroregion [106,107], Niecka Włoszczowska (Włoszczowa Basin) [108], Pagóry
Jaworznickie (Jaworzno Hills) [109], Pieniny Mountains [110,111], and Pogórze Przemyskie
(Przemyśl Foothills) [112].
The species is rare in Poland (Fig. 31), as it is known only from 13 localities in the Central Beskids or a few more localities in the Wyżyna Krakowsko-Częstochowska (KrakówCzęstochowa Upland), Wyżyna Kielecka (Kielce Upland), and Pogórze Przemyskie.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 31 Distribution and substrate preference of Alyxoria mougeotii in Poland.
4.6.3. Alyxoria ochrocheila (Nyl.) Ertz & Tehler
Fungal Diversity 49(1): 50 (2011).
Opegrapha ochrocheila Nyl., Flora (Regensburg) 48: 212 (1865).
= Opegrapha rubescens Sandst., Abh. Naturw. Ver. Bremen 21: 50 (1919).
Thallus thin, continuous, white or grey, covered by orange pruina. Ascomata numerous, solitary
or in groups, unbranched or branched, black, 0.5–2.0 × 0.2–0.4 mm. Discs narrow, sometimes
slightly wider in central part, pruinose with rusty-orange pruina. Excipulum relatively thick,
elevated, black, pruinose similarly to disc, continuous below the hypothecium, 20–30 µm thick
laterally and 40–50 µm at the base. Hypothecium pale brown, 20–35 µm high. Hymenium
50–65 µm high, I+ partly blue-green. Pseudoepithecium poorly developed, reddish-brown,
10–15 µm high. Asci club-shaped, 45–60 × 13–15 µm, inner layer of endoascus amyloid in
upper part (varia-type). Ascospores hyaline, ellipsoid, three-septate, 14–18 × 4–5 µm, with very
narrow, hardly visible perispore of 0.4–0.7 µm thick. Pycnidia numerous, small, black. Conidia
straight, two−three-septate, 12–19 × 1–2 µm (Fig. 32). Chemistry: thalli K−, C− PD−.
Discussion. The major microscopic feature that unambiguously distinguishes A. ochrocheila
from other epiphytic species are the two−three-septate conidia that are 12–19 µm long. Some
specimens that did not produce conidia were reported in the literature [35], but in Poland
no such materials were observed. Alyxoria ochrocheila is easy to distinguish with the use of
a magnifying glass due to the characteristic rusty-orange pruina on the ascomata. Pentecost
and James [59] and Ertz [35] described specimens with a nonpruinose ascomata, but all of the
specimens from Poland had scattered pruinose ascomata. With respect to asci and ascospores,
A. ochrocheila has slightly wider ascospores than Pseudoschismatomma rufescens, shorter
ascospores than Alyxoria culmigena, and when compared to Arthonia atra, A. ochrocheila has
no amyloid layer in the ascus and a short chamber without apical nasse.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 32 Alyxoria ochrocheila (Nyl.) Ertz & Tehler: (A,B) thallus and apothecia, LM;
(C–E) apothecia, SEM; (F) ascus with mature spores, LM; (G) ascospore, LM; (H)
pycnidia, LM; (I) conidium, LM (leg. J. Kiszka & J. Nowak, KRAM-L 17736). Scale
bars: (A,B) 1 mm; (C,D) 100 µm; (E) 50 µm; (F,G) 10 µm; (H) 500 µm; (I) 20 µm.
The five voucher specimens examined were insufficient to assess intraspecific variation.
However, the studied specimens produced mostly solitary, unbranched ascomata, with narrow,
exclusively concave discs that were widened sporadically in the central region. Few ascomata
were branched. All specimens had relatively numerous, convex pycnidia.
Nomenclatural notes. The species was described in 1865 by Nylander [113], who gave it the
binomial Opegrapha ochrocheila. Leighton [114] considered it as a form of O. atra, which was
accepted by Bouly de Lesdain [115] and Smith [116]. However, this treatment was rejected
by other researchers, who treated it as a separate species [1,34,38,117–119].
Distribution outside Europe. It is known from North America (e.g., from Canada [66]) and
Asia (Armenia [120] and Yemen [121]).
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], A. ochrocheila occurs in Central Europe, Mediterranean, and sub-Atlantic regions. It usually grows
on the smooth bark of Fagus sylvatica and Carpinus betulus and usually is found in moist
forest communities where the substrate is moderately acidic to neutral: pH 5.7–7.0 [38].
Known from Europe, e.g., from Austria [71], Denmark [122], Estonia [123], Germany
[90,124–126], Great Britain [92], Greece [127], Ireland [94], Italy [74], the Netherlands
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 33 Distribution and substrate preference of Alyxoria ochrocheila in Poland.
[96], Norway [97,128], Portugal (Azores [98]), Slovenia [76], Spain [99], Sweden [128], and
Switzerland [129].
Habitat and distribution in Poland. Two of the analyzed specimens grew on the bark of
Fagus sylvatica, while the third one grew on Carpinus betulus. The species was first reported
by Migula [130] in northern Poland, and it also was found in the Beskid Sądecki Mountains
[103], Beskid Żywiecki Mountains [131], Dolny Śląsk (Lower Silesia Province) [132], Pieniny Mountains [110], Równina Kozienicka (Kozienice Plain) [133], Wzgórza Łopuszańskie
(Łopuszańskie Hills) [134].
Based on the available herbarium material, the species occurs in central and southern
Poland (Fig. 33).
4.6.4. Alyxoria varia (Pers.) Ertz & Tehler
Fungal Diversity 49(1): 53 (2011).
Opegrapha varia Pers., Ann. Bot. (Usteri) 1: 30 (1794).
= Opegrapha diaphora Ach., Meth. Lich.: 19 (1803).
= Opegrapha lichenoides Pers., Ann. Bot. (Usteri) 7: 30 (1794).
= Opegrapha pulicaris (Hoffm.) Schrad., in Acharius, Lichenogr. Univ.: 251 (1810).
= Opegrapha variiformis Anzi Comm. Soc. crittog. Ital. 1(fasc. 3): 160 (1862).
Thallus thin, smooth or cracked, from white to grey, white-grey and brownish. Ascomata
numerous, variable in size and shape, solitary or in small groups, unbranched or branched,
0.3–2.0 × 0.2–0.8 mm. Discs concave, flat or convex, wide, in younger specimens sometimes
narrow, usually not, but sometimes pruinose with whitish or yellow pruina. Excipulum black,
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Fig. 34 Alyxoria varia (Pers.) Ertz & Tehler: (A) thallus and ascomata, LM
(leg. J. Nowak, KRAM-L 14551); (B–F) thallus and ascomata, SEM [(B,D)
leg. S. Cieśliński, 1993, KTC; (C) leg. J. Nowak, KRAM-L 14551; (E,F) leg.
S. Cieśliński, 1988, KTC]; (G) ascus with ascospores, LM (leg. J. Nowak,
KRAM-L 14551); (H) ascus, SEM (leg. S. Cieśliński, 1990, KTC); (I) ascospores,
SEM (leg. K. Toborowicz, 1976, KTC); (J,K) ascospores, LM (leg. S. Cieśliński,
1990, KTC); (L) pycnidium, SEM (leg. K. Glanc, KRAM-L 39086). Scale bars:
(A) 500 µm; (B–G) 100 µm; (H–J) 10 µm; (K) 20 µm; (L) 50 µm.
well defined, permanent, edge thick, lips strongly curved towards hymenium, continuous, rarely
discontinuous below the hypothecium, 20–45 µm thick laterally and 30–60 µm at the base.
Hypothecium 15–50 µm high, K−. Hymenium hyaline or pale yellowish, 40–80(90) µm high,
I+ red-yellow. Pseudoepithecium grey-brown, 10–25 µm high, K−. Asci club-shaped, 30–70 ×
12–18(20) µm, inner layer of endoascus amyloid to half of its length (varia-type). Ascospores
hyaline, spindle-shaped, straight, four–six-septate, 15–30 × 4–7(8) µm, with enlarged middle
cell, and with visible perispore of 0.6–1.5 µm thick. Pycnidia scattered, sunken. Conidia
straight or slightly curved, 3–4 × 1–1.5 µm (Fig. 34). Chemistry: thalli K−, C−, PD−.
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Discussion. Alyxoria varia is easy to distinguish from other epiphytic species. Its diagnostic
features are: the enlarged middle cell of ascospores, and the amyloid upper half of the inner
layer of the endoascus.
Alyxoria varia is highly variable with respect to thallus color, morphology, and anatomy of
the ascomata. Individual populations in Poland and other parts of Europe differ remarkably,
not only in thallus color and size and shape of ascomata, but also in ascospore size. These
features resulted in a taxonomic description of many varieties and forms of this species,
including O. lichenoides for. populina (Mong.) A. Zahlbr., for. confluens (Massal.) A. Zahlbr.,
for. cerebrina (Erichs.) J. Nowak, for. juglandis (Massal.) A. Zahlbr., var. nigrocaesia Chev., var.
chlorina (Pers.) Redgr., O. pulicaris for. lignicola (Harm.) A. Zahlb., for. minuta (Chev.) Oliv.,
for. incrustata (God.) Oliv., for. pollinii (Massal.) Redgr., for. phaea (Ach.) Oliv., O. diaphora
var. diaphora, for. signata (Ach.) J. Nowak, for. tigrine (Ach.) J. Nowak, for. saprophila (Nyl.)
Oliv., var. tridens (Ach.) Oliv., var. spurcata (Ach.) Ach., for. subgregaria Harm, O. rimalis
Pers. [10]. Its most frequent phenotype in Poland produces numerous circular to elliptic
ascomata, with a conspicuous disc and thin excipulum edge. The dics shapes also are variable.
Apothecia with flat, convex, and concave discs were observed. Moreover, in the examined
material, convex discs were found to have wide and narrow ascomata. The number and types
of pycnidia (Fig. 35) also were extremely variable. Similarly, the shape, length, and number
of septa vary widely, but their variation is continuous and fits within the range of variability
circumscribed for this species. Thus, attempts to identify a relationship between particular
phenotypes and distinct taxa at lower ranks were unsuccessful.
Nomenclatural notes. The extreme morphological and anatomical variation of Alyxoria varia
is due to its extensive geographic range. The variability resulted in a diverse circumscription of
the species throughout its history. Considering its complicated taxonomic history, as well as the
large numbers of varieties and forms of this species, it is challenging to identify correctly.
Difficulties with its classification probably result from different methods of taxonomic
analysis and concepts used by different researchers. The taxon was regarded for a long time
as a species aggregate, which could be divided into Opegrapha lichenoides (var. nigrocaesia,
and chlorina), O. pulicaris, O. rimalis, O. subrimalis, and O. diaphora (var. diaphora, tridens,
spurcata, and angustata Bouly de Lesd. [1]). For three nomenclatural synonyms (O. lichenoides,
O. pulicaris, and O. diaphora), Nowak [10] compiled a list of as many as 22 heterotypic names
of intraspecific taxa. Hawksworth et al. [62] excluded O. pulicaris from this group, as it was
considered an allied fungus growing on lichens. Pentecost and Coppins [119] included only
two names in the aggregate species: O. lichenoides and O. varia (synonyms: O. diaphora and
O. rimalis). Clauzade and Roux [64] maintained that O. varia was a collective species. Torrente and Egea [34] attempted to clarify the classification of this group, but failed to find any
relationships that determined limits of the taxonomic variation. Because of the discrepancies
in systematic classification of this species, I used the concept of Santesson [63], such that all
examined material was regarded as A. varia.
Distribution outside Europe. Widespread in North America [135–139]. Occasionally recorded
from South America, Africa, and Asia. Reported from, e.g., Algeria [140], Argentina [141], the
Bahamas [142], Brasil [143], Canada [66], China [144], Costa Rica [145], Hong Kong [146],
India [147], Israel [148], Japan [149], Maldives [150], Morocco [65], in eastern Russia [151],
Syria [152], Tunisia [88], Turkey [153–155], Uruguay [156,157], and the USA [135–139].
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100
E
C
90
C
C
D
D
B
80
70
%
60
B
C
B
C
B
B
50
40
B
A
30
20
A
B
A
0
A
A
10
A
A
1
2
3
4
5
6
7
Fig. 35 Variation in selected morphological and anatomical features Alyxoria varia.
1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 ascomata: A – unbranched,
B – branched; 3 – disc type: A – concave, B – flat, C – convex; 4 – thallus color: A – white,
B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia:
A – few, B – numerous, C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], A. varia
has a Central European-Mediterranean and Atlantic distribution. In most European countries,
it is usually found on the bark of deciduous trees, including Carpinus betulus, Acer spp., Fagus
sylvatica, and Fraxinus excelsior. It was the least recorded from the bark of Quercus spp, and
occasionally it was found on conifers. Like most of the species of the group, Alyxoria varia
prefers moderately acidic to neutral substrates: pH 4.9–7.0 [38,39].
Widely distributed in Europe, reported, e.g., from Albania [158,159], Austria [71,160],
Bulgaria [161], Cyprus [162], Denmark [163], Estonia [123,164], Finland [165], France
[166,167], Germany [168–170], Greece [127], Hungary [93], Italy [171], Latvia [172], Lithuania
[173–175], Malta [95], Montenegro [176], Norway [97], Portugal (Madeira [75,177]), Romania
[178,179], Slovenia [76], Spain [99], Switzerland [129], and Ukraine [100,180,181].
Habitat and distribution in Poland. Alyxoria varia grows on the bark of various deciduous
tree species in Poland, including old, well-preserved forests and moderately sunny sites. Most
frequently, it can be found on Fagus sylvatica, Acer spp., Fraxinus excelsior, Carpinus betulus,
and Quercus spp., but it is less frequently found on Acer pseudoplatanus, Ulmus spp., Betula
spp., Tilia spp., and Populus spp. It is found also along streams and rivers (Salix spp.) and
on roadside trees (Acer spp., Aesculus hippocastanum, etc.). Occasionally, it can be found on
decaying wood, the bark of coniferous trees, and in sandstone.
In the nineteenth century, Alyxoria varia was described by Körber [21] and Stein [182],
although they did not provide detailed localities. It also was reported without localities by
Ohlert [77] and Lettau [78] from Prussia and Galicia [183–185]. During the last 100 years,
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there have been a large number of reports of A. varia published in the Polish lichenological
literature, but many of them are undocumented in the Polish herbaria.
The species was reported from: the Beskid Makowski Mountains [186], Beskid Mały
Mountains [187], Beskid Niski Mountains [188], Beskid Śląski Mountains [189,190], Beskidy
Lesiste Mountains [191], Bieszczady Mountains [192,193], Bory Tucholskie (Tuchola Forest)
[194,195], Dolina Dolnego Sanu (Lower San River Valley) [104], Dolina Konińska (Konin
Valley) [196], Dolny Śląsk (Lower Silesia Province) [132], Gorce Mountains [197], Stołowe
Mountains [198], Świętokrzyskie Mountains [105,199–201], Karkonosze Mountains [202],
Carpathians [203,204], Kotlina Biebrzańska (Biebrza Basin) [205], Kotlina Toruńska (Toruń
Basin) [206], Kotlina Żywiecka (Żywiec Basin) [207], Łańcuch Tatrzański macroregion
[107,208–211], Małopolski Przełom Wisły (Lesser Poland Gorge of the Vistula) [212], Masyw
Ślęży (Ślęża Massif) [213,214], Mierzeja Wiślana (Vistula Spit) [215], Niecka Włoszczowska
(Włoszczowa Basin) [216], Nizina Mazowiecka (Mazovian Lowland) [217], northeastern
Poland [218], Pieniny Mountains [111,219], Płaskowyż Nałęczowski (Nałęczów Plateau)
[220], Pobrzeże Kaszubskie (Kashubian Coastland) [79,221], Pobrzeże Szczecińskie (Szczecin
Coastland) [222], Podgórze Rzeszowskie (Rzeszów Foothills) [223], Podhale [224], Pogórze
Rożnowsko-Ciężkowickie (Rożnowsko-Ciężkowickie Foothills) [81], Pogórze Spiskie (Spisz
Foothills) [225], Pojezierze Choszczeńskie (Choszczno Lake District) [226], Pojezierze
Gnieźnieńskie (Gniezno Lake District) [227–229], Pojezierze Iławskie (Iława Lake District)
[230,231], Pojezierze Kaszubskie (Kashubian Lake District) [232], Pojezierze Krajeńskie
(Krajna Lake District) [233], Pojezierze Mrągowskie (Mrągowo Lake District) [234], Pojezierze
Olsztyńskie (Olsztyn Lake District) [235,236], Pradolina Wieprza (Wieprz River Urstromtal)
[237], Pradolina Wrocławska (Wrocław Urstromtal) [238], Równina Augustowska (Augustów
Plain) [239–242], Równina Bielska (Bielsk Plain) [55,243], Równina Drawska (Drawno Plain)
[244], Równina Kozienicka (Kozienice Plain) [245,246], Równina Łęczyńsko-Włodawska
(Łęczna-Włodawa Plain) [247,248], Równina Łowicko-Błońska (Łowicz-Błonie Plain)
[249], Równina Łukowska (Łuków Plain) [250,251], Równina Piotrkowska (Piotrków Plain)
[252,253], Równina Słupska (Słupsk Plain) [86], Roztocze [254–258], Sudetes [259–261], Śląsk
Opolski (Opole Silesia) [262], West Beskids [131,190,191], Western Pomerania [56,263,264],
Wybrzeże Słowińskie (Slovincian Coast) [265,266], Wysoczyzna Bełchatowska (Bełchatów
Highland) [267,268], Wysoczyzna Białostocka (Białystok Highland) [269–272], Wysoczyzna Elbląska (Elbląg Highland) [273], Wysoczyzna Kaliska (Kalisz Highland) [274,275],
Wysoczyzna Łaska (Łask Highland) [276], Wysoczyzna Siedlecka (Siedlce Highland) [277],
Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa Upland) [278], Wyżyna Sandomierska (Sandomierz Upland) [279], Wyżyna Wieluńska (Wieluń Upland) [280],Wzgórza
Radomszczańskie (Radomsko Hills) [216], Wzniesienia Mławskie (Mława Hills) [281], and
Zaklęsłość Sosnowicka (Sosnowica Depression) [282].
Throughout Poland, 334 specimens were located in the lowlands and mountains (Fig. 36).
A majority of its known localities are in the West Carpathians and East Beskids. It has been
reported from the Western Tatras by Tobolewski at 1,150 m in altitude [209]. In the Sudetes,
it is known from a few localities. Outside the mountainous regions, it is frequent in the East
Baltic-Belorussian Lowlands on phorophytes in extensive, dense woodlands. In northern
Poland, it was scattered in the Pojezierze Południowobałtyckie (South Baltic Lakeland) and
Pobrzeże Południowobałtyckie (South Baltic Coastland). In other parts of the country, it was
observed more often in the eastern Nizina Środkowopolska (Central Poland Lowland). In
the western part of the country, it was recorded only occasionally.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 36 Distribution and substrate preference of Alyxoria varia in Poland.
4.6.5. Arthonia atra (Pers.) A. Schneid.
Guide Study Lich.: 131 (1898).
Opegrapha atra Pers., Ann. Bot. (Usteri) 7: 30 (1794).
Thallus thin, external, smooth or cracked, ranging in color from white and white-grey to
brown. Ascomata numerous, solitary or clustered, unbranched or branched, sometimes
forming large and continuous patches, fused terminally, 0.1–0.8 mm long and 0.1–0.3 mm
wide. Discs concave (groove-like), flat or, more often, convex, central part rarely ± wider
(trough-like), black, not pruinose, 13–133 µm wide. Excipulum relatively thick, elevated,
permanent, black, naked, continuous below the hypothecium, rarely discontinuous, 20–40 µm
thick laterally and 30–60 µm at the base. Hypothecium 12–25(30) µm high, K−. Hymenium
hyaline, 30–60 µm high, I+ blue. Pseudoepithecium brownish, 6–14 µm high, K−. Asci 30–50
× 10–15 µm, calcarea-type. Ascospores hyaline, ellipsoid, three-sepate, 13–17(20) × 4–6 µm,
with perispore of 0.4–0.6 µm thick. Pycnidia frequent, black, usually convex, 17–68 µm in
diameter. Conidia straight or curved, 3.5–5 × 0.5–1.5 µm (Fig. 37). Chemistry: thalli K−,
C−, PD−.
Discussion. Arthonia atra is characterized by short ascospores (up to 17 µm), a thin perispore
(0.4–0.6 µm), calcarea-type asci, and a short conidium (3.5–5 µm). According to Nowak [10],
this species is most similar to Alyxoria culmigena (= O. herbarum) and Pseudoschismatomma
rufescens (= O. rufescens). However, its ascospores are shorter than in A. culmigena, slightly
wider than in P. rufescens, and have a conspicuous but thin perispore. Moreover, in contrast to
the vulgata-type asci in P. rufescens and the varia-type asci in A. culmigena, A. atra produces
calcarea-type asci. Another species that can be confused with A. atra is Alyxoria ochrocheila.
Both species have ascospores of similar size and with perispores, but A. ochrocheila has conidia
that are much longer and two−three-septate.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 37 Arthonia atra (Pers.) A. Schneid.: (A) thallus and ascomata, LM (leg. S. Cieśliński,
1993, KTC); (B–F) thallus and ascomata, SEM [(B,D,E) leg. S. Cieśliński, 1985, KTC; (C)
leg. S. Cieśliński, 1993, KTC]; (G) ascus with ascospores, LM (leg. A. Wieczorek, 2002,
herb. Wieczorek); (H) ascospore, SEM (leg. S. Cieśliński, 1985, KTC); (I,J) ascospores, LM
(leg. A. Wieczorek, 2002, herb. Wieczorek); (K) pycnidium, SEM (leg. Z. Tobolewski, 1951,
POZ); (L) conidia, LM (leg. A. Wieczorek, 1996, herb. Wieczorek). Scale bars: (A) 2 mm;
(B,C,E,F) 100 µm; (D) 50 µm; (G,J) 10 µm; (H) 2 µm; (I) 5 µm; (K) 250 µm; (L) 3 µm.
Arthonia atra is phenotypic plastic among individuals, depending on changing environmental
conditions. In total, 24 intraspecific taxa have been described within A. atra, and their names
are regarded as synonyms. Based on the shape of their ascomata, Nowak [10] distinguished
11 forms of O. atra [for. atra – O. cerasi Chev., for. fibricola B. de Lesd., for. reticulata (DC.)
J. Nowak, for. rimosa (DC.) J. Nowak, for. denigrata (Ach.) Leight., for. stenocarpa (Ach.) –
O. stenocarpa Ach., for. arthonioidea (Leight.) Leight., for. nigrita Leight., for. platanoides
Nyl., for. stellulata Mereschk., for. orbicularis B. de Lesd.], including two new ones described
from Poland (for. reticulata and for. rimosa). However, these forms do not appear to deserve
separate taxonomic rankings, as the character variation is continuous throughout all examined
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
material, with many specimens having intermediate features and parameters. In the analyzed
material, the ascomata usually were elliptic or elongate, and circular ascomata were very
rare (Fig. 38). The ascomata length ranged from 154 µm to 797 µm. Their distribution was
scattered or clumped (with similar frequency). The examined specimens had flat or slightly
convex discs, rarely concave. In addition to the size and shape of their ascomata, the color
of the thalli also varied. They usually contain abundant, convex pycnidia, but the shape of
conidia was variable. They usually were slightly curved, but sometimes the same specimen
showed both straight and curved conidia.
100
D
90
80
C
C
C
B
70
C
C
60
B
%
B
50
40
30
B
B
A
B
B
20
A
10
0
A
A
1
2
3
4
A
A
5
6
A
7
Fig. 38 Variation in selected morphological and anatomical features Arthonia atra.
1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata: A – unbranched,
B – branched; 3 – disc type: A – concave, B – flat, C – convex; 4 – thallus color: A – white,
B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia:
A – few, B – numerous, C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
Nomenclatural notes. The taxon was first described by Persoon in 1794 [283], who named it
Opegrapha atra. It is noteworthy that Schneider in 1898 [284] was the first to call it Arthonia
atra, but this name was not commonly used. However, according to the latest taxonomic
classification [6], the genus Arthonia in the order Arthoniales is confirmed. Previously,
O. calcarea was included in Arthonia as well [6] due to the different structure of their asci
[34]. This conclusion is confirmed by the present study. The genus Arthonia is a heterogeneous
group of lichens [33,285] that now contains about 400 species, the largest genus in the order
Arthoniales [286].
Distribution outside Europe. Species with a wide geographic range. Recorded from North
America (Canada [66], Alabama [287], the Bahamas [142], Florida [139], Louisiana [138],
New York [288], South Carolina [289], Wisconsin [290]), Bermudas [291]), in South America
(Argentina [141]), in Africa (Lebanon [292], Morocco [65], Tunisia [88]), and in Oceania
[293].
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Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], Arthonia
atra is a boreal-Mediterranean, sub-Atlantic species. It grows on the bark of various deciduous tree species, but is less often found on conifers. It is most frequently found on Carpinus
betulus, Fagus sylvatica, Quercus spp., and Acer spp., but less frequently on the bark of Fraxinus
excelsior, Abies spp., and Picea spp. Only occasionally is it found on Pinus spp. This species
usually colonizes moderately acidic substrates, with pH 4.9–5.6 [38].
In Europe, it is known from numerous localities, reported, e.g., from Albania [294],
Austria [71], the Czech Republic [295], Denmark [122,163], Estonia [123], Finland [165],
France [166], Germany [124,168,170,296–300], Great Britain [92], Greece [127], Hungary
[93], Ireland [94], Italy [74,171], Latvia [172], Lithuania [173], Macedonia [301], Malta [95],
the Netherlands [96], Norway [97], Portugal (Madeira [75]), Romania [302], Slovenia [76],
Spain [99], Sweden [97], Switzerland [129], Turkey [153–155], and Ukraine [100].
Habitat and distribution in Poland. In Poland, Arthonia atra grows commonly on the bark
of deciduous trees, including Fagus sylvatica, Quercus spp., Fraxinus excelsior, Acer pseudoplatanus. It can be found less often on Acer spp., Carpinus betulus, or Salix spp, and very
rarely on Betula spp., Corylus avellana, Populus spp., and Ulmus spp. Nowak [10] reported it
on coniferous trees, including twigs of Myrica gale, and on decaying wood.
Most A. atra localities are found in northern Poland, while several isolated specimens
can occur in central Poland. Additionally, the species was recorded from the Sudetes and
Carpathians (Fig. 39).
The first record of A. atra was published by Körber [21]. It was also reported by Ohlert
[77], Stein [182], Eitner [303], Lettau [78], Boberski [183–185]. Later, the species was many
times reported in the Polish lichenological literature from: the Beskid Sądecki Mountains
[103], Beskidy Lesiste Mountains [191], Bory Tucholskie (Tuchola Forest) [194,195], Dolny
Fig. 39 Distribution and substrate preference of Arthonia atra in Poland.
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Śląsk (Lower Silesia Province) [132], Gorce Mountains [197], Stołowe Mountains [198],
Świętokrzyskie Mountains [105,199–201], Kotlina Biebrzańska (Biebrza Basin) [205], Kotlina Toruńska (Toruń Basin) [206], Łańcuch Tatrzański macroregion [107,208], Małopolski
Przełom Wisły (Lesser Poland Gorge of the Vistula) [212], Masyw Ślęży massif [213,214],
Niecka Włoszczowska basin [216], Pieniny Mountains [111,219], Płaskowyż Nałęczowski
(Nałęczów Plateau) [220], Pobrzeże Kaszubskie (Kashubian Coastland) [79,221,304], Pobrzeże
Szczecińskie (Szczecin Coastland) [80,222], Podgórze Rzeszowskie (Rzeszów Foothills)
[223], Pogórze Przemyskie (Przemyśl Foothills) [112], Pojezierze Ełckie (Ełk Lakeland)
[305], Pojezierze Gnieźnieńskie (Gniezno Lake District) [227], Pojezierze Iławskie (Iława
Lake District) [230], Pojezierze Kaszubskie (Kashubian Lake District) [83,232], Pojezierze
Krajeńskie (Krajna Lake District) [233], Pojezierze Mrągowskie (Mrągowo Lake District)
[234], Pradolina Wieprza (Wieprz River Urstromtal) [237], Pradolina Wrocławska (Wrocław
Urstromtal) [238], Równina Augustowska (Augustów Plain) [240–242,306,307], Równina
Bielska (Bielsk Plain) [55], Równina Biłgorajska (Biłgorajska Plain) [308], Równina Drawska
(Drawno Plain) [85,244], Równina Łęczyńsko-Włodawska (Łęczna-Włodawa Plain) [247,248],
Równina Łukowska (Łuków Plain) [250,251], Równina Piotrowska (Piotrków Plain) [252,253],
Roztocze [254–258], Sudetes [260,261], Śląsk Opolski (Opole Silesia) [262], West Beskids
[190,191], West Bieszczady Mountains [309], Western Pomerania [56,263,264], Wzgórza
Radomszczańskie (Radomsko Hills) [216], Wybrzeże Słowińskie (Slovincian Coast) [266],
Wysoczyzna Białostocka (Białystok Highland) [269–272], Wysoczyzna Elbląska (Elbląg
Highland) [273], Wysoczyzna Kaliska (Kalisz Highland) [274], Wysoczyzna Łaska (Łask
Highland) [276], Wysoczyzna Siedlecka (Siedlce Highland) [277], Wyżyna Lubelska (Lublin
Upland) [87], and Zaklęsłość Sosnowicka (Sosnowica Depression) [282].
Most of A. atra localities are placed in northern Poland, while several isolated ones are
in central Poland. Additionally, the species was recorded from the Sudetes and Carpathians
(Fig. 39).
4.6.6. Arthonia calcarea (Turner ex Sm.) Ertz & Diederich
in Ertz, Miądlikowska, Lutzoni, Dessein, Raspe, Vigneron, Hofstetter & Deiderich, Mycol. Res. 113(1): 146
(2009).
Opegrapha calcarea Turner ex Sm., in Smith & Sowerby, Engl. Bot. 25: Tab. 1790 (1807).
= Opegrapha chevallieri Leight., Ann. Mag. Nat. Hist., Ser. 213: 90 (1854).
Thallus thin, white. Ascomata black, not pruinose, elevated, solitary or in small groups, unbranched or branched, 0.3–1.2 × 0.15–0.3 mm. Discs narrow, reduced to a crevice, concave,
15–35 µm wide. Excipulum thick, elevated, permanent, black, naked, continuous below the
hypothecium, 20–35 µm thick laterally and 30–45 µm at the base. Hypothecium grey-brown,
15–25 µm high. Hymenium 40–70 µm high, I+ blue. Pseudoepithecium hardly visible. Asci
club-shaped, 35–50 × 15–20 µm, inner layer of endoascus slightly amyloid, limited to the
dome zone (calcarea-type). Ascospores hyaline, ellipsoid, three-septate, (13)15–18 × 4–6 µm,
with perispore of 0.5 µm thick, sometimes hardly visible. Pycnidia sunken in thallus, rarely
convex, conidia straight 5–8 × 0.8–1(1.2) µm (Fig. 40). Chemistry: thalli K−, C−, PD−.
Discussion. Arthonia calcarea can be distinguished from other epilithic taxa found in Poland
primarily by its longer conidia, the presence of an amyloid layer around the ascus, and based on
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 40 Arthonia calcarea (Turner ex Sm.) Ertz & Diederich: (A) thallus
and ascomata, LM (leg. A. Wieczorek, 1997, herb. Wieczorek); (B–D)
thallus and ascomata, SEM [(B) leg. A. Wieczorek, July 20, 1997, herb.
Wieczorek; (C) s. coll., KRAM-L 27263; (D) s. coll., KRAM-L 26972); (E)
ascus with ascospores, LM (s. coll., KRAM-L 27263); (F) ascospores, SEM
(leg. A. Wieczorek, 1997, herb. Wieczorek); (G,H) ascospores, LM (s. coll.,
KRAM-L 27263); (I,J) pycnidium, SEM (s. coll., KRAM-L 27263); (K)
conidium, SEM (leg. A. Wieczorek, 1997, herb. Wieczorek); (L) conidia,
LM (leg. A. Wieczorek, 1997, herb. Wieczorek). Scale bars: (A,B) 200 µm;
(C,D) 300 µm; (E) 20 µm; (F–H) 10 µm; (J) 50 µm; (K) 3 µm; (L) 5 µm.
the structure of the apical apparatus. Identification on the basis of other anatomical structures
can be difficult. Ascospore size and shape are similar to those in Opegrapha saxicola Ach,
particularly for individuals that develop a visible perispore. However, perispore’s thickness in
A. calcarea does not exceed 1 µm, but mature, well-developed spores of O. saxicola contain
thicker perispores. Arthonia calcarea differs from O. dolomitica by having ascospores shorter
than 20 µm.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Since Arthonia calcarea is rare in Poland and there are few herbarium specimens, it is
difficult to assess its intraspecific variability. Most populations vary primarily in shape of the
ascomata. In some individuals, the ascomata are circular, solitary, unbranched, and resemble
those in Opegrapha dolomitica. Occasionally, the ascomata are ellipsoid to elongate. Only
concave discs were observed in the studied specimens. The ascomata with two types of
excipulum were observed: Types A and B. Usually, there are few pycnidia, both sessile and
convex (Fig. 41). Perispore thickness also is variable in ascospores. In some individuals, it is
reduced and poorly developed, whilst in others, it is narrow but well defined.
100
90
C
80
70
C
B
A
A
6
7
B
%
60
50
B
A
A
A
40
30
20
10
0
A
A
1
2
3
4
5
Fig. 41 Variation in selected morphological and anatomical features of Arthonia
calcarea. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata:
A – unbranched, B – branched; 3 – disc type: A – concave, B – flat, C – convex;
4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very numerous;
7 – type of pycnidia: A – sessile, B – convex.
Nomenclatural notes. Opegrapha calcarea was described by Smith and Sowerby [310]. The
name was taken from Turner’s manuscripts, who described the species from “the mortar of
old walls in Norfolk and Suffolk”. The name was accepted by Acharius [2,3]. Stizenberger [4]
proposed a different approach and regarded this species, with other epilithic lichens (O. trifurcata Hepp and O. chevallieri), as synonymous with O. atra, on the basis of similar ascospore
shape and size. His approach, however, was not accepted by a majority of lichenologists, who
continued using the name O. calcarea [1,116,311–313].
Considering ascospore dimensions, Pentecost and Coppins [119] as well as Purvis et al.
[40] treated A. calcarea as an ecotype of Opegrapha saxatilis. However, according to Clauzade
and Roux [64], O. saxatilis has a discontinuous excipulum below the hypothecium, therefore
the species cannot be treated as synonyms.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Detailed studies on this species were conducted by Torrente and Egea [34], who compared
three similar, closely related species (O. calcarea, O. chevallieri, and O. trifurcata). They pointed
out that the three species are conspecific and represent O. calcarea. Smith et al. [41] deviated
from their earlier classification by assuming that O. saxatilis was a synonym of O. calcarea,
and they took into account other morphotypes of this taxon, including O. conferta and
O. chevallieri.
The treatment of Arthonia calcarea as a separate species is justified by the characteristic
ascus type and the continuous excipulum below the hypothecium. This was confirmed by
Clauzade and Roux [64], Torrente and Egea [34], and by the results of my research.
Distribution outside Europe. It was recorded from North America (Canada [66], the Bahamas
[142]), in Asia (Taiwan [69], Armenia [120]), and in North Africa (Morocco [65] and Tunisia
[88]). Moreover, it has been reported from the Canary Islands [314].
Habitat and distribution in Europe. According to Smith et al. [41], the species distribution
is Central European-Mediterranean and sub-Atlantic. In Europe, it is found on calcareous
rocks and occasionally sandstone. According to Torrente and Egea [34], they are observed
in Western Europe mostly in coastal zones, but they are known also from a few localities in
Central Europe, including Styria in Austria [160].
The species was reported, e.g., from Austria [71], Cyprus [162], Denmark [122,163], Germany [91,168,298,299], Greece [315], Hungary [93], Italy [74], Montenegro [316], Norway
[97], Portugal (Madeira [75]), Slovenia [76], Spain [99], and Ukraine [100].
Habitat and distribution in Poland. Like most of the epilithic species, Arthonia calcarea
grows on rocks of various types (limestone and dolomite, rarely sandstone), usually on dry,
sunny, vertical sites. The various local climates and microclimates associated with the varied
relief of mountainous terrain, likely provide favorable conditions for this species. Arthonia
calcarea was reported for the first time by Eitner [303] in the Sudetes. It was later reported
from the Łańcuch Tatrzański macroregion [106,107], Sudetes [260], and Dolny Śląsk (Lower
Silesia Province) [132].
Only two contemporary collections were available to study and they both originated from
the Carpathians (Fig. 42). However, the KRAM herbarium includes three specimens (one
dating back to 1847), with no location or author given.
4.6.7. Gyrographa gyrocarpa (Flot.) Ertz & Tehler
in Ertz, Tehler, Irestedt, Frisch, Thor & van den Boom, Fungal Diversity 70: 42 (2015).
Opegrapha gyrocarpa Flot., Flora, (Regensburg) 8: 345 (1825).
Thallus with very characteristic, yellowish soralia, poorly developed, often forming larger
aggregations. Ascomata rare, if present, usually circular to ellipsoid, solitary or in groups, very
rarely branched, 0.5–2.0 mm in diameter. Discs strongly wrinkled, sometimes limited from the
center with a roundish ridge or irregularly twisted, black, not pruinose. Excipulum thick, permanent, undulated or convoluted, black, naked, dull or slightly shiny, brittle, continuous below
the hypothecium, up to 500 µm thick at the base, K+ olive-green. Hypothecium pale to nearly
hyaline, 20–25 µm high. Hymenium hyaline, ca. 100 µm high, I+ reddish. Pseudoepithecium
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 42 Distribution of Arthonia calcarea in Poland.
inconspicuous. Asci club-shaped or cylindrical, 80–90 × 12–15 µm. Ascospores hyaline, narrowly spindle-shaped, three-septate, 20–26 × 3–5 µm, without perispore. Pycnidia scattered,
sunken, visible as very small, blackish dots. Conidia straight, 5–6 × 0.3−0.5 µm (Fig. 43).
Thallus K−, C+ reddish, KC+ reddish (mainly the inner part and soralia are stained), PD−.
Three chemotypes were described by Tønsberg [317].
Discussion. In Poland, Gyrographa gyrocarpa is the most distinctive species among the
epilithic representatives of the group. It rarely is confused with other species. Due to the
characteristic soredia, it easily can be identified, even when sterile. In addition to the color
reaction of the soredia, it differs from other species in the following characteristics: (i) large
(but not always present), nearly circular or irregular ascomata and (ii) three-septate ascospores
without perispore.
The species is usually sterile. In the sterile form, it is similar to Enterographa zonata, but
it differs in the color reaction of the medulla and soredia (C+ and KC+ reddish). In some
localities (e.g., in the Sudetes), G. gyrocarpa produces numerous ascomata. Like in other taxa
of the genus Opegrapha s. l., the ascomata are variable, usually elliptic or circular, very rarely
elongate. They usually are solitary, producing various types of discs from concave to flat or
convex. The species is variable in thallus color and in the abundance of pycnidia (Fig. 44).
Nomenclatural notes. Gyrographa gyrocarpa was described in 1825 as Opegrapha gyrocarpa
[318]. Previously, two varieties were distinguished: var. arenaria Körber [319] and var. schisticola
(Eitn.) Migula [313]. The genus Gyrographa was described by Ertz et al. [9]. In the earlier
literature, polymorphic characters were noted by Tønsberg [317], and it was even suggested
that the species should be moved to the related genus Lecanactis [57].
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 43 Gyrographa gyrocarpa (Flot.) Ertz & Tehler: (A,B) thallus and ascomata, LM (leg. M. Kossowska, 2008, herb. Kossowska); (C–G) thallus and
ascomata, SEM [(C–E) leg. K. Szczepańska, 2003, herb. Szczepańska; (F,G) leg.
P. Czarnota, GPN 2145]; (H) ascus with ascospores, LM (leg. Z. Tobolewski,
1960, POZ); (I) ascospore, SEM (leg. K. Szczepańska, 2003, herb. Szczepańska);
(J,K) ascospores, LM (leg. A. Wieczorek, 1989, herb. Wieczorek); (L) conidia,
LM (leg. A. Wieczorek, 1989, herb. Wieczorek). Scale bars: (A,C,E,G) 100 µm;
(B) 500 µm; (D) 200 µm; (F) 250 µm; (H–K) 10 µm; (L) 5 µm.
Distribution outside Europe. Species with a wide geographic range, including North America,
e.g., Canada [66], and part of Asia, e.g., Japan [149].
Habitat and distribution in Europe. According to Wirth [38], Wirth et al. [39], and Purvis
et al. [40], the species distribution is boreal-Central European and sub-Atlantic. It is most
frequently found on sandstone and granite.
In Europe it is known, e.g., from Austria [71,160], the Czech Republic [295], Finland
[165], Germany [124–126,168,170,320], Great Britain [92], Italy [74], the Netherlands [96],
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
100
C
90
C
B
C
80
B
D
70
C
B
%
60
B
50
40
C
A
B
30
A
B
20
B
A
10
A
A
A
A
0
1
2
3
4
5
6
7
Fig. 44 Variation in selected morphological and anatomical features Gyrographa gyrocarpa. 1 – ascomata shape: A – circular, B – elliptic, C – elongate;
2 – ascomata: A – unbranched, B – branched; 3 – disc type: A – concave, B – flat,
C – convex; 4 – thallus color: A – white, B – grey, C – white-grey, D – brownish;
5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very
numerous; 7 – type of pycnidia: A – sessile, B – convex.
Norway [97], Portugal (Madeira, Azores [75,98]), Romania [302], Slovenia [76], Spain [99],
and Sweden [97].
Habitat and distribution in Poland. Gyrographa gyrocarpa can be found on rocks of woodlands
located in valleys and gorges, and on vertical or hanging rocks that are dry or moist, with
north or northeastern exposure. They are less often found in open or sunny habitats.
The first record of Gyrographa gyrocarpa was published by Körber [21]. It was also recorded
by Stein [182] and Eitner [303], but it was erroneously reported from calcareous rocks in
some southern parts of Poland.
It was later reported in the Polish lichenological literature from the Beskid Mały Mountains
[321], Beskid Niski Mountains [322], Beskid Sądecki Mountains [102,103,323], Beskid Śląski
Mountains [190], Beskidy Lesiste (Wooded Beskids) [191], Gorce Mountains [324], Stołowe
Mountains [198], Sowie Mountains [325], Karkonosze Mountains [326,327], Łańcuch Tatrzański
macroregion [107,210], Pogórze Rożnowsko-Ciężkowickie (Rożnowsko-Ciężkowickie Foothills) [81], Sudetes [261,328], West Beskids [329], West Bieszczady Mountains [309].
Gyrographa gyrocarpa was recorded exclusively from the southern part of Poland (Fig. 45),
with a center of distribution in the Polish Carpathians. It can also be found in the West
Beskids and Beskidy Lesiste. It is more rare to find it in the Sudetes, Karkonosze Mountains,
and Stołowe Mountains.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 45 Distribution and substrate preference of Gyrographa gyrocarpa in Poland.
4.6.8. Opegrapha dolomitica (Arnold) Clauzade & Cl. Roux ex Torrente & Egea
Bibliotheca Lichenol. 32: 146 (1989).
Opegrapha rupestris var. dolomitica Arnold, Flora (Regensburg) 43: 78 (1860).
Opegrapha saxicola var. dolomitica (Arnold) V. Wirth, Flechtenflora (Stuttgart): 531 (1980).
Thallus grey to grey-white, sometimes with black prothallus. Ascomata elliptic, sometimes
curved and branched in maturity, 0.2–0.8 × 0.4–0.8 mm, solitary or in groups of various size;
surface not pruinose, protrude above the thallus, very rarely sunken. Discs black, narrow,
rarely broad. Excipulum thick, permanent, black, naked, continuous below the hypothecium,
40−60 µm thick laterally and 80–150 µm at the base. Hypothecium grey to grey-brown, dark,
20–25 µm high. Hymenium 50–70 µm high, I+ reddish. Pseudoepithecium brown, K−.
Asci broadly club-shaped, mature 60–70 × 19–22 µm, young 45–57 × 16 µm, inner layer of
endoascus amyloid in upper part, ring and ocular chamber reduced (varia-type). Ascospores
hyaline, ellipsoid, three-septate, (18)22–29 × 5–6 µm, with perispore of 0.5–0.7(1.0) µm thick.
Pycnidia elevated or sessile, black. Conidia straight, 4–6 × 0.5–1 µm (Fig. 46). Chemistry:
thalli K−, C−, PD−.
Discussion. Opegrapha dolomitica can be distinguished from the similar Opegrapha saxicola by
its markedly longer ascospores, which are more than 22 µm long, with a thin hyaline perispore.
Opegrapha dolomitica differs from Gyrographa gyrocarpa by the presence of a perispore and
the lack of a color reaction on the thallus.
Most populations of O. dolomitica vary in size and shape of the ascomata. In some specimens, all of the ascomata were elliptic, unbranched, and elevated. In others, the ascomata were
branched and meandering. However, in some specimens, a broad range of shape variation was
observed. For this reason, like in other Opegrapha species, the shape and size of the ascomata
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Fig. 46 Opegrapha dolomitica (Arnold) Clauzade & Cl. Roux ex Torrente
& Egea: (A,B) thallus and ascomata, LM [(A) leg. J. Nowak, KRAM-L 12158;
(B) leg. J. Nowak, KRAM-L 9661]; (C–E) thallus and ascomata, SEM [(C,E)
leg. J. Nowak, KRAM-L 30447; (D) leg. J. Nowak, KRAM-L 12158); (F)
ascus with ascospores, LM (leg. J. Nowak, KRAM-L 30447); (G) ascospore,
SEM (leg. J. Nowak, KRAM-L 30447); (H,I) ascospores, LM (leg. J. Nowak,
KRAM-L 30447); (J,K) pycnidia, SEM (leg. J. Nowak, KRAM-L 30447);
(L) conidia, LM (leg. J. Nowak, KRAM-L 30447). Scale bars: (A,B,D,E,K)
100 µm; (C) 150 µm; (F,G) 25 µm; (H,I) 10 µm; (J) 250 µm; (L) 5 µm.
cannot be regarded as a good diagnostic feature. Disc type also is variable within populations
and on individual specimens. They have an excipulum of Type C, but A or B may be found
occasionally. The pycnidia are numerous, sessile, and convex (Fig. 47).
Nomenclatural notes. Opegrapha dolomitica is one of the newest species of this genus to
be distinguished at the species rank [34]. Stizenberger [4] wrote “a frequently noticeable
form of this species (O. saxicola) is dolomitica Arn., but I do not attempt to establish its
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
100
C
C
D
90
B
C
80
B
C
B
70
C
B
B
%
60
50
40
B
A
30
A
A
20
A
B
A
10
A
A
0
1
2
3
4
5
6
7
Fig. 47 Variation in selected morphological and anatomical features Opegrapha
dolomitica. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata:
A – unbranched, B – branched; 3 – disc type: A – concave, B – flat, C – convex;
4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – cross
section of a typical ascoma; 6 – number of pycnidia: A – few, B – numerous,
C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
systematic position”. Most often it was regarded as a variety or subspecies of O. rupestris or
O. saxicola [O. rupestris var. dolomitica Arnold [330], O. saxicola subsp. dolomitica (Arnold)
Roux [331]].
Nowak [10] discussed the species under the name O. saxicola for. dolomitica (Arnold)
Arnold.
Distribution outside Europe. Opegrapha dolomitica has a Holarctic distribution, covering
the whole Europe and North America (the Bahamas [142], Canada [332]). Recorded also
from Africa (Morocco) [65].
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], this
species distribution is boreal Mediterranean. In Europe, it grows on limestone or dolomitic
rocks.
Reported, e.g., from Austria [71], the Czech Republic [295], Denmark [122,163], Great
Britain [92], Italy [74], Norway [97], Romania [178,302], Russia (Dagestan [333]), Spain [99],
Sweden [97], and Ukraine [334].
Habitat and distribution in Poland. In Poland, Opegrapha dolomitica occurs less frequently
on limestone than on other rocks, such as sandstones. It occurs typically on vertical and
smooth rock walls. The first record of O. dolomitica was published by Körber [21]. It was also
reported by Stein [182], Eitner [303], Migula [130], and a few times in the Polish lichenological
literature from Dolny Śląsk (Lower Silesia Province) [132], Łańcuch Tatrzański macroregion
[106,107], Pieniny Mountains [110,111], Sudetes [260], and West Beskids [131].
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 48 Distribution and substrate preference of Opegrapha dolomitica in Poland.
Like most of the epilithic species of the studied genera, O. dolomitica is found in the
southern part of the country, with a few scattered localities in the western part of the Polish
Carpathians and uplands (Fig. 48).
4.6.9. *Opegrapha geographicola (Arnold) Hafellner
Herzogia 10: 12 (1994).
Phaeospora geographicola Arnold, Verh. Zool.-Bot. Ges. Wien 46: 140 (1896).
It is a lichenicolous nonlichenized fungus. It is parasitic on the host lichen thallus (Rhizocarpon
badioatrum). Ascomata black, roundish, not pruinose, usually elevated above the host thalli.
Discs black, narrow, rarely broad. Excipulum permanent, thick, black, continuous below
the hypothecium. Hypothecium 10−15 µm high. Hymenium 45–80 µm high, I+ reddish.
Pseudoepithecium inconspicuous. Asci broadly club-shaped, mature 55–70 × 15–20 µm.
Ascospores pale brown, ellipsoid, two−three-septate, 18–29 × 7–9 µm, darkening in herbarium
specimens, with perispore of 0.8–1.4 µm thick. Wall of ascospores I+ blue. Pycnidia elevated
or sessile, black. Conidia straight, 4–6 × 1 µm (Fig. 49).
Discussion. This species is morphologically similar to Opegrapha dolomitica and Phacographa
glaucomaria, especially to the individuals that develop ascospores of the same size. It differs
from the above taxa primarily due to its pale brown ascospores, as well as the color reaction
of the transverse walls of the ascospores (I+ blue). Additionally, O. geographicola produces
both three-septate and two-septate ascospores. It is difficult to understand the variability of
this species, because only few herbarium specimens are available.
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 49 Opegrapha geographicola (Arnold) Hafellner: (A,B) host thallus
of Rhizocarpon badioatrum and apothecia, LM; (B) apothecium, LM;
(D) ascus with ascospores, LM; (E) ascospore, LM; (F) pycnidia, LM
(leg. A. Flakus, 2740/1, KRAM). Scale bars: (A,F) 1 mm; (B) 2 mm;
(C,D) 30 µm; (E) 20 μm.
Fig. 50 Distribution of Opegrapha geographicola in Poland.
General distribution. Recorded from Alaska [67]. Known also from a few localities in Europe,
e.g., from Austria [335] and Spain [99].
Habitat and distribution in Poland. This is a lichenicolous fungus that grows on Rhizocarpon
badioatrum and is known only from one locality in the Western Tatras [336] (Fig. 50).
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
4.6.10. Opegrapha lithyrga Ach.
Lichenogr. Univers.: 247 (1810).
Thallus poorly developed, thin, smooth or granular, greyish. Ascomata solitary or in small
groups, unbranched, rarely branched, straight or curved, 0.2–1 × 0.15–0.25 mm. Discs narrow, concave, not pruinose. Excipulum thick, permanent, black, naked, continuous below
the hypothecium, 13–18 µm thick laterally and 40–60 µm at the base. Hypothecium greybrown, 15–25 µm high. Hymenium hyaline, 60–70 µm high, I+ reddish. Pseudoepithecium
insignificant, very narrow. Asci cylindrical-clavate, mature 40−60 × 15−18 µm, inner layer of
the endoascus amyloid along its whole length, ring hardly visible (vulgata-type). Ascospores
hyaline, spindle-shaped, straight or slightly curved, five–seven-septate, 21–35 × 3–5 µm, with
perispore of 1–1.5 µm thick. Pycnidia numerous, black, very small, sunken in thallus. Conidia
straight, 3–5 × 1–1.5 µm (Fig. 51). Chemistry: thalli K−, C−, PD−.
Discussion. The species is morphologically similar to Arthonia calcarea and Alyxoria mougeotii,
especially the individuals that develop solitary, unbranched ascomata. The differences between
the species primarily concern the structure of the apical apparatus and the color reaction of
the inner layer of the endoascus: calcarea-type in A. calcarea, varia-type in A. mougeotii, and
vulgata-type in O. lithyrga. Moreover, O. lithyrga differs from A. calcarea by five–seven-septate
ascospores, and from A. mougeotii by narrower ascospores, with all cells of similar size.
Opegrapha lithyrga is closely related to O. vulgata [7]. However, the former is epilithic,
while the latter is epiphytic. Moreover, O. lithyrga has much larger, mature asci, slightly wider
ascospores, and shorter conidia. Undoubtedly, the species evolved by adapting to its substrate,
which was confirmed by recent molecular analyses [7] that support their species status in
spite of the close relationship.
Opegrapha lithyrga varies remarkably in size and shape of their ascomata. Nowak [10]
distinguished three varieties with two forms (var. lithyrga, for. lithyrga, [for.] b. steriza Ach.,
[var]. β. ochracea Körb., var. vestita Redgr.) to reflect this variation.
This species has elliptic and elongate ascomata, with narrow concave or flat discs, and one
excipulum type. Some specimens only have short and straight conidia (e.g., a specimen from
WA herbarium), while others have long and curved conidia (Fig. 52). Opegrapha lithyrga is
a well described species that is easy to distinguish from other epilithic species. Its spore size is
uniform, 23–24 × 3–3.5 µm, and its perispore is well developed. All of the analyzed specimens
had single and unbranched ascomata, with narrow and concave discs. They mostly represent
the typical var. lithyrga described by Acharius [2].
Nomenclatural notes. The species was described by Acharius [2], who gave it the binomial
name that is still in usage. Its similarity to O. vulgata was noted by Stizenberger [4], who
classified it as O. vulgata var. lithyrga. Opegrapha lithyrga was listed in many works published
in the nineteenth century as O. lithyrga [2,3,25], as O. vulgata for. lithyrga, [285], as O. lithyrgiza [5], or as synonymous with O. vulgata [4]. In the early twentieth century, it was treated
as the distinct species O. lithyrga (e.g., [60]). Redinger [1] distinguished two varieties: var.
grisea Körb. and var. ochracea Redgr., whereas Nowak [10] discussed three varieties: var.
lithyrga, var. ochracea, and var. vestita (see above). Both authors distinguished them on the
basis of morphological variation of the thalli. The lack of synonyms indicates that this taxon
is interpreted rather unambiguously.
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Fig. 51 Opegrapha lithyrga Ach.: (A,B) thallus and ascomata, LM [(A)
leg. J. Nowak, KRAM-L 13909; (B) leg. J. Nowak, KRAM-L 5059]; (C–F)
thallus and ascomata, SEM [(C,E) leg. J. Nowak, KRAM-L 5059; (D,F) leg.
J. Nowak, KRAM-L 13909]; (G) ascus with ascospores, LM (leg. J. Nowak,
KRAM-L 5557); (H,I) ascospores, LM (leg. J. Nowak, KRAM-L 5557); (J)
pycnidium, LM (leg. J. Nowak, KRAM-L 13909); (K) pycnidium, SEM (leg.
J. Nowak, KRAM-L 13909); (L) conidia, LM (leg. J. Nowak, KRAM-L 5557).
Scale bars: (A,B,D,F,J) 100 µm; (C,E,K) 50 µm; (G,I) 10 µm; (L) 5 µm.
Distribution outside Europe. They are distributed in North America in mountainous regions,
such as the high altitudes in Canada.
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], it is
a Central European and Mediterranean species. In Europe, it is found on acidic shale/slate,
granite, gneiss, and quartzite. It prefers smooth and vertical or hanging surfaces, in shaded
habitats, usually with northern exposure that are dry or at least periodically slightly moist
[10]. It probably spread from the south of Sweden and Norway to the Alps and southwestern
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100
90
D
80
B
C
70
B
B
A
A
6
7
%
60
50
C
B
40
30
A
B
20
A
10
0
1
2
4
5
Fig. 52 Opegrapha lithyrga, variability of selected morphological-anatomical features. 1 – ascomata
shape: A – circular, B – elliptic, C – elongate; 2 – ascomata: A – unbranched, B – branched; 4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of
pycnidia: A – few, B – numerous, C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
Europe [34]. In Central Europe, it grows together with Enterographa zonata (Körb.) Källsten,
E. hutchinsiae (Leight.) A. Massal. [38], whereas on the Atlantic coast of the Iberian Peninsula,
it grows with O. gyrocarpa and O. saxigena Taylor [34].
In Europe, known, e.g., from Austria [71,160], Denmark [122,163], France [167], Germany
[124–126,170,320,337], Great Britain [92], Ireland [94], Italy [74], Malta [95], Portugal (Madeira, Azores [75,98]), Spain [99], Turkey [153,154], and Ukraine [100].
Habitat and distribution in Poland. It grows on sandstone, but less commonly on other rock
types. It is a very rare species. The first record of O. lithyrga was published by Nowak [187].
It has been reported from the Beskid Makowski Mountains [186], Beskid Mały Mountains
[187], Beskid Sądecki Mountains [102], Beskid Śląski Mountains [189], Łańcuch Tatrzański
macroregion [210], Podhale [224], and Wyżyna Wieluńska (Wieluń Upland) [280].
Only three voucher specimens support records of this species in the western part of the
Polish Carpathians (Fig. 53).
4.6.11. Opegrapha niveoatra (Borrer) J. R. Laundon
Lichenologist 2(2): 138 (1963).
Verrucaria niveoatra Borrer, in Hooker, Engl. Bot., Supplement. 1: Tab. 2637 (1831).
= Opegrapha vulgata var. subsiderella Nyl., Lich. Scand. (Helsinki): 255 (1861).
= Opegrapha subsiderella (Nyl.) Arnold, Flora (Regensburg) 67: 656 (1884).
Thallus thin, continuous, sometimes in the form of spots, whitish or greyish, sometimes with
green tint, with dark prothallus. Ascomata numerous, solitary or in groups, unbranched or
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Fig. 53 Distribution of Opegrapha lithyrga in Poland.
more often branched, 0.3–1.5 × 3–5 mm. Discs narrow or delicately widened, concave, black,
not pruinose. Excipulum of various thickness, elevated, permanent, black, naked, continuous below the hypothecium, 25–30 µm thick laterally and 40–50 µm at the base. Hymenium
50–60 µm high, I+ reddish. Hypothecium 10–20 µm high. Pseudoepithecium very poorly
developed, ca. 5–10 µm high. Asci cylindrical-clavate, mature 45–55 × 10–13 µm, inner layer
of endoascus amyloid along its whole length, small ring around ocular chamber (vulgata-type).
Ascospores hyaline, spindle-shaped, straight or slightly curved, four–five(six)-septate, 20–25
× 2.5–3.5 µm, with inconspicuous perispore of 0.5–0.7 µm thick. Pycnidia numerous, small,
black. Conidia highly variable, straight or slightly or very strongly curved, 4–9 × 0.9–1.5 µm
(Fig. 54). Chemistry: thalli K−, C−, PD−.
Discussion. An exceptional feature that differentiates O. niveoatra from the other Opegrapha
species is the length of its conidia. In its typical form, O. niveoatra produces conidia that are
4–8(9) µm long. However, specimens without mature pycnidia are impossible to be distinguished from the closely related O. vulgata.
Morphological and anatomical variability of O. niveoatra (as in case of other species of the
genus) is driven by the environmental factors. Nowak [10] distinguished three forms within
the species [for. niveoatra, for. intermedia (B. de Lesd.) J. Nowak, and for. olivaceonigra (B. de
Lesd.) J. Nowak] that differ in size and shape of their ascomata. However, it should be noted
that the species was misinterpreted by the author (see “Nomenclatural notes”).
My observations show that conidial size and shape differ markedly between and within
populations of O. niveoatra. Specimens from Southern Europe are characterized by straight
conidia up to 9 µm long [34]. By contrast, in material from Poland, both straight and slightly
curved conidia are observed, sometimes on the same individual. Their length ranges from
4 µm to 8 µm, regardless of the curve angle (Fig. 55).
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Fig. 54 Opegrapha niveoatra (Borrer) J. R. Laundon: (A) thallus and
ascomata, LM (leg. W. Fałtynowicz, UGDA-L 3435); (B–F) thallus and
ascomata, SEM [(B,C) leg. S. Cieśliński & Z. Tobolewski, 1984, KTC;
(D) leg. W. Fałtynowicz, UGDA-L 3435; (E,F) leg. W. Fałtynowicz,
UGDA-L 3435); (G) ascus with ascospore, SEM (leg. W. Fałtynowicz,
UGDA-L 3435); (H,I) ascospore, SEM (leg. S. Cieśliński & Z. Tobolewski,
1983, KTC); (J) ascospore, LM (leg. S. Cieśliński & Z. Tobolewski, 1983,
KTC); (K) pycnidium, SEM (leg. W. Fałtynowicz, UGDA-L 3435); (L)
conidia, LM (leg. W. Fałtynowicz, UGDA-L 3435). Scale bars: (A) 200 µm;
(B–F) 100 µm; (G) 25 µm; (H–J) 10 µm; (K) 50 µm; (L) 5 µm.
Nomenclatural notes. Until the twentieth century, the taxon was named Opegrapha vulgata
var. subsiderella [338], and then it was named Opegrapha subsiderella [339]. The names
appeared, however, synonymously with O. niveoatra [340]. During the revision of voucher
specimens, I found some Polish collections that were misidentified following Nowak [10]. In
the treatment, Nowak described O. niveoatra that was characterized by a conidium length of
12−18 µm. This was misleading, as this size range of conidia is characteristic for O. vulgata.
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100
90
C
D
C
80
70
C
C
60
%
C
B
B
50
B
40
B
B
B
30
A
B
20
10
A
A
A
A
A
0
A
1
2
3
4
5
6
7
Fig. 55 Opegrapha niveoatra, variability of selected morphological-anatomical
features. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata:
A – unbranched, B – branched; 3 – disc type: A – concave, B – flat, C – convex;
4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very numerous;
7 – type of pycnidia: A – sessile, B – convex.
This mistake was due to the fact that Nowak [10] accepted Redinger’s concept of this species
[1]. The latter author treated O. niveoatra under the name O. cinerea Chevall. This contradicts the currently valid classifications, where O. cinerea is a synonym of O. vulgata, which is
characterized by curved conidia of such length [34,119,122]. However, according to Torrente
and Egea [34], O. niveoatra is characterized by straight conidia that are 4–8 µm long. This is
confirmed by the results of my study (Fig. 56). As a result of the revision of material in Polish
herbaria, O. niveoatra was renamed and included under O. vulgata.
Distribution outside Europe. This species is distributed in North Africa (Morocco [65],
Algeria [140]) and in a few localities in North America (Canada [66], USA [341]).
Fig. 56 Conidia of Opegrapha niveoatra, LM. (A) leg. Z. Tobolewski
& S. Cieśliński, 1987, KTC; (B,C) leg. S. Cieśliński & Z. Tobolewski,
1983, KTC.
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Habitat and distribution in Europe. Opegrapha niveoatra is a very rare sub-Atlantic species
[38]. It grows in Central Europe on the bark of deciduous trees, especially in open sites. It is
most frequently found on Fagus sylvatica, Fraxinus excelsior, and Quercus spp. Sometimes it
grows together with O. vulgata, O. ochrocheila, and Pertusaria leioplaca.
According to Torrente and Egea [34], this species is known from Central and northwestern
Europe. It is reported from, e.g., Austria [71], Denmark [122,163], Estonia [164], Finland
[128], France [166,167], Germany [168,300], Great Britain [92], Ireland [73], Italy [74], Latvia
[342], Lithuania [173], the Netherlands [96], Portugal (Madeira, Azores [75,98]), Romania
[179], Russia [343–345], Spain [99], Sweden [97].
Habitat and distribution in Poland. In Poland, O. niveoatra grows in open and sparsely
wooded habitats, on the smooth bark of these deciduous trees: Fagus sylvatica, Quercus spp.,
Acer pseudoplatanus, Carpinus betulus. Less often, it is found on other Acer spp., Ulmus spp.,
and Fraxinus excelsior. In mountains, it is found frequently on the bark of Abies alba.
Its first record was published by Grummann [346]. It also was reported by Eitner [303,347]
and Migula [130]. The species also was reported from the Beskid Makowski Mountains [186],
Beskid Niski Mountains [322], Beskid Sądecki Mountains [102,323,348], Beskid Śląski Mountains [189,190], Gorce Mountains [349], Karkonosze Mountains [350], Łańcuch Tatrzański
macroregion [107,210,211,351], Mierzeja Wiślana (Vistula Spit) [215], northeastern Poland
[218], Pieniny Mountains [110], Pogórze Przemyskie (Przemyśl Foothills) [112], Pogórze
Rożnowsko-Ciężkowickie (Rożnowsko-Ciężkowickie Foothills) [81], Pogórze Spiskie (Spisz
Foothills) [225], Pojezierze Ełckie (Ełk Lakeland) [82,305], Pojezierze Kaszubskie (Kashubian
Lake District) [83], Pojezierze Olsztyńskie (Olsztyn Lake District) [235,352,353], Równina
Augustowska (Augustów Plain) [306], Równina Bielska (Bielsk Plain) [55,243], Równina
Drawska (Drawno Plain) [244], Równina Kozienicka (Kozienice Plain) [245,246], Równina
Piotrkowska (Piotrków Plain) [252], Roztocze [255,258], Sudetes [260], Uznam and Wolin
islands [354], West Beskids [131,190,191], West Bieszczady Mountains [309,355], Western
Pomerania [56], Wysoczyzna Białostocka (Białystok Highland) [271,356], Wysoczyzna Elbląska
(Elbląg Highland) [273], and Wzgórza Radomszczańskie (Radomsko Hills) [216].
The species occurs in the mountains, where it is quite frequent in the western part of the
Polish Carpathians and in the Bieszczady Mountains, and in northern Poland where it is scattered along the Pobrzeże Koszalińskie (Koszalin Coastland), Pojezierze Wschodniopomorskie
(East Pomeranian Lake District), Pojezierze Mazurskie (Masurian Lake District), and Nizina
Północnopodlaska (North Podlachian Lowland). In addition, it is reported from the Wyżyna
Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Świętokrzyskie Mountains, and
Równina Kozienicka (Fig. 57).
4.6.12. Opegrapha saxicola Ach.
Syn. Meth. Lich. (Lund): 71 (1814).
= Opegrapha persoonii Ach., Methodus, Sectio prior: 17 (1803).
= Opegrapha rupestris auct. polon. (non Pers.).
Thallus epilithic, thin, smooth or delicately wrinkled, dull, usually composed of small grannules, sometimes powdery-granular, whitish, light grey. Ascomata not numerous, solitary or in
groups, elevated, 0.4–1 × 0.1–0.2 mm, variable in shape. Discs narrow or wide, not pruinose,
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Fig. 57 Distribution and substrate preference of Opegrapha niveoatra in Poland.
dull. Excipulum relatively thick, elevated, permanent, black, naked, continuous below the
hypothecium, 40–50 µm thick laterally and up to 90 µm at the base. Hypothecium from pale
brownish-yellowish to nearly hyaline, 20– 40 µm high. Hymenium hyaline, 60–80 µm high,
I+ blue. Pseudoepithecium poorly developed, 10–15 µm high. Asci club-shaped, 55–65 ×
15–20 µm, inner layer of endoascus amyloid in upper half (varia-type). Ascospores hyaline,
spindle-shaped, three-septate, 18–22(24) × 5–7 µm, with perispore of 1.0–1.2 µm thick.
Pycnidia numerous, small, black. Conidia straight, 4–6 × 0.5–1 µm, cylindrical (Fig. 58).
Chemistry: thalli K−, C−, PD−.
Discussion. Opegrapha saxicola differs from the otherwise similar Arthonia calcarea by the
varia-type ascus, ascospores longer than 18 µm, and shorter conidia that are up to 6 µm long
in Opegrapha saxicola and up to 8 µm long in Arthonia calcarea.
The species is not highly variable, as pointed out by Nowak [10]. This taxon is easy to
recognize due to its relatively thin thallus that usually is whitish and gray, and its narrow,
slender, elliptic, elongate, unbranched, or branched ascomata. It produces the ascomata with
relatively narrow discs (Fig. 59).
Nomenclatural notes. Throughout the nineteenth century, Opegrapha saxicola was treated
under various names, and the group of species has a complicated taxonomy. In 1989, Egea
and Torrente [34] treated O. saxicola with O. saxatilis, and O. persoonii was a synonym
of O. rupestris. A different concept was adopted by Purvis et al. [40], who accepted three
species in this group based on ascospore length: O. saxicola, O. saxatilis, and O. parasitica
(= O. persoonii). The authors’ concept later was modified to distinguish: O. dolomitica (=
O. saxicola auct., non Ach.) and O. rupestris (= O. persoonii), with the latter species as a strictly
lichenicolous, parasitic species growing on crustose Verrucariaceae, especially V. baldensis
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Fig. 58 Opegrapha saxicola Ach.: (A–F) thallus and ascomata, LM [(A,B)
leg. J. Nowak, KRAM-L 12277]; (C–F) thallus and ascomata, SEM [(C,D) leg.
J. Nowak, KRAM-L 12277; (E,F) leg. K. Toborowicz, 1975, KTC); (G) ascus
with ascospores, LM (leg. J. Nowak, KRAM-L 12277); (H,I) ascospores, LM
(leg. U. Bielczyk, KRAM-L 44572); (J,K) pycnidium, SEM (leg. J. Nowak,
KRAM-L12277); (L) conidium, LM (leg. J. Nowak, KRAM-L 12277). Scale bars:
(A,B,E,K) 100 µm; (C,D,F) 200 µm; (G) 20 µm; (H,I) 10 µm; (J) 50 µm; (L) 5 µm.
[41]. In the Polish material, the two species O. dolomitica and O. saxicola were distinguished
based on length of the ascospores. However, O. rupestris, as described by Smith et al. [41],
was not found in the material.
Distribution outside Europe. Known from a few localities in North Africa, e.g., in Morocco
[357].
Habitat and distribution in Europe. According to Purvis et al. [40], Opegrapha saxicola is
a Central European and Mediterranean species. It is found on limestone or calcareous schist,
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
100
D
B
C
90
C
C
B
80
70
C
B
B
%
60
B
50
A
B
40
A
30
A
B
20
A
A
4
5
10
A
A
0
1
2
3
6
7
Fig. 59 Variation in selected morphological and anatomical features Opegrapha
saxicola. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata:
A – unbranched, B – branched; 3 – disc type: A – concave, B – flat, C – convex;
4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very numerous;
7 – type of pycnidia: A – sessile, B – convex.
and rarely on sandstone or shale/slate. It is known from Central, Southern, and Eastern
Europe, and has been recorded from Austria [358] and Bosnia and Herzegovina [359].
Habitat and distribution in Poland. Opegrapha saxicola is found in southern Poland, on
limestone or occasionally on other rocks. Its first record was published by Błoński (as O. rupestris
[360]). It was reported from the Beskid Mały Mountains [187], Stołowe Mountains [198],
Świętokrzyskie Mountains [105,201,361], Kotlina Orawsko-Nowotarska (Orava-Nowy Targ
Basin) [362], Łańcuch Tatrzański macroregion [363–365], Pieniny Mountains [110,366],
Pradolina Wrocławska (Wrocław Urstromtal) [238], Eastern Sudetes [367], Wyżyna KrakowskoCzęstochowska (Kraków-Częstochowa Upland) [278], Wysoczyzna Bełchatowska (Bełchatów
Highland) [268] and Wyżyna Wieluńska (Wieluń Upland) [280].
The species occurs in the western part of the Polish Carpathians and in the Wyżyna Krakowska upland. It also is recorded from the Sudety Mountains (Fig. 60).
4.6.13. Opegrapha vermicellifera (Kunze) J. R. Laundon
Lichenologist 2(2): 139 (1963).
Pyrenothea vermicellifera Kunze, in Reichenbach. & Schubert, Lich. Exsicc. No. 60 (1823).
Thallus thin, well developed, white-grey to dark grey, usually sterile. Ascomata very rare,
unbranched or branched, usually sessile, 115–118 × 76–375 µm. Discs of ascomata narrow
or wider, often concave, not pruinose, 11–264 µm wide. Excipulum black, slightly elevated,
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Fig. 60 Distribution and substrate preference of Opegrapha saxicola in Poland.
continuous below the hypothecium, 29–40 µm thick laterally and 40–50 µm at the base. Hypothecium brownish, 10–15 µm high. Hymenium hyaline, ca. 60 µm high. Pseudoepithecium
very poorly developed. Asci club-shaped, 40–50 × 12–15 µm, inner layer of endoascus amyloid
nearly along its whole length (vulgata-type). Ascospores hyaline, straight, spindle-shaped,
five–six-septate, 16–24 × 3–5 µm, with perispore of 0.8–1.0 µm thick. Pycnidia numerous,
very characteristic whitish pruinose. Conidia straight, widened in the middle part, 3–4 ×
1 µm (Fig. 61). Chemistry: thalli K−, C−, PD−.
Discussion. Opegrapha vermicellifera is a very characteristic, epiphytic species that has a large,
whitish pruinose and nearly spherical pycnidia. The pycnidia help to distinguish it from the
closely related species (O. vulgata, O. niveoatra, and O. lithyrga). This diagnostic feature is
particularly useful for species identification in its sterile form and in material mixed with other
taxa. Additionally, O. vermicellifera differs from the above mentioned Opegrapha spp. due to
the presence of conidia that are widened in the middle part and only sometimes elongate.
The species varies remarkably in size and shape of ascomata. Nowak [10] accepted five
forms of this species: for. vermicellifera, for. discrepans (Erichs) J. Nowak, for. deplanata (Erichs)
J. Nowak, for. subpulicaris (Erichs) J. Nowak, and for. spermogonifera (Arnold?). The species is
found usually in sterile form (for. spermogonifera) with numerous, convex, whitish pruinose
pycnidia (Fig. 62). Specimens producing the ascomata are infrequent, and ascomata are elliptic, elongate, unbranched, and branched with a convex or rarely concave or flat disc.
Nomenclatural notes. This species was described in 1823 as Pyrenothea vermicellifera [368],
and Laundon referred to it as Opegrapha vermicellifera in 1963 [369].
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
Fig. 61 Opegrapha vermicellifera (Kunze) J. R. Laundon: (A–C) thallus and ascomata, LM
[(A) leg. A. Wieczorek, 2001, herb. Wieczorek; (B) leg. J. Nowak, KRAM-L 19963; (C) leg.
A. Wieczorek, 1995, herb. Wieczorek]; (D–F) thallus and ascomata, SEM [(D,F) leg. A. Wieczorek, 2001, herb. Wieczorek; (E) leg. A. Wieczorek, 1997, herb. Wieczorek); (G) ascus, SEM
(leg. A. Wieczorek, 1997, herb. Wieczorek); (H) ascospore, SEM (leg. A. Wieczorek, 1997,
herb. Wieczorek); (I) six-septate ascospore, LM (leg. A. Wieczorek, 2001, herb. Wieczorek);
(J) pycnidia, SEM (leg. P. Czarnota, GPN 2946); (K) conidia, SEM (leg. P. Czarnota, GPN
2946); (L) conidia, LM (leg. P. Czarnota, GPN 2946). Scale bars: (A) 300 µm; (B) 400 µm;
(C,J) 200 µm; (D,F) 100 µm; (E) 250 µm; (G–J) 10 µm; (K) 1 µm; (L) 5 µm.
Distribution outside Europe. The species is recorded primarily from Europe but also from
Africa (Morocco) [65].
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], this
species is found in Central European, Mediterranean, and sub-Atlantic regions. It is found on
the bark of deciduous trees, including Carpinus betulus, Quercus sp., and Fraxinus excelsior.
It is less often found on conifers. As late as the 1980s, it has not been reported from Southern
Europe [10].
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100
D
90
B
C
C
80
C
B
B
C
70
%
60
50
B
40
B
A
B
30
B
A
A
20
A
10
A
0
A
1
A
2
3
4
5
6
7
Fig. 62 Variation in selected morphological and anatomical features Opegrapha
vermicellifera. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 –
ascomata: A – unbranched, B – branched; 3 – disc type: A – concave, B – flat,
C – convex; 4 – thallus color: A – white, B – grey, C – white-grey, D – brownish;
5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very
numerous; 7 – type of pycnidia: A – sessile, B – convex.
Currently it is known from, e.g., Austria [71], the Czech Republic [295], Denmark [122,163],
France [166], Germany [90,91,124–126,168,337], Great Britain [92], Ireland [94], Italy [74],
Latvia [172], Lithuania [173], the Netherlands [96], Norway and Sweden [128], Portugal
(Azores [98]), Slovenia [76], Spain [99], Switzerland [129], and Ukraine [100].
Habitat and distribution in Poland. Opegrapha vermicellifera clearly prefers extensive woodlands. It is found on the bark of deciduous trees, most often on Fagus sylvatica and Ulmus
spp. However, it is less often found on Quercus spp., Fraxinus excelsior, Acer spp., Carpinus
betulus, and occasionally it can be found on Aesculus hippocastanum, Populus spp., Pyrus
sp., and Robinia pseudacacia [369]. Its preferred habitats are moderately acidic (pH 4.9–5.6)
to neutral (pH 5.7–7.0) [38]. It is a typical skiophyte, hence it needs shaded conditions for
proper development, which is similar to O. atra or Zwackhia viridis [38].
The first record of O. vermicellifera was published by Eitner [370] and Migula [130].
The species was reported from: Dolny Ślask (Lower Silesia Province) [132], Świętokrzyskie
Mountains [199–201], Karkonosze Mountains [202], Mierzeja Wiślana (Vistula Spit) [215],
Niecka Włoszczowska (Włoszczowa Basin) [216], northeastern Poland [218], Pobrzeże
Szczecińskie (Szczecin Coastland) [80], Pojezierze Ełckie (Ełk Lakeland) [305], Pojezierze
Olsztyńskie (Olsztyn Lake District) [235,236], Pogórze Przemyskie (Przemyśl Foothills) [112],
Równina Bielska (Bielsk Plain) [55,243], Równina Drawska (Drawno Plain) [244], Równina
Kozienicka (Kozienice Plain) [245,246,371,372], Śląsk Opolski (Opole Silesia) [262], Uznam
and Wolin islands [354], West Bieszczady Mountains [373–376], West Carpathians [12],
Western Pomerania [56,377], Wybrzeże Słowińskie (Slovincian Coast) [378], Wysoczyzna
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Fig. 63 Distribution and substrate preference of Opegrapha vermicellifera in Poland.
Białostocka (Białystok Highland) [356], and Wzgórza Radomszczańskie (Radomsko Hills)
[216]. According to Nowak [10], this species was often misidentified and overlooked during
field research because of its usually sterile thalli.
The species is scattered in the lowlands, and it is rare in the uplands and mountains (Fig. 63).
In the mountains, it has been recorded only from the West Bieszczady Mountains and from
a few localities in the Western Tatras.
4.6.14. Opegrapha vulgata (Ach.) Ach.
Meth. Lich.: 20 (1803).
Lichen vulgatus Ach., Lichenogr. Suec. Prodr. 21 (1799) [1798].
= Opegrapha cinerea Chevall., J. Phys. Chim. Hist. Nat. Arts 94: 41 (1822).
= Opegrapha devulgata Nyl., Flora (Regensburg) 62: 358 (1879).
= Opegrapha niveoatra auct. polon. (according to Nowak 1983) [non (Borrer) J. R. Laundon].
Thallus thin, continuous or cracked, smooth or granular, white-grey to green-brown. Ascomata numerous, solitary or in small groups, straight or curved, rarely branched, 0.5–1.5 ×
0.15–0.25 mm. Discs narrow, very rarely widened, sunken, black, not pruinose. Excipulum
permanent, elevated, black, naked, continuous below the hypothecium, 20–45 µm thick laterally
and 30–70 µm at the base. Hypothecium dirty yellowish, 15–35 µm high. Hymenium hyaline,
40–60 µm high, I+ reddish. Pseudoepithecium poorly developed. Asci cylindrical-clavate,
mature 40–50 × 11–18 µm, young 30–40 × 10–13 µm, inner layer of endoascus amyloid
along its whole length, ring around small ocular chamber (vulgata-type). Ascospores hyaline,
spindle-shaped, usually straight or slightly curved, five–six-septate, 19–28 × 3–5 µm, with
thin perispore of 1 µm, sometimes inconspicuous. Pycnidia numerous, sunken, small, black.
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Fig. 64 Opegrapha vulgata (Ach.) Ach.: (A) thallus and ascomata, LM (leg.
J. Nowak, KRAM-L 16211); (B–D) thallus and ascomata, SEM (leg. J. Nowak,
KRAM-L 16211); (E) three-septate ascospore, LM (leg. J. Nowak, KRAML 7288); (F) six-septate ascospore, LM (leg. J. Nowak, KRAM-L 10012); (G)
ascospores, SEM (leg. J. Nowak, KRAM-L 7252); (H) pycnidium, SEM (leg.
J. Nowak, KRAM-L 16211); (I–L) conidia, LM (leg. J. Nowak, KRAM-L 7252).
Scale bars: (A) 200 µm; (B,D) 100 µm; (C) 50 µm; (E–G,I–L) 10 µm; (H) 30 µm.
Conidia highly variable, straight, or slightly or strongly curved, 9–15 × 0.9–1.5 µm (Fig. 64).
Chemistry: thalli K−, C−, PD−.
Discussion. Opegrapha vulgata is closely related to O. vermicellifera and O. niveoatra [7]. Its
long conidia and small, hardly visible, but not pruinose pycnidia help to separate it from the
two morphologically similar species.
In Opegrapha vulgata, inter- and intrapopulation variation is observed in size and shape
of the ascomata (Fig. 65). The variation is associated with diverse habitat conditions. The
conidia size and shape also are highly variable. In general, the species has long and strongly
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100
D
C
90
80
C
C
D
C
B
70
%
60
B
50
B
B
C
40
B
30
B
A
20
A
B
10
0
A
A
A
1
2
3
4
A
A
5
6
7
Fig. 65 Variation in selected morphological and anatomical features Opegrapha
vulgata. 1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata:
A – unbranched, B – branched; 3 – disc type: A – concave, B – flat, C – convex;
4 – thallus color: A – white, B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very numerous;
7 – type of pycnidia: A – sessile, B – convex.
curved conidia. However, sometimes highly variable conidia can be observed within the
same population: from straight to slightly or strongly curved. This also was noted by Purvis
et al. [40], who distinguished three types of conidia on the basis of their size and shape. In
this study, shape and size of conidia were used as the most reliable diagnostic features for
this species.
Nomenclatural notes. The species was described by Acharius [18] as Lichen vulgatus, and
it was moved to Opegrapha later by the some author [17]. In the eighteenth and nineteenth
century, the taxon was treated under various names (O. cinerea, O. confluens, O. devulgata)
because of the high variation in size and shape of the ascomata, ascospores, and conidia (e.g.,
[1]). To show this morphological variation, 22 heterotypic names of intraspecific taxa also
were introduced [1]. Nowak [10] accepted two species: O. devulgata and O. vulgata.
Distribution outside Europe. This species has a wide geographic range. It can be found in
the USA [135,136,138,139,379], the Republic of Khakassia in Asia [380], and Australia [381].
The taxon is frequent in oceanic climates in the Euro-Siberian and Mediterranean regions,
e.g., in Israel [148], Iran [382], and Morocco [65]. It rarely occurs in dry habitats [34].
Habitat and distribution in Europe. Wirth [38] and Wirth et al. [39] defined it as having
a Central European distribution where it can be found in lowland forests, in shaded habitats
on slightly acidic bark. Reported, e.g., from Austria [71], the Czech Republic [295], Denmark
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[122,163,383], Estonia [123], Finland [165], France [166], Germany [124,168,297], Great
Britain [92], Greece [384], Hungary [93], Ireland [73], Italy [74,171], Latvia [172], Lithuania
[173,174], Montenegro [176], the Netherlands [96], Norway [97], Portugal (Madeira, Azores
[75,177]), Romania [179], Slovenia [76,385], Spain [99], Sweden [97], Switzerland [129],
Turkey [153,154], and Ukraine [100].
Habitat and distribution in Poland. Opegrapha vulgata is found on the bark of both deciduous
and coniferous trees in Poland. In the lowlands, it has been found on Carpinus betulus, Fagus
sylvatica, Quercus spp., and Acer spp.; in the mountains, it is found on the bark of coniferous
trees: Abies alba, Picea abies, and Pinus spp. It grows on slightly acidic substrates, with pH
4.1–4.8 [38]. The first records of O. vulgata were published by Körber [21], Ohlert [77], Stein
[182], Eitner [303], and Lettau [78]. Later it was reported from the Beskid Makowski Mountains [186], Beskid Mały Mountains [187,321], Beskid Niski Mountains [188], Beskid Sądecki
Mountains [102,103,323], Beskid Śląski Mountains [189], Bory Tucholskie (Tuchola Forest)
[194], Dolny Śląsk (Lower Silesia Province) [132], Gorce Mountains [349], Świętokrzyskie
Mountains [105,201,361], Kotlina Biebrzańska (Biebrza Basin) [205], Łańcuch Tatrzański
macroregion [107,365], northeastern Poland [218], Pieniny Mountains [111,366], Płaskowyż
Nałęczowski (Nałęczów Plateau) [220], Pobrzeże Szczecińskie (Szczecin Coastland) [80],
Pogórze Spiskie (Spisz Foothills) [225], Pojezierze Ełckie (Ełk Lakeland) [305], Pojezierze
Kaszubskie (Kashubian Lake District) [232], Pojezierze Olsztyńskie (Olsztyn Lake District)
[235,353], Pradolina Wrocławska (Wrocław Urstromtal) [238], Roztocze [258,386], Równina
Augustowska (Augustów Plain) [239], Równina Drawska (Drawno Plain) [244], Równina
Kozienicka (Kozienice Plain) [246], Równina Piotrkowska (Piotrków Plain) [252,253], Równina
Słupska (Słupsk Plain) [86], Sudetes [260,367], Śląsk Opolski (Opole Silesia) [262], Uznam
and Wolin islands [354], West Beskids [131], West Bieszczady Mountains [309], Western
Pomerania [56,84], Wybrzeże Słowińskie (Slovincian Coast) [378], Wysoczyzna Białostocka
(Białystok Highland) [356], Wyżyna Wieluńska (Wieluń Upland) [280], Wyżyna Lubelska
(Lublin Upland) [87], and Wzgórza Radomszczańskie (Radomsko Hills) [216].
The species is frequent in various areas of Poland (Fig. 66). In the south, it is known in the
western part of the Polish Carpathians (mostly the Beskid Wysoki Mountains, Beskid Średni
Mountains, Tatras, and also East Beskids) and Kotlina Kłodzka (Kłodzko Basin). Outside of
the mountains, it is frequent in the lowlands [mostly in the Pojezierze Kaszubskie, Warmia
and Masuria, and Nizina Podlaska (Podlachian Lowland)]. It also is scattered in the northern
part of the Wyżyna Małopolska (Lesser Poland Upland).
4.6.15. *Phacographa glaucomaria (Nyl.) Hafellner
Bibliotheca Lichenol. 100: 102 (2009).
Lecidea glaucomaria Nyl., Bot. Notiser: 177 (1852).
Opegrapha glaucomaria (Nyl.) Källsten ex Hafellner, Bull. Soc. Linn. Provence 45: 227 (1994).
It is a lichenicolous nonlichenized fungus. It is parasitic on the host lichen thallus (Protoparmelia badia). Ascomata black, roundish, naked, usually elevated above host thalli, very rarely.
Discs black, narrow, rarely broad. Excipulum permanent, thick, black, continuous below
the hypothecium. Hypothecium grey to grey-brown, dark, 10−14 µm high,upper part of
hypothecium K+ yellowish. Hymenium 50–75 µm high, I+ reddish. Pseudoepithecium very
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Fig. 66 Distribution and substrate preference of Opegrapha vulgata in Poland.
poorly developed, brown. Asci broadly club-shaped, mature 60–75 × 18–30 µm. Ascospores
hyaline, ellipsoid, two–three-septate, 18–29 × 7–9 µm, darkening in herbarium specimens,
with perispore of 0.6−1.4 µm thick. Pycnidia elevated or sessile, black. Conidia straight, 4–6
× 1 µm (Fig. 67).
Discussion. Phacographa glaucomaria is characterized by its epruinose ascomata, with
exposed discs. It differs from Opegrapha dolomitica in its color reaction in the upper part of
the hypothecium (K+ yellowish) and its slightly wider ascospores. It differs from Arthonia
calcarea due to its much longer and wider ascospores, and the presence of a well-developed
perispore. It is difficult to assess its intraspecific variability due to too few herbarium specimens available. The two specimens studied were morphologically similar. Small differences
were noted in ascospore size, but they are typical for this taxon. This species was not treated
by Nowak [10].
Nomenclatural notes. The species was described by Nylander [387] as Lecidea glaucomaria.
It was moved to Opegrapha in 1994, and recently to Phacographa [388].
General distribution. It has been reported from Canada [66], Austria [335], Portugal (Madeira [75]), and Spain [99].
Habitat and distribution in Poland. In Poland, this parasitic fungus grows on the thalli
of Protoparmelia badia. It was recorded only from the High Tatras at the Mięguszowiecka
Przełęcz Wyżnia (Mięguszowiecka Wyżnia Pass) and from Mięguszowiecki Szczyt Czarny
(Mięguszowiecki Black Peak) [336] (Fig. 68).
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Fig. 67 Phacographa glaucomaria (Nyl.) Hafellner: (A,B) thallus and ascomata,
LM (leg. Arnold, July 21, 1887, herb. Seaward); (C,D) thallus and ascomata,
SEM; (E) ascus with ascospores, LM; (F) ascospore, LM (leg. A. Flakus, KRAML 52516).Scale bars: (A,B) 1 mm; (C) 50 µm; (D) 150 µm; (E) 40 µm; (F) 15 µm.
Fig. 68 Distribution of Phacographa glaucomaria in Poland.
4.6.16. Pseudoschismatomma rufescens (Pers.) Ertz & Tehler
in Ertz, Tehler, Irestedt, Frisch, Thor & Boom, Fungal Diversity 70: 42 (2015) [2014].
Opegrapha rufescens Pers., Ann. Bot. (Usteri) 1: 29 (1794).
= Opegrapha herpetica (Ach.) Ach., Meth. Lich. 23 (1803).
Thallus thin, often with dark prothallus, smooth or granular-nodular, uniform or delicately
wrinkled, from grey-whitish to brownish-grey to dark brown. Ascomata numerous, more or
less evenly distributed, solitary or in small groups, unbranched or branched, sunken, rarely
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Fig. 69 Pseudoschismatomma rufescens (Pers.) Ertz Tehler: (A) thallus and ascomata, LM (leg. W. Fałtynowicz, UGDA-L 2568); (B–E) thallus and ascomata, SEM
[(B) leg. W. Fałtynowicz, UGDA-L 2568; (C,D) leg. J. Nowak, KRAM-L 2051; (E)
leg. S. Cieśliński, 1989, KTC]; (F) ascus with ascospore, SEM (leg. S. Cieśliński,
1989, KTC); (G) ascospore, SEM (leg. D. Kubiak, 2003, OLTC); (H,I) ascospores,
LM (leg. P. Czarnota, GPN 1181/94); (J,K) pycnidium, SEM (leg. J. Kiszka, 1968,
KRAP); (L) conidium, LM (leg. P. Czarnota, GPN 1181/94). Scale bars: (A) 500 µm;
(B,C) 200 µm; (D,K) 100 µm; (E,H,I) 10 µm; (F,G,L) 5 µm.
convex, 0.4–1.2 × 0.1–0.3 mm. Discs narrow, rarely widened, sunken, black, not pruinose.
Excipulum permanent, thick, elevated, black, naked, continuous below the hypothecium,
20–40 µm thick laterally and 30–60 µm at the base. Hypothecium 10–25 µm high, I+ blue.
Hymenium ca. 35–70 µm high, I+ reddish. Asci club-shaped, mature 40–60 × 10–12 µm,
young 30–40 × 10–11 µm, inner layer of endoascus amyloid nearly along its whole length
(vulgata-type). Ascospores hyaline, spindle-shaped, straight or slightly curved, three-septate,
15–25 × 3–4 µm, without perispore. Pycnidia numerous, small, black. Conidia straight or
slightly curved, 4–7 × 0.8–2 µm (Fig. 69). Chemistry: thalli K−, C−, PD−.
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Discussion. Pseudoschismatomma rufescens differs from the otherwise similar Arthonia
atra, Alyxoria culmigena, and A. ochrocheila due to its narrower ascospores and the lack of
a perispore. The primary diagnostic features of Pseudoschismatomma rufescens are: (i) the
ascomata are relatively short and usually sunken or surrounded by the thallus, (ii) there is
a lack of a perispore, and (iii) it has narrow ascospores.
Pseudoschismatomma rufescens is morphologically variable. Two varieties were described:
var. rufescens (Pers.) Mudd and var. subocellata Ach., with a total of 10 forms [var. rufescens
for. rufescens, for. disparate (Ach.) Grumm., for. rubella (Pers.) Grumm., for. elegans (Borr.)
Grumm, for. arthonioidea (Schaer), for. herpetica (Ach.), for. fuscata (Turn. Ach.), for. stellaris
(Müll. Arg.), var. δ. subocellata (Ach.) Schaer., for. subocellata, for. albicans (Chev.) Grumm.],
which seem to be of little taxonomic value [10]. There is intraspecific variation in thallus color,
and size and shape of the ascomata (Fig. 70). The characters are of little diagnostic value, as
they change considerably with environmental factors. The size and shape of the conidia are
quite variable as well, with either straight or mixed conidia (straight and slightly curved)
occurring occasionally in the same population or on the same thallus.
100
C
90
C
C
80
D
B
B
70
B
60
%
B
50
C
B
40
30
A
A
A
A
20
B
10
0
A
A
2
3
4
5
6
7
Fig. 70 Variation in selected morphological and anatomical features of Pseudoschismatomma rufescens. 1 – ascomata shape: A – circular, B – elliptic, C – elongate;
2 – ascomata: A – unbranched, B – branched; 3 – disc type: A – concave, B – flat,
C – convex; 4 – thallus color: A – white, B – grey, C – white-grey, D – brownish;
5 – excipulum types; 6 – number of pycnidia: A – few, B – numerous, C – very
numerous; 7 – type of pycnidia: A – sessile, B – convex.
Nomenclatural notes. The species was described by Persoon [283] as Opegrapha rufescens.
Ertz et al. [6] placed it in Pseudoschismatomma and noted its close morphological and anatomical similarity to species of the family Roccellaceae, including the genera Lecanactis and
Schismatomma. This similarity was mentioned by Diederich et al. [36], who pointed out the
lack of a perispore in ascospores of the taxa.
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Distribution outside Europe. This species has a wide geographic range, being distributed in
China [144], Morocco [65], Mongolia [389], the USA [138,288,390], and Canada [66].
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], this species occurs in boreal Mediterranean and sub-Atlantic regions. Pseudoschismatomma rufescens
grows on the bark of Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, and Quercus spp.,
and occasionally on Abies spp. and Picea spp. It usually is found in natural broad-leaved forest stands, where it is sometimes seen on trees growing along rivers and streams. It prefers
moderately acidic to neutral substrates, with pH 4.9–7.0 [38].
This species has been reported from Austria [71,160], the Czech Republic [295], Denmark
[122,163], Estonia [123], Finland [165], France [166,167], Germany [124,170,297], Great Britain
[92], Greece [127], Hungary [93], Ireland [94], Italy [74], Latvia [172], Lithuania [173,174],
Malta [95], Montenegro [316], the Netherlands [96], Norway [97], Portugal (Madeira, Azory
[75,98]), Romania [178], Russia (Dagestan [333]), Slovenia [76], Spain [99], Sweden [97],
Switzerland [129], and Ukraine [100,180,181].
Habitat and distribution in Poland. Pseudoschismatomma rufescens occurs primarily in
extensive natural woodlands and sometimes on isolated trees. It prefers shaded and slightly
moist habitats. It is widespread (in some places common) in the lowlands and mountains to
the lower montane zone of Poland. It grows on the bark of deciduous trees (Carpinus betulus,
Fagus sylvatica, Fraxinus excelsior, Quercus spp., less often on Salix spp.), occasionally on
Abies alba, exceptionally on Picea abies.
The first records of Pseudoschismatomma rufescens were published by Körber [21], Ohlert
[77], Stein [182], Eitner [303,347], and Lettau [78]. It was reported from Galicia [391], the
Beskid Makowski Mountains [186], Beskid Mały Mountains [321], Beskid Niski Mountains
[188,322], Beskid Sądecki Mountains [103,323], Beskid Śląski Mountains [189,190], Beskidy
Lesiste Mountains [191], Bieszczady Mountains [193,376], Bory Tucholskie (Tuchola Forest)
[194], Dolina Dolnego Sanu (Lower San River Valley) [104,392], Dolina Konińska (Konin
Valley) [196], Dolina Wisły (Vistula Valley) [393,394], Dolny Śląsk (Lower Silesia Province)
[132], Gorce Mountains [349], Stołowe Mountains [198], Świętokrzyskie Mountains [199–201],
Kotlina Gorzowska (Gorzów Basin) [395], Kotlina Orawsko-Nowotarska (Orava-Nowy
Targ Basin) [362], Kotlina Sandomierska (Sandomierz Basin) [396,397], Kotlina Żywiecka
(Żywiec Basin) [207], Łańcuch Tatrzański macroregion [210,211], Masyw Ślęży (Ślęża Massif)
[213], Nizina Mazowiecka (Mazovian Lowland) [217], Padół Zamojski (Zamość Depression)
[223], Pieniny Mountains [110,111,398], Płaskowyż Nałęczowski (Nałęczów Plateau) [220],
Pobrzeże Kaszubskie (Kashubian Coastland) [79,221], Pobrzeże Szczecińskie (Szczecin
Coastland) [80,222], Podgórze Rzeszowskie (Rzeszów Foothills) [223], Pogórze Przemyskie
(Przemyśl Foothills) [112], Pogórze Rożnowsko-Ciężkowickie (Rożnowsko-Ciężkowickie
Foothills) [81], Pogórze Spiskie (Spisz Foothills) [225], Pojezierze Ełckie (Ełk Lakeland)
[305,399], Pojezierze Gnieźnieńskie (Gniezno Lake District) [229], Pojezierze Kaszubskie
(Kashubian Lake District) [232], Pojezierze Krajeńskie (Krajna Lake District) [233], Pojezierze
Olsztyńskie (Olsztyn Lake District) [235], northeastern Poland [218], Pradolina Wieprza
(Wieprz River Urstromtal) [237], Równina Drawska (Drawno Plain) [85,244], Roztocze
[255,256,258], Równina Augustowska (Augustów Plain) [239,241,307], Równina Bielska
(Bielsk Plain) [55,243], Równina Biłgorajska (Biłgorajska Plain) [308], Równina Drawska,
Równina Kozienicka (Kozienice Plain) [245,373,400], Równina Łęczyńsko-Włodawska
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(Łęczna-Włodawa Plain) [247], Równina Łukowska (Łuków Plain) [251], Równina Słupska
(Słupsk Plain) [86,401], Sudetes [260,261,367], Śląsk Opolski (Opole Silesia) [262], West
Beskids [131,190,191], Western Pomerania [264,402], Wybrzeże Słowińskie (Slovincian Coast)
[378], Wysoczyzna Białostocka (Białystok Highland) [272],Wysoczyzna Elbląska (Elbląg
Highland) [273], Wysoczyzna Kaliska (Kalisz Highland) [274,275], Wysoczyzna Łaska (Łask
Highland) [276], Wyżyna Lubelska (Lublin Upland) [87], Wyżyna Krakowsko-Częstochowska
(Kraków-Częstochowa Upland) [278], Wyżyna Sandomierska (Sandomierz Upland) [279],
and Zaklęsłość Sosnowicka (Sosnowica Depression) [282].
In comparison to other species of the genus Opegrapha s. l., P. rufescens is most widespread
in the Polish lowlands, with a majority of its known localities in the northern and eastern parts
of the country, mostly in Warmia, Masuria, the Nizina Podlaska (Podlachian Lowland), and
Wyżyna Lubelska. In southern Poland, it is found in the western part of the Polish Carpathians and in the East Beskids (Fig. 71).
Fig. 71 Distribution and substrate preference of Pseudoschismatomma rufescens in Poland.
4.6.17. Zwackhia sorediifera (P. James) Ertz
in Diederich, Ertz, Eichler, Cezanne, Boom, Fischer, Killmann, van den Broeck & Sérusiau, Bull. Soc. Nat.
Luemb. 113: 106 (2012).
Opegrapha sorediifera P. James, Lichenologist 2: 86 (1962).
Thallus thin, partly sunken, with characteristic orange soralia. Ascomata black, unbranched,
0.2–0.7 × 0.1–0.3 mm, not pruinose, elevated, very rarely sunken. Discs black, narrow, rarely
broad. Excipulum continuous below the hypothecium, 80−100 µm thick at the base, K+ olivegreen. Hypothecium grey to grey-brown, dark, 10−15 µm high. Hymenium pale, 80–110 µm
high, I+ reddish. Asci club-shaped, 60–100 × 16–21 µm. Ascospores hyaline, spindle-shaped,
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Fig. 72 Zwackhia sorediifera (P. James) Ertz: (A,B) thallus with soredia, LM (leg. A. Wieczorek, 2014, herb. Wieczorek); (C) thallus with soredia, SEM (leg. A. Wieczorek, 2014, Sept.
22, herb. Wieczorek); (D) apothecia, SEM (leg. A. Wieczorek, 2014, herb. Wieczorek); (E)
ascus with ascospore, LM (leg. A. Wieczorek, 2014, herb. Wieczorek); (F,G) ascospore, LM
(leg. A. Wieczorek, 2014, herb. Wieczorek); (H) pycnidium, LM (leg. A. Wieczorek, 2014,
herb. Wieczorek); (I) pycnidium, SEM (leg. A. Wieczorek, 2014, herb. Wieczorek). Scale
bars: (A,B,G) 2 mm; (C) 200 µm; (D) 30 µm; (E)15 µm; (F) 10 µm; (H) 40 µm; (I) 200 µm.
30–45 × 4–5 µm, 10–14-septate, with well-developed perispore of 0.5–0.9 µm thick. Pycnidia
elevated or sessile, black. Conidia straight, 4–6 × 0.5–0.8 µm (Fig. 72). Soredia K−, C− or +
reddish, PD−.
Discussion. Zwackhia sorediifera is often mistaken with Z. viridis. Although both species grow
together in mixed populations, they are relatively easy to distinguish. Zwackhia sorediifera
can be recognized due to its characteristic orange soralia, multicellular ascospores, and markedly thinner perispore. The minor diagnostic features of Z. sorediifera are the shorter and
slightly wider ascospores compared to Z. viridis. Because of its rarity in Poland, intraspecific
variability cannot be assessed.
Nomenclatural notes. Opegrapha sorediifera was described by James [403] based on specimens
collected near Middleton in Ireland. In Europe, this taxon had been overlooked until recently.
According to the latest taxonomic treatment, the species is now placed in the genus Zwackhia
[29]. Both species in this genus, Z. sorediifera and Z. viridis, differ from other species in the
genus Opegrapha s. l. by their multicellular spindle-shaped ascospores and vulgata-type asci,
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which are characterized by an inner layer of endoascus amyloid along most of its length and
a conspicuous ring with no apical nasse.
Distribution outside Europe. This species is known from North America, recorded, e.g.,
from Canada [66].
Habitat and distribution in Europe. Zwackhia sorediifera was reported, e.g., from the Azores
[404], Denmark [163], Estonia [123], France [405], Great Britain [92], Lithuania [173,174],
and Norway and Sweden [97].
Habitat and distribution in Poland. This is the first record of Z. sorediifera reported from
Poland. The species was found on the bark of Fraxinus excelsior and Carpinus betulus in
extensive woodlands, in the northwestern part of the country (Fig. 73).
Fig. 73 Distribution of Zwackhia sorediifera in Poland.
4.6.18. Zwackhia viridis (Ach.) Poetsch & Schied.
System. Aufzähl. Samenlos. Pflanzen (Krypt.): 186 (1872).
Opegrapha rubella var. viridis Ach., Methodus, Sectio prior Stockholmiæ: 22 (1803).
= Opegrapha viridis (Ach.) Behlen & Desberger, Naturgesch. und Beschreib. Deutsch Forst.-Kryptog.: 110
(1835).
Thallus thin, often with lines of dark prothallus, smooth or delicately cracked in some parts,
grey-brown. Ascomata numerous, solitary or in large groups, usually evenly distributed,
unbranched or branched, 0.4–5.0 × 0.1–0.4 mm. Discs flat or convex, black and not pruinose,
at early stages of development very narrow, but during maturation some of them widened.
Excipulum edge thick, permanent, elevated, black, naked, continuous below the hypothecium,
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Fig. 74 Zwackhia viridis (Ach.) Poetsch & Schied.: (A,B) thallus and ascomata, LM [(A) leg.
A. Wieczorek, 1997, herb. Wieczorek; (B) leg. W. Fałtynowicz & J. Miądlikowska UGDA-L 3496];
(C–F) thallus and ascomata, SEM [(C,E) leg. A. Wieczorek, July 12, 1997, herb. Wieczorek;
(D,F) leg. W. Fałtynowicz & J. Miądlikowska UGDA-L 3496]; (G) ascus with ascospores, LM
(leg. A. Wieczorek, 1997, herb. Wieczorek); (H,I) ascospores, LM (leg. A. Wieczorek, 1997, herb.
Wieczorek); (J,K) pycnidium, SEM (leg. W. Fałtynowicz & J. Miądlikowska, UGDA-L 3496);
(L) conidia, LM (leg. W. Fałtynowicz & J. Miądlikowska, UGDA-L 3496). Scale bars: (A,B) 200
(C–F) 200 µm; (G) 30 µm; (H,I) 20 µm; (J,K) 100 µm; (L) 10 µm.
20–60 µm thick laterally and 30–80 µm at the base. Hypothecium brownish, 10–35 µm
high. Hymenium hyaline, ca. 40–85 µm high, I+ reddish-yellow. Pseudoepithecium poorly
developed. Asci club-shaped, mature 50–90 × 15–25 µm, young 30–50 × 10–15 µm − inner
layer of endoascus amyloid nearly along its whole length, chamber small, with no apical nasse
(vulgata-type). Ascospores hyaline, narrowly spindle-shaped, straight or slightly curved,
(5)6−15-septate, 20–60 × 3–10 µm, with well-developed perispore of 1.5–2.0 µm thick. Pycnidia numerous, small, black. Conidia straight or slightly curved, 6–11 × 0.8–2 µm (Fig. 74).
Chemistry: thalli K± dark rusty-brownish, C−, PD−.
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100
C
D
90
80
D
70
B
C
60
B
C
B
%
C
50
40
C
30
20
A
B
B
B
0
1
2
A
B
10
A
A
3
4
A
A
5
6
7
Fig. 75 Variation in selected morphological and anatomical features Zwackhia viridis
1 – ascomata shape: A – circular, B – elliptic, C – elongate; 2 – ascomata: A – unbranched,
B – branched; 3 – disc type: A – concave, B – flat, C – convex; 4 – thallus color: A – white,
B – grey, C – white-grey, D – brownish; 5 – excipulum types; 6 – number of pycnidia:
A – few, B – numerous, C – very numerous; 7 – type of pycnidia: A – sessile, B – convex.
Discussion. Zwackhia viridis is easy to recognize due to its very long and narrow, multicellular ascospores and its well-developed hyaline perispore. Its thallus habit and ascomata
shape sometimes resemble Opegrapha vulgata and O. niveoatra, but the ascospore size and
structure generally differ in the latter species.
Similar to other species with wide geographic ranges, Zwackhia viridis shows high phenotypic
variability. Nowak [10] distinguished five forms of the species: for. viridis, for. albicans Kremp.
ex Redgr., for. perminuta Erichs. ex Redgr. in herb., for. subatra Erichs. ex Redgr. in herb., and
for. ferruginea (Kremp.) A. Zahlbr. The size and shape of ascomata are highly variable in this
species (Fig. 75), as is ascospore length (with a range from 20 µm to 60 µm), and the number
of cells in ascospores. The majority of specimens (76) have 10-septate ascospores, followed
by 47 specimens with 11-septate ascospores and 20 specimens with 12-septate ascospores.
The remaining specimens have 13–15-septate ascospores [34].
A recent study on the genetic diversity of seven populations of Z. viridis revealed high
intraspecific genetic variability in different regions of eastern Poland [406].
Nomenclatural notes. Zwackhia viridis was described by Acharius [17] in 1803, under the name
Opegrapha rubella var. viridis. In 1814, the species name O. viridis began being used widely
[3]. An old synonym for the species is Lecanactis involuta (Wallr.) A. Massal. [407]. In 1869,
O. viridis was transferred to Graphis by Branth and Rostrup [408]. The species was treated by
Poetsch and Schiedermayr [409], who due to its specific features, called it Zwackhia viridis,
adopting the generic epithet from Körber [21]. Research by Ertz and Tehler [7] confirmed the
genotypic differentiation of this taxon, and the genus Zwackhia was reinstated by the authors,
who considered the morphological and anatomical structure of its representatives.
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Distribution outside Europe. Zwackhia viridis has a wide geographic range, comprising Africa,
North America, and Asia. It has been recorded, e.g., from China [144], Hong Kong [146],
Korea [68], Taiwan [69], Morocco [65], Canada [66], and the USA [135,138,139,410].
Habitat and distribution in Europe. According to Wirth [38] and Wirth et al. [39], Zwackhia
viridis occurs in Central European, Mediterranean, and sub-Atlantic regions. In Europe, it
grows on the bark of deciduous trees, most often on Quercus spp., Fraxinus excelsior, Fagus
sylvatica, and Carpinus betulus. Occasionally it can be found on coniferous trees.
It has been recorded, e.g., from Austria [71], the Czech Republic [295], Denmark [122,163],
Estonia [411], Germany [168,412], Great Britain [92], Hungary [93], Latvia [172], Lithuania
[175,413], Romania [178,179], Sweden and Norway [128], Switzerland [129], and Ukraine
[100].
Habitat and distribution in Poland. Zwackhia viridis grows mostly on the bark of Fagus
sylvatica, Carpinus betulus, and Fraxinus excelsior, in well-preserved forests of Poland. It is
found less frequently on isolated trees or in open forests, such as on Betula spp., Acer spp.,
Quercus spp., or Populus spp. Occasionally, it is found on Abies alba [367]. Zwackhia viridis
grows on trees whose bark is moderately acidic to neutral, with pH 4.9–7.0. It grows in the
shade, preferring moist habitats [38].
The first records of Zwackhia viridis were published by Körber [21], Ohlert [77], Stein
[182], Eitner [303,347], and Lettau [78]. It was reported from Galicia [185], the Beskid Mały
Mountains [321], Beskid Niski Mountains [188,322], Beskid Sądecki Mountains [102,323],
Beskid Śląski Mountains [189,190], Beskidy Lesiste Mountains [191], Bory Tucholskie
(Tuchola Forest) [194,195], Dolina Dolnego Sanu (Lower San River Valley) [104,397],
Dolina Konińska (Konin Valley) [194], Dolina Wisły (Vistula Valley) [394], Dolny Śląsk
(Lower Silesia Province) [132], Gorce Mountains [349,414], Stołowe Mountains [198],
Świętokrzyskie Mountains [199–201], Kotlina Gorzowska (Gorzów Basin) [395], Kotlina
Orawsko-Nowotarska (Orava-Nowy Targ Basin) [362], Kotlina Żywiecka (Żywiec Basin)
[207], Łańcuch Tatrzański macroregion [210,211], Masyw Ślęży (Ślęża Massif) [213], Mierzeja Wiślana (Vistula Spit) [215], Niecka Włoszczowska (Włoszczowa Basin) [216]. Nizina
Mazowiecka (Mazovian Lowland) [217], Padół Zamojski (Zamość Depression) [223], Pieniny Mountains [110,111,219], Płaskowyż Nałęczowski (Nałęczów Plateau) [220], Pobrzeże
Kaszubskie (Kashubian Coastland) [83,221], Pobrzeże Szczecińskie (Szczecin Coastland)
[80,222], Podgórze Rzeszowskie (Rzeszów Foothills) [223], Pogórze Przemyskie (Przemyśl
Foothills) [112], Pogórze Rożnowsko-Ciężkowickie (Rożnowsko-Ciężkowickie Foothills)
[81], Pogórze Spiskie (Spisz Foothills) [225], Pojezierze Ełckie (Ełk Lakeland) [82,305,399],
Pojezierze Gnieźnieńskie (Gniezno Lake District) [227–229], Pojezierze Iławskie (Iława Lake
District) [231], Pojezierze Kaszubskie (Kashubian Lake District) [232], Pojezierze Krajeńskie
(Krajna Lake District) [233], Pojezierze Olsztyńskie (Olsztyn Lake District) [235,353],
northeastern Poland [218], Pradolina Wieprza (Wieprz River Urstromtal) [237], Równina
Łukowska (Łuków Plain) [251], Roztocze [255,256,258], Równina Augustowska (Augustów
Plain) [239,241,242,307], Równina Bielska (Bielsk Plain) [55,243], Równina Biłgorajska
(Biłgorajska Plain) [308], Równina Drawska (Drawno Plain) [85,244], Równina Kozienicka
(Kozienice Plain) [245,372,400], Równina Łęczyńsko-Włodawska (Łęczna-Włodawa Plain)
[247], Równina Łowicko-Błońska (Łowicz-Błonie Plain) [249], Równina Piotrkowska
(Piotrków Plain) [252,253], Równina Słupska (Słupsk Plain) [86], Sudetes [259–261,367],
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Śląsk Opolski (Opole Silesia) [262], West Beskids [131,190,191], West Bieszczady Mountains
[309,355,376], Western Pomerania [84,264,402], Wybrzeże Słowińskie (Slovincian Coast)
[378], Wysoczyzna Białostocka (Białystok Highland) [271,272,356], Wysoczyzna Elbląska
(Elbląg Highland) [273], Wysoczyzna Kaliska (Kalisz Highland) [274,275], Wysoczyzna Łaska
(Łask Highland) [276], Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa Upland)
[278], Wyżyna Lubelska (Lublin Upland) [87], Wyżyna Sandomierska (Sandomierz Upland)
[279], and Zaklęsłość Sosnowicka (Sosnowica Depression) [282].
Zwackhia viridis is one of the most frequent epiphytic species in Poland, and it is distributed
in both lowlands and uplands [e.g., the Pojezierze Pomorskie (Pomeranian Lake District),
Warmia and Masuria, Wyżyna Małopolska (Lesser Poland Upland), and Wyżyna Lubelska)
as well as in the mountains (Tatras, Beskids, and Bieszczady Mountains) (Fig. 76). Herbarium
specimen labels indicate that in the Tatras, it is frequent in both the lower and upper montane
belts, but it reaches the subalpine belt (altitudes of up to about 1,700 m).
Fig. 76 Distribution and substrate preference of Zwackhia viridis in Poland.
5. Summary of results and conclusions
■ About 1,400 specimens of the genus Opegrapha s. l. were examined from the Polish herbaria
(BDPA, KRA, KRAM, KRAP, KTC, LBL, LOD, OLTC, POZ, TRN, UGDA, WA, WRSL),
from private collections, and obtained during the author’s field research. Reference material
from several European herbaria also was studied (e.g., BILAS, GSU, NNSU, OHHI, WI).
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
■ In total, 18 species of the genus Opegrapha s. l. were found to occur in Poland, including,
eight of the genus Opegrapha s.s, four of the genus Alyxoria, two of the genus Arthonia,
two of the genus Zwackhia, one of the genus Gyrographa, one of the genus Phacographa,
and one of the genus Pseudoschismatomma.
■ Zwackhia sorediifera is reported from Poland for the first time.
■ An identification key for the species of the genus Opegrapha s. l. found in Poland was
compiled based on morphological and anatomical characters.
■ Most of the species in the genus Opegrapha s. l. in Poland are simple to identify. The only
exception is Arthonia calcarea, as its ascospores are of similar size and shape to those of
O. saxicola. However, A. calcarea differs from O. saxicola by having a narrow perispore
and different ascus structure.
■ Within the genus Alyxoria, the shape of the hymenial disc and ascomata varied widely, so
the characters do not seem to be good diagnostic features.
■ In the genus Opegrapha, conidia size and shape are the most reliable diagnostic characters
at the species level.
■ All species of genus Opegrapha s. l. grow on natural substrates in Poland.
■ Out of the 18 species, 10 are epiphytes that grow on the bark of various tree species, including: Alyxoria culmigena, A. ochrocheila, A. varia, Arthonia atra, Opegrapha niveoatra,
O. vermicellifera, O. vulgata, Pseudoschismatomma rufescens, Zwackhia sorediifera, and
Z. viridis.
■ The epilithic species of the genus Opegrapha s. l. are Alyxoria mougeotii, Arthonia calcarea,
Gyrographa gyrocara, Opegrapha dolomitica, O. lithyrga, and O. saxicola, and they occur
predominantly in the south of the country.
■ New regional records for the number of species extended our knowledge of their distribution in Poland.
■ Several species are rare in Poland and are known only from a few localities. These include
Alyxoria culmigena, A. mougeotii, A. ochrocheila, Arthonia calcarea, Opegrapha dolomitica,
O. geographicola, O. lithyrga, Phacographa glaucomaria, and Zwackhia sorediifera.
■ A large number of the examined specimens originated from the northeastern and southern parts of Poland, likely a consequence of the more intensive collecting activity there
compared to other areas.
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1. Appendix: list of examined herbarium specimens,
literature records, and list of surveyed localities
Alyxoria culmigena (Lib.) Ertz
Specimens examined. AB-84 – Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe
(Beech Hills): Szczecin Landscape Park, Puszcza Bukowa (Beech Forest), Fagus sylvatica,
forest section No. 240, July 1997, leg. A. Wieczorek (herb. Wieczorek). AC-48 – Pojezierze
Południowopomorskie (South Pomeranian Lake District), Równina Gorzowska (Gorzów
Plain): Puszcza Barlinecka (Barlinek Forest), Wilanów Nature Reserve, Populus spp., December
3, 1989, leg. L. Lipnicki (herb. Lipnicki).
Additional material examined. BELARUS: Gomel Region, Gomel District, Pribor forest,
52°22' N, 30°45' E, Vaccinium myrtillus-type old-growth pine forest, Pinus sp., October 10, 2012,
leg. A. Tsurykau 01382 (GSU); Kalinino forest, 52°15' N, 30°59' E, Pleurozium schreberi-type
pine forest, Pinus sp., October 2, 2012, leg. A. Tsurykau 01383 (GSU); Kalinino forest, 52°15'
N, 30°59' E, Pleurozium schreberi-type pine forest, Pinus sp., August 3, 2011, leg. A. Tsurykau,
01439 (GSU); Kalinino forest, 52°15' N, 30°58' E, Pteridium aquilinum-type pine forest, Pinus
sp., August 3, 2011, leg. A. Tsurykau 01440 (GSU). FRANCE: forêt de Hardelot, Quercus sp.,
August 29, 1954, leg. J. Barkman. RUSSIA: Orel region, T. 204, Khotynetsky distr., Orlovskoye
Polesie National Park, L’govskoye Forest Division, dried wood, 53°17.249' N, 35°20.544' E,
alt. 239 m, June 9, 2012, leg. E. E. Muchnik 9635 (OHHI).
Alyxoria mougeotii (A. Massal.) Ertz, Frisch & G. Thor
Specimens examined. DF-48 – Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa
Upland), Wyżyna Olkuska (Olkusz Upland): Ojców village, Dolina Sąspowska (Sąspowska
Valley), limestone, May 1, 1956, leg. J. Nowak (KRAM-L 4128). DF-58 – Wyżyna KrakowskoCzęstochowska (Kraków-Częstochowa Upland), Wyżyna Olkuska (Olkusz Upland): Kochanów
village near Rudawa village, limestone, August 6, 1956, leg. J. Nowak (KRAM-L 4132); Ojców
National Park, Dolina Prądnika (Prądnik River Valley), rock, May 3, 2005, leg. J. Kiszka
(KRAP); Dolina Sąspowska (Sąspowska Valley), Jarki Gorge, rock, September 29, 2005, leg.
J. Kiszka (KRAP). EE-82 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
Polichno village, limestone, April 10, 1977, leg. Sepski (KTC). EG-32 – Obniżenie OrawskoPodhalańskie (Orava-Podhale Depression), Pieniny Mountains: rocks in Czorsztyn village,
limestone, October 11, 1993, leg. J. Kiszka (KRAP); Zielone Skałki, on N from Falsztyn
village, limestone, August 18, 1995, leg. J. Kiszka (KRAM-L 60008). EG-33 – Obniżenie
Orawsko-Podhalańskie (Orava-Podhale Depression), Pieniny Mountains: Wąwóz Szopczański
[Sobczański] (Szopczański Gorge), limestone, September 11, 1957, leg. K. Glanc (KRAML 39372); Wąwóz Szopczański [Sobczański] (Szopczański Gorge), alt. 560 m, limestone,
September 18, 1957, leg. Z. Tobolewski (POZ); limestone, August 6, 1954, leg. Z. Tobolewski
(POZ). EG-36 – West Beskids, Beskid Sądecki Mountains: Pasmo Jaworzyny Krynickiej (Jaworzyna Krynicka Range), sandstone, August 23, 1985, leg. A. Wieczorek (herb. Wieczorek).
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FF-98 – Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie
(Przemyśl Foothills): Krzeczkowa village, rock, stones, May 22, 1986, leg. J. Piórecki (BDPA).
FF-99 − Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie
(Przemyśl Foothills): Przemyśl City, on the way to Kruhel Wielki village, No. 127, concrete
wall, October 16, 1996, leg. J. Kiszka (KRAM-L 60007).
Additional material examined. GREAT BRITAIN: Chudleigh cave, S Devon, on basic rock
in deep shade, August 15, 1975, leg. C. J. B. Hitch (herb. Seaward).
Alyxoria ochrocheila (Nyl.) Ertz & Tehler
Specimens examined. DG-24 – West Beskids, Beskid Żywiecki Mountains: Rysianka
Mount, Fagus sylvatica, July 3, 1978, leg. J. Kiszka & J. Nowak (KRAM-L 17736); Hala
Rysianka, Fagus sylvatica, September 7, 1964, leg. J. Nowak (KRAM-L 17579). EE-09 − Nizina
Środkowomazowiecka (Central Mazovian Lowland), Równina Kozienicka (Kozienice Plain):
Puszcza Kozienicka (Kozienice Forest), Jedlnia Nature Reserve, Quercus sp., September 1,
2003, leg. S. Cieśliński (KTC). EE-60 – Wyżyna Przedborska (Przedborska Upland), Wzgórza
Łopuszańskie (Łopuszańskie Hills), Oleszno Nature Reserve, forest section No. 53, Acer
pseudoplatanus, February 2008, leg. A. Donica (KTC). FE-75 – Wyżyna Lubelska (Lublin
Upland), Wzniesienia Urzędowskie (Urzędów Hills): Trzydnik village near Kraśnik village,
Carpinus betulus, 1975, leg. Z. Flisińska (LBL).
Additional material examined. DENMARK: Bornholm, Dondalen, forest, Carpinus betulus,
July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek). GERMANY: SchleswigHolstein, Ostholstein, Fehmarn, Staberhuk, Staber Holz, Sambucus nigra, October 30,
2010, leg. U. Schiefelbein 2468 (herb. Schiefelbein); Mecklenburg-Vorpommern, Landkreis
Mecklenburg-Strelitz, Schlosspark, Mirow, concrete, November 2, 2002, leg. U. Schiefelbein
1213 (herb. Schiefelbein); Vorpommern-Rügen, Isle of Rügen, forest Goor, ca. 1 km E of
Badehaus, alt. 15 m, Carpinus betulus, October 24, 2014, leg. U. Schiefelbein 3732 (herb.
Schiefelbein); Niedersachsen, Landkreis Cloppenburg, NSG Urwald Baumweg, Carpinus
betulus, October 25, 2004, leg. U. Schiefelbein 1887 (herb. Schiefelbein).
Alyxoria varia (Pers.) Ertz & Tehler
Specimens examined. BA-59 – Pobrzeże Koszalińskie (Koszalin Coastland), Wybrzeże
Słowińskie (Slovincian Coast): Orzechowo village near Ustka town, Fagus sylvatica, July 17,
1986, leg. W. Fałtynowicz (UGDA-L 2824) as O. variaeformis. BA-98 − Pobrzeże Koszalińskie
(Koszalin Coastland), Równina Słupska (Słupsk Plain): Chomiec village near Janiewice village,
Fagus sylvatica, August 27, 1987, leg. W. Fałtynowicz & J. Miądlikowska (UGDA-L 3893) as
O. variaeformis. BB-23 – Pobrzeże Koszalińskie (Koszalin Coastland), Równina Białogardzka
(Białogard Plain): Krępa village near Białogard town, Acer sp., October 17, 1987, leg.
W. Fałtynowicz (KRAM-L 21815, UGDA-L 3602) as O. variaeformis. BB-59− Pojezierze
Południowopomorskie (South Pomeranian Lake District), Dolina Gwdy (Gwda River Valley);
Cisy w Czarnem Nature Reserve, beech forest, Fagus sylvatica, November 13, 2004, leg.
M. Kukwa 3709 (UGDA-L 12641). BC-12 − Pojezierze Południowopomorskie (South Pomeranian Lake District), Równina Drawska (Drawno Plain): Drawieński Park Narodowy
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(Drawa National Park), Tragankowe Urwisko Nature Reserve, E slopes of Drawa River, ca.
2 km NE of Zatom village, Fagus sylvatica, May 6, 2010, leg. U. Schiefelbein 2617, 2625 (herb.
Schiefelbein). BC-22 – Pojezierze Południowopomorskie (South Pomeranian Lake District),
Równina Drawska (Drawno Plain): Drawieński Park Narodowy (Drawa National Park),
Ostrowiec Lake, Okrzei Island, Fagus sylvatica, August 17, 1989, September 18, 1989, leg.
L. Lipnicki (herb. Lipnicki); decaying wood, May 27, 1989, leg. L. Lipnicki (herb. Lipnicki);
Województwo lubuskie (Lubusz Province), Acer pseudoplatanus, May 7, 2010, leg. U. Schiefelbein 2670 (herb. Schiefelbein); Dąbrowa village, ca. 2 km E of Moczele, Populus tremula, May
7, 2010, leg. U. Schiefelbein 2642 (herb. Schiefelbein); Wydrzy Głaz Nature Reserve, ca. 3 km
N of Moczele village, Fagus sylvatica, May 8, 2010, leg. U. Schiefelbein 2721 (herb. Schiefelbein).
BC-32 – Pojezierze Południowopomorskie (South Pomeranian Lake District), Równina
Drawska (Drawno Plain): Drawieński Park Narodowy (Drawa National Park), by Płociczna
River, forest section No. 253, Carpinus betulus, August 17, 1989, leg. L. Lipnicki (herb. Lipnicki).
BC-46 – Pradolina Toruńsko-Eberswaldzka (Toruń-Eberswalder Urstromtal), Kotlina Gorzowska (Gorzów Basin): Noteć region, near Czarnków town, Ulmus laevis, March 12, 1967,
leg. K. Glanc (KRAM-L 39090). BD-96 – Obniżenie Milicko-Głogowskie (Milicz-Głogów
Depression), Kotlina Żmigrodzka (Żmigród Basin): Kowalowo village, Populus sp., May 21,
1969, leg. K. Glanc (KRAM-L 39069). BF-37 – Eastern Sudetes, Góry Złote (Golden Mountains): Lądek Zdrój town, decaying wood, July 23, 1997, leg. E. Kozioł (WRSL). BF-47 – Eastern
Sudetes, Masyw Śnieżnika (Śnieżnik Massif): Bialskie Mountains, Dolina Kleśnicy (valley of
Kleśnica Stream), Acer sp., June 30, 2001, leg. K. Szczepańska (herb. Szczepańska); Puszcza
Jaworowa Nature Reserve, Fagus sylvatica, July 8, 2003, leg. K. Szczepańska (herb. Szczepańska);
Acer sp., July 8, 2003, leg. K. Szczepańska (herb. Szczepańska). BF-48 – Eastern Sudetes, Góry
Złote (Golden Mountains): Nowa Morawa village, roadside trees, September 18, 1992, leg.
E. Kozioł (WRSL); above Bielice village, old tree, September 20, 1991, leg. E. Kozioł (WRSL).
CA-36 – Pobrzeże Koszalińskie (Koszalin Coastland), Wybrzeże Słowińskie (Slovincian
Coast): Białogóra village, forest section No. 25Z, Fagus sylvatica, July 29, 1982, leg. W. Fałtynowicz
(UGDA-L 4236) as O. variaeformis. CA-37 − Pobrzeże Koszalińskie (Koszalin Coastland),
Pradolina Redy-Łeby (Reda-Łeba Urstromtal): ca. 2 km of Wejherowo town, near road to
Krokowa village, Acer sp., April 16, 1983, leg. A. Zalewska (UGDA-L 2880) as O. variaeformis.
CA-51 – Pobrzeże Koszalińskie (Koszalin Coastland), Wysoczyzna Damnicka (Damnica
Highland): Smołdzino village, March 19, 1995, leg. W. Fałtynowicz (herb. Seaward). CA-58 –
Pobrzeże Koszalińskie (Koszalin Coastland), Pradolina Redy-Łeby (Reda-Łeba Urstromtal):
ca. 4 km N of Wejherowo town, Ulmus sp., July 15, 1983, leg. A. Zalewska (UGDA-L 2869).
CA-66 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze
Kaszubskie (Kashubian Lake District): 1.5 km S of Porzecze village, Carpinus betulus, July
11, 1985, leg. W. Fałtynowicz (UGDA-L 2871) as O. variaeformis. CA-69 – Pojezierze
Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian
Lake District): between Rumia and Zagórze towns, Quercus sp., March 28, 1987, leg.
W. Fałtynowicz (UGDA-L 3241) as O. variaeformis. CA-86 – Pojezierze Wschodniopomorskie
(East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): near
Staniszewskie Błota Nature Reserve, Acer sp., April 23, 1977, leg. W. Fałtynowicz (UGDAL 1849, 1861); Fagus sylvatica, August 7, 1984, leg. W. Fałtynowicz (UGDA-L 2563) as
O. variaeformis; May 9, 2013, leg. M. Kukwa 12175, A. Kowalewska (UGDA-L 19348). CA-89
− Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie
(Kashubian Lake District): Gdańsk Oliwa City, Dolina Radości (Valley of Joy), Fagus sylvatica,
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June 7, 1963, leg. W. Fałtynowicz (UGDA-L 3384) as O. variaeformis. CB-04 – Pojezierze
Zachodniopomorskie (West Pomeranian Lake District), Pojezierze Bytowskie (Bytowskie
Lake District): Glinowo Forest District, SW edge of Matusz Lake, Fagus sylvatica, April 3,
1986, leg. W. Fałtynowicz (UGDA-L 3418). CB-09 – Pojezierze Wschodniopomorskie (East
Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Gdańsk City,
Zaskoczyn near Kościerzyna town, Aesculus hippocastanum, April 5, 1986, leg. W. Fałtynowicz
& Z. Tobolewski (KRAM-L 34074, UGDA-L 3014) as O. variaeformis. CB-24 – Pojezierze
Południowopomorskie (South Pomeranian Lake District), Równina Charzykowska (Charzykowy Plain): Bory Tucholskie (Tuchola Forest), Przymuszewo Forest District, Acer sp., July
28, 1976, leg. W. Fałtynowicz (UGDA-L 248); Parzyn village, Acer sp., June 17, 1975, leg.
W. Fałtynowicz (UGDA-L 605). CB-33 – Pojezierze Południowopomorskie (South Pomeranian
Lake District), Równina Charzykowska (Charzykowy Plain): Bory Tucholskie (Tuchola Forest), Laska village, Aesculus hippocastanum, August 2, 1983, leg. L. Lipnicki (TRN); July 6,
1985, leg. L. Lipnicki (herb. Lipnicki); Laska Forest District, forest section No. 766, Acer sp.,
March 27, 1976, leg. W. Fałtynowicz (UGDA-L 621). CB-68 – Pojezierze Południowopomorskie
(South Pomeranian Lake District), Bory Tucholskie (Tuchola Forest): stream below Cieściernicka
meadow, open place in oak-hornbeam forest, Quercus sp., June 14, 2004, leg. M. Kukwa 3325
(UGDA-L 11802). CB-75 – Pojezierze Południowopomorskie (South Pomeranian Lake
District), Równina Charzykowska (Charzykowy Plain): Bory Tucholskie (Tuchola Forest),
Tuchola Landscape Park, Brda River Valley Nature Reserve, Quercus sp., March 30, 1985, leg.
L. Lipnicki (herb. Lipnicki, TRN); Brda River valley, Carpinus betulus, August 11, 1989, leg.
L. Lipnicki (herb. Lipnicki); Corylus avellana, August 11, 1989, leg. L. Lipnicki (herb. Lipnicki).
CC-70 – Pojezierze Wielkopolskie (Wielkopolska Lake District), Pojezierze Gnieźnieńskie
(Gniezno Lake District): Zielonka Forest District near Poznań City, forest section No. 8,
Carpinus betulus, October 3, 1960, leg. K. Glanc (KRAM-L 39089); forest section No. 8g,
Quercus sp., August 11, 1963, leg. K. Glanc (KRAM-L 39093); Acer pseudoplatanus, August
11, 1963, leg. K. Glanc (KRAM-L 39099). CC-80 − Pojezierze Wielkopolskie (Wielkopolska
Lake District), Pojezierze Gnieźnieńskie (Gniezno Lake District): Zielonka Forest District
near Poznań City, forest section No. 27c, Quercus sp., September 18, 1961, leg. K. Glanc
(KRAM-L 39071); forest section No. 72c, Quercus sp., October 24, 1960, leg. K. Glanc (KRAML 39072); forest section No. 73, Acer sp., September 15, 1961, leg. K. Glanc (KRAM-L 39094);
Tilia sp., June 17, 1971, leg. K. Glanc (KRAM-L 39098); Ulmus sp., October 31, 1962, leg.
K. Glanc (KRAM-L 39100); Quercus sp., August 16, 1963, leg. K. Glanc (KRAM-L 39095);
forest section No. 73, Acer sp., June 17, 1961, leg. K. Glanc (KRAM-L 39097); forest section
No. 110f, Ulmus laevis, November 9, 1962, leg. K. Glanc (KRAM-L 39096); Ulmus sp., September 9, 1962, leg. K. Glanc (UGDA-L 5078) as O. variaeformis. DA-85 – Pobrzeże Gdańskie
(Gdańsk Coastland), Mierzeja Wiślana (Vistula Spit): Przebrno village, forest section No.
128, Betula sp., June 21, 1981, leg. E. Budzbon (UGDA-L 2189). DB-42 – Pojezierze
Południowobałtyckie (South Baltic Lakeland), Pojezierze Iławskie (Iława Lake District):
Kwidzyń Forest District, Populus tremula, April 10, 2004, leg. M. Kukwa 3026 (UGDA-L 10685).
DB-49 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn
Lake District): ca. 1.5 km S of Kotkowo village, Acer sp., August 22, 1993, leg. S. Cieśliński
(KTC); ca. 2 km S of Kot village, Acer sp., May 8, 1990, leg. S. Cieśliński (KTC). DB-53 –
Pojezierze Południowobałtyckie (South Baltic Lakeland), Pojezierze Iławskie (Iława Lake
District): Liwa River, Ulmus sp., April 12, 2004, leg. M. Kukwa 3056 (UGDA-L 10267).
DE-46 – Wyżyna Przedborska (Przedborska Upland), Wzgórza Radomszczańskie (Radomsko
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Hills): Fryszerka village by Widawka River, Ulmus sp., October 28, 1969, leg. K. Czyżewska
(LOD). DE-48 – Wyżyna Przedborska (Przedborska Upland), Wzgórza Radomszczańskie
(Radomsko Hills): Masłowice village, in a former manor house park, Ulmus sp., November
12, 1971, leg. K. Czyżewska (LOD). DE-78 – Wyżyna Przedborska (Przedborska Upland),
Niecka Włoszczowska (Włoszczowa Basin): Dębowiec Nature Reserve, forest section No.
161, Silniczka Forest District, Acer platanoides, June 15, 1972, leg. K. Czyżewska (LOD); forest
section No. 162, Ulmus glabra, September 30, 1969, leg. K. Czyżewska (LOD). DF-48 – Wyżyna
Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Wyżyna Olkuska (Olkusz
Upland): southern part, Ojców village, in Dolina Sąspowska (Sąspowska Valley), Carpinus
betulus, May 1, 1956, leg. J. Nowak (KRAM-L 4123). DF-58 – Wyżyna KrakowskoCzęstochowska (Kraków-Częstochowa Upland), Wyżyna Olkuska (Olkusz Upland): Dolina
Prądnika (Prądnik River Valley), rock, May 3, 2005, leg. J. Kiszka (KRAP). DF-68 – Wyżyna
Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Garb Tenczyński (Tenczynek
Ridge): Brzoskwinia village near Mników village, limestone, June 17, 1956, leg. J. Nowak
(KRAM-L 4127). DF-69 – Brama Krakowska, Pomost Krakowski: Kraków City, November
4, 1800, leg. A. Rehmann (KRAM-L 10316); Kraków City, Salix sp., leg. A. Rehman (KRAML 3194). DF-94 – Pogórze Zachodniobeskidzkie (West Beskids Foothills), Beskid Mały
Mountains: valley stream on S slope of Wielki Cisownik Mount, Fagus sylvatica, April 13,
1961, leg. J. Nowak (KRAM-L 7291); Zwalisko Mount, S slope, alt. 700 m, Fagus sylvatica,
May 11, 1960, leg. J. Nowak (KRAM-L 6222). DF-96 – Pogórze Zachodniobeskidzkie (West
Beskids Foothills), Beskid Mały Mountains: Targoszów village, 12 km to Sucha Beskidzka
town, Fagus sylvatica, May 11, 1960, leg. J. Nowak (KRAM-L 6223). DG-01 – West Beskids,
Beskid Śląski Mountains: in stream valley under Lipowski Wierch Mountain, Populus sp.,
August 3, 1964, leg. J. Kiszka (KRAP). DG-03 – West Beskids, Beskid Śląski Mountains:
Skrzyczne Mount, Fagus sylvatica, September 1, 1964, leg. J. Kiszka (KRAP). DG-04 – Pogórze
Zachodniobeskidzkie (West Beskids Foothills), Beskid Mały Mountains: Dolina Roztoki
(Roztoka Stream Valley), near Czernichowa village, Populus sp., August 24, 1960, leg. J. Nowak
(KRAM-L 6225); Kocierz Moszczanicki village, Fraxinus excelsior, August 8, 1960, leg. J. Nowak
(KRAM-L 7290). DG-05 – Pogórze Zachodniobeskidzkie (West Beskids Foothills), Beskid
Makowski Mountains: Pasmo Pewelskie (Pewelskie Range), Kapałów Stream, Abies alba, July
25, 1965, leg. J. Nowak (KRAM-L 15231). DG-07 – West Beskids, Beskid Żywiecki Mountains:
Przysłop Wielki Mount, Fagus sylvatica, August 6, 1964, leg. J. Nowak (KRAM-L 14398);
Beskid Makowski Mountains, Groń Mount, Fagus sylvatica, April 28, 1966, leg. J. Nowak
(KRAM-L 17343). DG-11 – West Beskids, Beskid Śląski Mountains: slope of Soszów Wielki
Mount, Fagus sylvatica, July 13, 1963, leg. J. Kiszka (KRAP). DG-12 – West Beskids, Beskid
Śląski Mountains: Barania Góra (Ram Mountain), E slope, Populus sp., August 31, 1965, leg.
J. Kiszka (KRAP); Biała Wisełka Stream, Fagus sylvatica, September 2, 1966, leg. J. Kiszka
(KRAP); Fagus sylvatica, July 24, 1965, leg. J. Kiszka (KRAP); Czarna Wisełka Stream, Fagus
sylvatica, June 24, 1965, leg. J. Kiszka (KRAP). DG-17 – West Beskids, Beskid Żywiecki
Mountains: Pasmo Policy (Polica Range), Urwanica Mount, Fagus sylvatica, July 9, 1965, leg.
J. Nowak (KRAM-L 15167); Okrąglica Mount, Fagus sylvatica, May 9, 1965, leg. J. Nowak
(KRAM-L 15175); Sidzińskie Pasionki, Fagus sylvatica, May 8, 1964, leg. J. Nowak (KRAML 16204). DG-22 – West Beskids, Beskid Śląski Mountains: Gańczorka Mount, Fagus sylvatica,
August 3, 1962, leg. J. Kiszka (KRAP). DG-24 – West Beskids, Beskid Żywiecki Mountains;
Pilsko Massif, Hala Bacmańska, Fagus sylvatica, September 9, 1964, leg. J. Nowak (KRAML 16594); Pilsko Massif, Pod Rysianką Nature Reserve, Fagus sylvatica, September 7, 1964,
© The Author(s) 2018
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leg. J. Nowak, (KRAM-L 16384); October 12, 1967, leg. J. Kiszka (KRAP 10060, 10083, 10057).
DG-32 – West Beskids, Beskid Żywiecki Mountains: Wielka Racza Massif, Beskid Bednarów
Mount, Fagus sylvatica, September 4, 1964, leg. J. Nowak (KRAM-L 14866, 14873); beech
forest, on N slope of Orzeł Mount, Fagus sylvatica, August 9, 1964, leg. J. Nowak (KRAML 14550); Przysłop Wielki Mount, Fagus sylvatica, August 6, 1964, leg. J. Nowak (KRAML 14069); Wielka Racza Massif, Śrubita Nature Reserve, Fagus sylvatica, August 12, 1964, leg.
J. Nowak, (KRAM-L 14004, 14005, 14449); Kikula Mount, Fagus sylvatica, June 5, 1976, leg.
U. Bielczyk (KRAM-L 42067); Wielka Racza Massif, on N slope of Orzeł Mount, Fagus sylvatica, August 9, 1964, leg. J. Nowak (KRAM-L 14551). DG-38 – Obniżenie OrawskoPodhalańskie (Orava-Podhale Depression), Kotlina Orawsko-Nowotarska (Orava-Nowy Targ
Basin): by Piekielnik Stream, Fagus sylvatica, May 30, 1970, leg. J. Kiszka (KRAP 10816);
Fagus sylvatica, June 20, 1970, leg. J. Kiszka (KRAP 10847). DG-39 – Obniżenie OrawskoPodhalańskie (Orava-Podhale Depression), Pogórze Spisko-Gubałowskie (Spisz-Gubałówka
Foothills): Skalice Nowotarskie, Tilia cordata, August 12, 1968, leg. J. Kiszka (KRAP 10529).
DG-49 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Rów Podtatrzański
(Podtatrzański Trench): Ligasowski Potok (Ligasowski Stream), Fraxinus excelsior, September
20, 1964, leg. J. Kiszka (KRAP); by Cichy Potok (Cichy Stream), Acer pseudoplatanus, August
20, 1964, leg. J. Kiszka (KRAP). DG-59 – Łańcuch Tatrzański Mountains, Western Tatras:
Dolina Strążyska (Strążyska Valley), near trail to Wielka Rówień under Giewont Mount, Acer
pseudoplatanus, August 18, 1959, leg. Z. Tobolewski (POZ). EB-02 – Nizina Staropruska (Old
Prussian Lowland), Wzniesienia Górowskie (Górowo Hills): NE of Stabunity village, Ulmus
sp., August 28, 1993, leg. S. Cieśliński (KTC). EB-22 – Pojezierze Mazurskie (Masurian Lake
District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 3 km NNE of Smolajny village,
near Łyna River, Fraxinus excelsior, August 23, 1993, leg. S. Cieśliński (KTC); ca. 1.5 km NNE
of Smolajny village, Fraxinus excelsior, August 23, 1993, leg. S. Cieśliński (KTC). EB-26 –
Pojezierze Mazurskie (Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake
District): W edge of Legińskie Lake, SW of Leginy village, Acer sp., May 3, 1989, leg. S. Cieśliński
(KTC). EB-32 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie
(Olsztyn Lake District): ca. 2 km SSW of railway station in Cerkiewnik village, Fraxinus
excelsior, September 14, 1989, leg. S. Cieśliński (KTC). EB-40 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): near road between villages
Pełnik and Wołowo, forest section No. 242, Acer sp., July 18, 1988, leg. S. Cieśliński (KTC).
EB-42 − Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn
Lake District): Olsztyn City, Osiedle Redykejany, Acer sp., August 2000, leg. D. Kubiak (OLTC);
Olsztyn City, city forest, Acer sp., August 9, 2000, leg. D. Kubiak (OLTC); Malus sp., April
2001, leg. D. Kubiak (OLTC). EB-45 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 1 km S of Rzeck village, by road to Rasząg
village, Acer sp., August 28, 1993, leg. S. Cieśliński (KTC). EB-49 − Pojezierze Mazurskie
(Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): Puszcza Piska
(Pisz Forest), ca. 1.5 km S of Lipowo village, Quercus sp., September 1, 1987, leg. S. Cieśliński
(KTC). EB-59 – Pojezierze Mazurskie (Masurian Lake District), Kraina Wielkich Jezior
Mazurskich: Masurian Landscape Park, 2 km SE of Bobrówko village Fraxinus excelsior,
November 9, 1999, leg. D. Kubiak (OLTC); ca. 1 km E of Krutyń village, Acer sp., September
2, 1987, leg. S. Cieśliński (KTC); Krutynia Nature Reserve, Quercus robur, August 31, 1987,
leg. S. Cieśliński (KTC). EB-62 − Pojezierze Mazurskie (Masurian Lake District), Pojezierze
Olsztyńskie (Olsztyn Lake District): Las Warmiński Nature Reserve on Łyna River, Fraxinus
© The Author(s) 2018
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excelsior, May 28, 1994, leg. S. Cieśliński (KTC). EB-68 – Pojezierze Mazurskie (Masurian
Lake District), Równina Mazurska (Masurian Plain): Szklarnia Forest District, ca. 4 km SSE
of Spychowo village, Acer sp., May 4, 1990, leg. S. Cieśliński (KTC). EC-87 – Nizina
Północnomazowiecka (North Mazovian Lowland), Dolina Dolnej Narwi (Lower Narew River
Valley): Pogorzelec village, May 21, 1950, leg. S. Cieśliński (LBL). ED-13 –Nizina
Środkowomazowiecka (Central Mazovian Lowland), Kotlina Warszawska (Warsaw Basin):
Kampinos National Park, Dąbrowa Stara village, Salix sp., July 1958, leg. J. Zielińska (WA).
ED-73 – Wzniesienia Południowomazowieckie (South Mazovian Hills), Wysoczyzna Rawska
(Rawa Highland): Trębaczew Nature Reserve, forest section No. 92, Carpinus betulus, August
19, 1970, leg. K. Czyżewska (LOD, KRAM-L 29964). ED-99 – Nizina Środkowomazowiecka
(Central Mazovian Lowland), Równina Kozienicka (Kozienice Plain): Puszcza Kozienicka
(Kozienice Forest), Zagożdżon Nature Reserve, Carpinus betulus, August 12, 1998, leg. Gnat
& S. Cieśliński (KTC). EE-00 – Wzniesienia Południowomazowieckie (South Mazovian
Hills), Równina Piotrkowska (Piotrków Plain): Puszcza Pilicka (Pilica Forest), Spała Nature
Reserve, forest section No. 271, Carpinus betulus, August 23, 1970, leg. K. Czyżewska
(LOD-L 521); forest section No. 287, Quercus robur, October 20, 1970, leg. K. Czyżewska
(LOD-L 522); forest section No. 287, Quercus robur, November 10, 1970, leg. K. Czyżewska
(LOD-L 524); Dolina Białobrzeska (Białobrzegi Valley): Puszcza Pilicka (Pilica Forest), near
Sługocice village, forest section No. 205l, Carpinus betulus, May 16, 1974, leg. K. Czyżewska
(LOD-L 4447). EE-09 – Nizina Środkowomazowiecka (Central Mazovian Lowland), Równina
Kozienicka (Kozienice Plain): Puszcza Kozienicka (Kozienice Forest), 1 km SW of Pionki
village, 1978, leg. A. Anusiewicz & S. Cieśliński (KTC). EE-32 – Wyżyna Przedborska (Przedborska Upland), Wzgórza Opoczyńskie (Opoczno Hills): near Trzemoszna village, Acer sp.,
July 20, 1979, leg. Kurczyńska & K. Toborowicz (KTC). EE-60 – Wyżyna Przedborska
(Przedborska Upland), Wzgórza Łopuszańskie (Łopuszańskie Hills): Oleszno Nature Reserve,
forest section No. 53, Acer platanoides, June 19, 1974, leg. K. Czyżewska (LOD). EE-73 –
Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains: Zagórsko village near Kielce
town, Ulmus sp., September 4, 1976, leg. K. Toborowicz (KTC). EE-82 – Wyżyna Kielecka
(Kielce Upland), Świętokrzyskie Mountains: Góra Czubatka Mount, Fraxinus excelsior, May
8, 1989, leg. S. Cieśliński (KTC). EE-97 – Wyżyna Kielecka (Kielce Upland), Pogórze Szydłowskie
(Szydłów Foothills): Przyjmy Forest District, forest section No. 113, 5 km E of Chańcza village,
Carpinus betulus, July 1986, leg. K. Toborowicz (KTC). EG-10 – West Beskids, Gorce Mountains: near church in Niedźwiedź village, Tilia sp., September 16, 1965, leg. K. Glanc (KRAML 39082); S of Niedźwiedź village, by Konina Stream, Populus sp., September 16, 1965, leg.
K. Glanc (KRAM-L 39084, 39086); in former manor house park in Poręba Wielka village,
November 23, 1993, leg. P. Czarnota (GPN 13); park near Poręba Wielka Forest District
headquarters, Fraxinus excelsior, Ulmus sp., September 16, 1965, leg. K. Glanc (KRAML 39050). EG-20 – West Beskids, Gorce Mountains: W of Turbacz Mount, by Lepietnica
Stream, Fagus sylvatica, September 12, 1964, leg. K. Glanc (KRAM-L 39049); valley of Olszowy
Potok (Olszowy Stream), Fagus sylvatica, May 6, 1993, leg. P. Czarnota (GPN 514); on E slope
of Suhora Mount, Fagus sylvatica, July 4, 1994, leg. P. Czarnota (GPN 592); Suche Buczki
Forest District, Fagus sylvatica, July 31, 1995, leg. P. Czarnota (GPN 1849). EG-21 – West
Beskids, Gorce Mountains: Turbacz Nature Reserve, above Turbaczyk Stream, Fagus sylvatica,
September 6, 1959, leg. K. Glanc (KRAM-L 39067); Dolina Kamienicy (Kamienica River
Valley), Fagus sylvatica, June 11, 1996, leg. P. Czarnota (GPN 1405); Turbacz Nature Reserve,
Fagus sylvatica, February 5, 1993, leg. P. Czarnota (GPN 267); s. loc., May 23, 1996, leg.
© The Author(s) 2018
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P. Czarnota (GPN 12/95/94); Konina Stream Valley, Acer sp., January 9, 1994, leg. P. Czarnota
(GPN 878); Ustępny Stream Valley, Fagus sylvatica, February 20, 1995, leg. P. Czarnota (GPN
251); Łopuszna Nature Reserve, Fagus sylvatica, May 5, 1995, leg. P. Czarnota (GPN 1041);
near plot 329, Fagus sylvatica, February 20, 1995, leg. P. Czarnota (GPN); under ridge of
Przysłopek Mount, in Konina Stream catchment area, Fagus sylvatica, January 4, 1996, leg.
P. Czarnota (GPN 854); S slope of Mostownica Mount, Fagus sylvatica, April 3, 1954, leg.
K. Glanc (KRAM-L 39066); near source of Lepietnica Stream, at the eit of Turbacz Mount,
Fagus sylvatica, September 2, 1964, leg. K. Glanc (KRAM-L 39088); near blue tourist trail,
by Kamienica Stream, near temporary building of Polish Academy of Sciences, Fagus sylvatica,
June 11, 1996, leg. P. Czarnota (GPN 967); Turbacz Nature Reserve, Czoło Turbacza Mount,
Fagus sylvatica, September 9, 1964, leg. K. Glanc (KRAM-L 39076); Fagus sylvatica, September
11, 1966, leg. K. Glanc (KRAM-L 39092); decaying wood, June 20, 1967, leg. K. Glanc (KRAML 39091); Fagus sylvatica, August 28, 1961, leg. K. Glanc (KRAM-L 39074); E slope of Kiczora
Mount, Dolina Furcówki (Furcówka Stream Valley), decaying wood, August 27, 1968, leg.
K. Glanc (KRAM-L 39080). EG-22 – West Beskids, Gorce Mountains: S slope of Twarogi
Mount, Fraxinus excelsior, July 8, 1966, leg. K. Glanc (KRAM-L 39085); N slope of Lubań
Mount, in the valley of Kudowski Stream Fagus sylvatica, July 3, 1967, leg. K. Glanc (KRAML 39075); Ochotnica Dolna village, under Kasinka Mount, by the road to Ochotnica Górna
village, Salix sp., August 7, 1966, leg. K. Glanc (KRAM-L 39057); N slope of Lubań Mount,
SE of Skała Boginka, Acer pseudoplatanus, August 15, 1967, leg. K. Glanc (KRAM-L 39081);
E slope of Lubań Mount, above Tylmanowa village, Salix sp., September 9, 1973, leg. K. Glanc
(KRAM-L 39497); S slope of Lubań Mount, Dolina Kluszkowianki (Kluszkowianka Stream
Valley), Fagus sylvatica, July 6, 1967, leg. K. Glanc (KRAM-L 39087); S slope of Lubań Mount,
valley of Skalny Potok (Rocky Stream), decaying wood, May 10, 1959, leg. K. Glanc (KRAML 39070); Lubań Mount, NE slope, near route from Rzek to Biała Skała, Fagus sylvatica, August
4, 1968, leg. K. Glanc (KRAM-L 39083). EG-25 – West Beskids, Beskid Sądecki Mountains:
Pasmo Jaworzyny (Jaworzyna Range), Barnowiec Nature Reserve, Fagus sylvatica, September
12, 1976, leg. U. Bielczyk (KRAM-L 42062). EG-26 – West Beskids, Beskid Sądecki Mountains:
Pasmo Jaworzyny (Jaworzyna Range), Łabowiec Nature Reserve, Fagus sylvatica, June 13,
1991, leg. L. Śliwa (KRA). EG-33 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Pieniny Mountains: near road from Krościenko to Zawiasy villages by Dunajec
River, Salix sp., August 12, 1954, leg. Z. Tobolewski (POZ); near road from Krościenko to
Źlebie Żłobina villages, Robinia pseudacacia, June 10, 1956, leg. Z. Tobolewski (POZ); near
trail from Polana Wyrobek to Zamkowa Góra (Zamkowa Mount), Fagus sylvatica, June 12,
1956, leg. Z. Tobolewski (POZ); roadside near Szlachtowa village, Salix sp., June 9, 1956, leg.
Z. Tobolewski (POZ). EG-34 – West Beskids, Beskid Sądecki Mountains: Pasmo Radziejowej
(Radziejowa Range), S slope of Dzwonkówka Mount, Fagus sylvatica, June 13, 1965, leg.
M. Olech (KRA); by Potok Sopotnicki (Sopotnicki Stream), sandstone, June 13, 1965, leg.
M. Olech (KRA); Szlachtowa village, Fraxinus excelsior, May 2, 1991, leg. L. Śliwa (KRA);
Kłodzkie Nature Reserve, Tilia cordata, July 12, 1967, leg. M. Olech (KRA); Carpinus betulus,
August 12, 1967, leg. M. Olech (KRA); Pasmo Jaworzyny (Jaworzyna Range), at edge of Polana
Bukowa, Fagus sylvatica, September 23, 1966, leg. M. Olech (KRA). EG-35 – West Beskids,
Beskid Sądecki Mountains: Pasmo Jaworzyny (Jaworzyna Range), Potasznia Stream Valley,
Fagus sylvatica, September 9, 1967, leg. M. Olech (KRA); sandstone, October 22, 1966, leg.
M. Olech (KRA); Fagus sylvatica, July 28, 1967, leg. M. Olech (KRA). EG-36 – West Beskids,
Beskid Sądecki Mountains: Pasmo Jaworzyny (Jaworzyna Range), Składniszczan Stream
© The Author(s) 2018
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Valley, Fraxinus excelsior, July 25, 1966, leg. M. Olech (KRA); Feleczyn Stream Valley, Fagus
sylvatica, October 9, 1967, leg. M. Olech (KRA); Żebracze Nature Reserve, Fagus sylvatica,
July 5, 2001, leg. P. Czarnota (GPN 2520); Fagus sylvatica, July 6, 2001, leg. P. Czarnota (GPN
2549). EG-40 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Pogórze
Spisko-Gubałowskie (Spisz-Gubałówka Foothills); Pasmo Bukowiny Tatrzańskiej, Betula sp.,
August 17, 1967, leg. J. Kiszka (KRAP). EG-41 – Obniżenie Orawsko-Podhalańskie (OravaPodhale Depression), Pogórze Spisko-Gubałowskie (Spisz-Gubałówka Foothills): NW slope
of Piastowy Wierch Mountain, Fagus sylvatica, June 20, 1970, leg. J. Kiszka (KRAP 10894).
EG-50 – Łańcuch Tatrzański Mountains, Eastern Tatras: Dolina Białego (Valley of the White),
Fagus sylvatica, July 13, 1912, leg. Augustynowicz (KRAM-L 18076). FA-84 – Pojezierze
Litewskie (Lithuanian Lake District), Puszcza Romincka (Romincka Forest): near road from
Jurkiszki to Szyliny villages, forest section No. 304, Betula sp., May 17, 1985, leg. Z. Tobolewski
(KRAM-L 31851); s. loc., Betula sp., May 16, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC);
forest section No. 88, Fraxinus excelsior, May 16, 1985, leg. S. Cieśliński (KRAM-L 31864);
2 km N of Gołdap town, Ulmus sp., May 20, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC);
s. loc., leg. S. Cieśliński (KRAM-L 31912). FA-85 – Pojezierze Litewskie (Lithuanian Lake
District), Puszcza Romincka (Romincka Forest): Boczki Nature Reserve, forest section No.
86, Tilia cordata, May 16, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC); 0.7 km SE of
Pluszkiejmy village, E edge of Czarne Lake, Salix sp., May 19, 1985, leg. Z. Tobolewski &
S. Cieśliński (KTC). FA-87 – Pojezierze Litewskie (Lithuanian Lake District), Puszcza Romincka
(Romincka Forest): ca. 2.5 km SW of Żytkiejmy village, Acer sp., May 15, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC). FB-13 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze
Olsztyńskie (Olsztyn Lake District): Puszcza Borecka (Borki Forest), Walisko Lake, Fraxinus
excelsior, May 13, 1958, leg. K. Glanc (KRAM-L 39052); Borki Nature Reserve, forest section
No. 20, Tilia cordata, May 19, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC); Acer sp., May
19, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC). FB-16 – Pojezierze Litewskie (Lithuanian
Lake District), Pojezierze Zachodniosuwalskie (West Suwałki Lake District): near road from
Bakałarzewo village to Olecko town, Acer sp., May 16, 1987, leg. Z. Tobolewski & S. Cieśliński
(KTC). FB-29 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska
(Augustów Plain): Monkinie Nature Reserve, Corylus avellana, August 4, 1993, leg. J. Bystrek
& A. Matwiejuk (University of Bialystok, Institute of Biology). FB-31 – Pojezierze Mazurskie
(Masurian Lake District), Kraina Wielkich Jezior Mazurskich (Great Masurian Lakes Land):
ca. 2 km SE of Dąbrówka village, near road to Orzysz town, Acer sp., September 6, 1988, leg.
S. Cieśliński (KTC). FB-38 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Ełckie
(Ełk Lakeland): Jeziorki village near road from Ełk to Augustów towns, Alnus glutinosa, May
23, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC). FB-51 – Pojezierze Mazurskie (Masurian
Lake District), Kraina Wielkich Jezior Mazurskich (Great Masurian Lakes Land): S edge of
Kocioł Lake, Acer sp., September 7, 1988, leg. S. Cieśliński (KTC). FB-61 – Pojezierze Mazurskie (Masurian Lake District), Równina Mazurska (Masurian Plain): ca. 7 km NE of Pisz
village, near Roś Lake, Acer sp., May 10, 1990, leg. S. Cieśliński (KTC). FB-68 – Nizina
Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin): Czerwone Bagno Nature Reserve, forest section No. 246, Quercus robur, September 24, 1986, leg.
S. Cieśliński (KTC). FC-68 – Nizina Północnopodlaska (North Podlachian Lowland), Równina
Bielska (Bielsk Plain): ca. 5 km W of Szmurły village, Quercus robur, August 20, 1991, leg.
S. Cieśliński (KTC). FD-45 – Wzniesienia Południowomazowieckie (South Mazovian Hills),
Równina Łukowska (Łuków Plain): Jata Nature Reserve near Łuków town, Acer sp., November
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4, 1971, leg. M. Motyka-Zgłobicka (LBL); Sorbus sp., May 25, 1973, leg. M. Motyka-Zgłobicka
(LBL). FD-68 – Nizina Południowopodlaska (South Podlachian Lowland), Równina Łukowska
(Łuków Plain): Omelno Nature Reserve near Radzyń Podlaski town, Quercus sp., August 3,
1972, leg. B. Ordyczyńska (LBL); Acer sp., August 3, 1972, leg. B. Ordyczyńska (LBL). FE-07 –
Nizina Południowopodlaska (South Podlachian Lowland), Wysoczyzna Lubartowska (Lubartów
Highland): Kozłówka Forest District, Dąbrówka Forest District, Nowodwór Forest District,
Betula sp., September 20, 1962, leg. L. Jędrzejewska (LBL); Nasutów Forest District, Populus
sp., September 12, 1962, leg. L. Jędrzejewska (LBL). FE-13 – Wyżyna Lubelska (Lublin Upland),
Małopolski Przełom Wisły (Lesser Poland Gorge of the Vistula): Puławy town, Quercus sp.,
1970, leg. H. Stasiak (LBL). FF-01 – Kotlina Sandomierska (Sandomierz Basin), Równina
Tarnobrzeska (Tarnobrzeg Plain): Tarnobrzeg town, Quercus sp., 1968, leg. J. Piórecki (LBL).
FF-36 – Kotlina Sandomierska (Sandomierz Basin), Płaskowyż Kolbuszowski (Kolbuszowa
Plateau): Brzóza Królewska Forest District, forest section No. 218, Acer sp., 1963, leg.
B. Ordyczyńska (LBL); Jelana Forest District, Fraxinus sp., 1963, leg. B. Ordyczyńska (LBL);
forest section No. 119, Sorbus sp., leg. B. Ordyczyńska (LBL). FF-37 – Kotlina Sandomierska
(Sandomierz Basin), Płaskowyż Kolbuszowski (Kolbuszowa Plateau): Leżajsk County, Puszcza
Sandomierska (Sandomierz Forest), Julin Forest District, forest section No. 253, Quercus sp.,
August 1960, leg. B. Ordyczyńska (LBL); forest section No. 271, Fraxinus sp., 1963, leg.
B. Ordyczyńska (LBL). FF-45 − Kotlina Sandomierska (Sandomierz Basin), Płaskowyż
Kolbuszowski (Kolbuszowa Plateau): Marynin Forest District, Acer sp., 1963, leg. B. Ordyczyńska
(LBL). FF-99 – Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills): Korytniki village, Morus sp., October 13, 1983, leg. J. Kiszka &
J. Piórecki (KRAM-L 30517). FG-13 − Central Beskids, Beskid Niski Mountains: near RymanówZdrój town, Wisłoczek Stream Valley, Fraxinus excelsior, August 20, 1974, leg. J. Nowak
(KRAM-L 33137). FG-14 − Central Beskids, Beskid Niski Mountains: Bukowica, grassland
near Odrzechowa village, Juglans regia, October 10, 1974, leg. J. Nowak (KRAM-L 32416).
FG-19 – Beskidy Lesiste Mountains, Sanocko-Turczyńskie Mountains: Pasmo Turnicy, under
Arłamów village, Acer pseudoplatanus, May 22, 1986, leg. J. Piórecki (BDPA), leg. J. Kiszka
& J. Piórecki (KRAM-L 60047). FG-20 − Central Beskids, Beskid Niski Mountains: Nieznajowa
near Świątkowa villages, Spełzły forest, No. 600, Abies alba, October 4, 1979, leg. J. Nowak
(KRAM-L 32441). FG-22 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Puszcza
Bukowa (Beech Forest), between Czerkownik and Szypowaty streams, Fagus sylvatica, August
22, 1957, leg. K. Glanc (KRAM-L 39061). FG-23 – Central Beskids, Beskid Niski Mountains:
valley stream, between Bania Mount and Daňová Mount, Fagus sylvatica, May 22, 1974, leg.
M. Olech (KRA). FG-24 − Central Beskids, Beskid Niski Mountains: near Huczkowa village,
on a clearing in the forest No. 415, Julans regia, September 4, 1977, leg. J. Nowak (KRAML 23014). FG-33 − Central Beskids, Beskid Niski Mountains: near Jaśliska village, between
Spalona and Stasiana farmsteads No. 410, Acer platanoides, September 11, 1974, leg. J. Nowak
(KRAM-L 32287). FG-34 − Central Beskids, Beskid Niski Mountains: Berdo village near
Moszczaniec village, by the Wisłok River, Fraxinus excelsior, September 16, 1974, leg. J. Nowak
(KRAM-L 32739). FG-45 – Bieszczady Lesiste Mountains, West Bieszczady Mountains:
S slope of Dział Mount, Acer pseudoplatanus, April 24, 1958, leg. K. Glanc (KRAM-L 39048).
FG-59 – Bieszczady Lesiste Mountains, West Bieszczady Mountains: Dwerniczek village,
near road by San River, Salix sp., August 15, 1958, leg. Z. Tobolewski (POZ). FG-68 – Beskidy
Lesiste Mountains, West Bieszczady Mountains: Puszcza Bukowa (Beech Forest), on S slope
of Dział Mount, Fagus sylvatica, August 19, 1958, leg. K. Glanc (KRAM-L 39064); Puszcza
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Bukowa (Beech Forest), Fagus sylvatica, August 22, 1957, leg. K. Glanc (KRAM-L 39058);
above Beskidnik Stream, Fagus sylvatica, August 28, 1957, leg. K. Glanc (KRAM-L 39060);
Lesko County, near Beskidnik Stream, Acer sp., August 28, 1957, leg. Z. Tobolewski (POZ);
Acer pseudoplatanus, August 1956, leg. Z. Tobolewski (KTC); Dział Mount, below the top of
the mountain, Acer pseudoplatanus, April 26, 1956, leg. K. Glanc (KRAM-L 39045); Fagus
sylvatica, April 26, 1956, leg. K. Glanc (KRAM-L 39055); by Wielki Lutowy Stream, Puszcza
Bukowa (Beech Forest), Fagus sylvatica, August 18, 1958, leg. K. Glanc (KRAM-L 39059);
Puszcza Bukowa (Beech Forest), near Wetlina village, by Wielki Lutowy Stream, Fagus sylvatica,
August 18, 1958, leg. Z. Tobolewski (POZ); slope on right bank of Wielki Lutowy Stream,
Fagus sylvatica, August 23, 1957, leg. Z. Tobolewski (KTC). FG-69 – Bieszczady Lesiste
Mountains, West Bieszczady Mountains: Ustrzyki Górne village, by road to Bereżki village,
Ulmus montana, August 5, 1959, leg. K. Glanc (KRAM-L 39053); Ustrzyki Górne village,
above Wołosaty Stream, Acer pseudoplatanus, July 8, 1960, leg. J. Nowak (KRAM-L 8821) as
O. variaeformis; Bereżki village, Acer pseudoplatanus, September 20, 1968, leg. K. Glanc
(KRAM-L 39046); valley of Rzeczyca Stream, near Ustrzyki Górne village, Acer pseudoplatanus,
August 27, 1957, leg. K. Glanc (KRAM-L 39056); valley of Rzeczyca Stream, above road from
Berehy (now: Brzegi Górne) village to Ustrzyki Górne village, Acer pseudoplatanus, September
27, 1958, leg. K. Glanc (KRAM-L 39047); between Caryńskie and Bereżki villages, Acer
pseudoplatanus, September 22, 1958, leg. K. Glanc (KRAM-L 39062); Bereżki village, road
to Przełęcz Przysłup (Przysłup Pass), Acer pseudoplatanus, July 17, 2001, leg. J. Kiszka &
R. Kościelniak (KRAP 1888). FG-79 – Beskidy Lesiste Mountains, West Bieszczady Mountains:
S slope of Tarnica Mount, above Wołosatka Stream Valley, Acer pseudoplatanus, June 1956,
leg. K. Glanc (KRAM-L 39063). GB-21 – Pojezierze Litewskie (Lithuanian Lake District),
Równina Augustowska (Augustów Plain): Puszcza Augustowska (Augustów Primeval Forest),
Łoska Forest District, forest section No. 7, Fraxinus excelsior, November 1, 1958, leg. K. Glanc
(KRAM-L 39051). GB-30 – Pojezierze Wschodniosuwalskie (East Suwałki Lake District),
Równina Augustowska (Augustów Plain): Puszcza Augustowska (Augustów Primeval Forest),
Płaska Forest District, forest section No. 23, Quercus robur, November 1, 1958, leg. K. Glanc
(KRAM-L 39054); Płaska Forest District, Acer platanoides, November 1, 1958, leg. K. Glanc
(KRAM-L 39079); Quercus robur, November 1, 1958, leg. K. Glanc (KRAM-L 39078); Żyliny
village, Salix sp., November 3, 1958, leg. K. Glanc (KRAM-L 39073). GB-31 – Pojezierze
Wschodniosuwalskie (East Suwałki Lake District), Równina Augustowska (Augustów Plain):
Puszcza Augustowska (Augustów Primeval Forest), Starożyn Nature Reserve, Acer platanoides,
September 17, 1986, leg. S. Cieśliński (KTC); Acer sp., July 18, 1965, leg. J. Zielińska (WA);
Puszcza Augustowska (Augustów Primeval Forest), forest section No. 51/24, Łozy Forest
District, Fraxinus excelsior, September 13, 1986, leg. S. Cieśliński (KTC); Puszcza Augustowska
(Augustów Primeval Forest), forest section No. 442, Salix sp., September 12, 1986, leg.
S. Cieśliński (KTC). GB-32 – Pojezierze Wschodniosuwalskie (East Suwałki Lake District),
Równina Augustowska (Augustów Plain): Puszcza Augustowska (Augustów Primeval Forest),
Kudrynki village near Rudawki village, Acer platanoides, September 16, 1986, leg. S. Cieśliński
(KTC). GB-40 – Nizina Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska
(Biebrza Basin): Puszcza Augustowska (Augustów Primeval Forest), forest section No. 153,
ca. 1 km of Lebiedzin village, Fraxinus excelsior, September 21, 1986, leg. S. Cieśliński (KRAML 31931). GB-91 – Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna
Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn Forest), 1 km NWW of
Machnacz village, forest section No. 130, Fraxinus excelsior, September 1997, leg. M. Murati,
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K. Glanc & Z. Tobolewski (KTC). GB-92 – Nizina Północnopodlaska (North Podlachian
Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn
Forest), Budzisk Nature Reserve, Carpinus betulus, August 5, 1994, leg. S. Cieśliński (KTC);
Fraxinus excelsior, August 5, 1994, leg. S. Cieśliński (KTC); Acer platanoides, May 13, 1993,
leg. K. Kolanko (University of Bialystok, Institute of Biology). GC-02 – Nizina Północnopodlaska
(North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): ca. 4 km SW
of Sokołda village, forest section No. 120, Fraxinus excelsior, August 4, 1994, leg. S. Cieśliński
(KTC); Knyszyn Forest Landscape Park, Czołnowo Nature Reserve, Acer sp., March 29, 1994,
s. coll. (University of Bialystok, Institute of Biology); Carpinus betulus, March 29, 1994, s.
coll. (University of Bialystok, Institute of Biology); Quercus robur, March 29, 1994, leg.
K. Kolanko (University of Bialystok, Institute of Biology). GC-03 – Nizina Północnopodlaska
(North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza
Knyszyńska (Knyszyn Forest), Stare Biele Nature Reserve, s. d., s. coll. (University of Bialystok,
Institute of Biology). GC-44 – Nizina Północnopodlaska (North Podlachian Lowland),
Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest), Browsk
Forest District, forest section No. 185c,d, Fraxinus excelsior, 1983, leg. Z. Tobolewski &
S. Cieśliński (KTC); Browsk Forest District, Ulmus sp., August 21, 1962, leg. J. Rydzak (LBL);
Acer platanoides, August 21, 1962, leg. J. Rydzak (LBL). GC-45 – Nizina Północnopodlaska
(North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Narewka Forest District, Fraxinus excelsior, August 25, 1968, leg. J. Rydzak
(LBL). GC-54 – Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska
(Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest), Zwierzyniec village, forest
section No. 447b, Quercus sp., 1982, leg. Z. Tobolewski & S. Cieśliński (KTC); forest section
No. 281d, Fraxinus excelsior, 1981, leg. Z. Tobolewski & S. Cieśliński (KTC); Zwierzyniec
Forest District, Ulmus sp., August 24, 1965, leg. J. Rydzak (LBL); Acer sp., August 25, 1965,
leg. J. Rydzak (LBL); forest section No. 220, Fraxinus excelsior, August 25, 1965, leg. J. Rydzak
(LBL); Acer platanoides, August 26, 1965, leg. J. Rydzak (LBL); Acer sp., September 13, 1965,
leg. J. Rydzak (LBL). GC-55 – Nizina Północnopodlaska (North Podlachian Lowland),
Równina Bielska (Bielsk Plain): Białowieża National Park, Quercus robur, August 12, 2002,
leg. P. Czarnota (GPN 2949); forest section No. 283, S of Orłówka River, Tilia cordata, 1982,
leg. Z. Tobolewski & S. Cieśliński (KTC); forest section No. 342b, Carpinus betulus, 1979, leg.
Z. Tobolewski & S. Cieśliński (KTC); forest section No. 256, Carpinus betulus, 1998, leg.
S. Cieśliński (KTC); forest section No. 256, Carpinus betulus, 1988, leg. K. Glanc (KTC);
Pogorzelce village, Quercus robur, May 21, 1950, leg. S. Cieśliński (LBL); forest section No.
256, Carpinus betulus, August 2014, leg. M. Kukwa 12891, A. Łubek (UGDA-L 20440); May
2014, leg. M. Kukwa 12737, 12610, A. Łubek (UGDA-L 20334, 20246); forest section No. 256,
Acer platanoides, August 2014, leg. M. Kukwa 13288, A. Łubek (UGDA-L 20734); May 2014,
leg. M. Kukwa 12754, A. Łubek (UGDA-L 20345); on bark of snag (Fraxinus excelsior), October
2014, leg. M. Kukwa 13062, A. Łubek (UGDA-L 20572). GC-65 – Nizina Północnopodlaska
(North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Białowieża Forest District, Białowieża Forest Division, forest section No.
609a,b, on Podcerkowka River, Fraxinus excelsior, 1983, leg. Z. Tobolewski & S. Cieśliński
(KTC); Fraxinus excelsior, August 21, 1961, leg. J. Rydzak (LBL). GD-01 – Wysoczyzny
Podlasko-Białoruskie (Podlachian-Belarusian Highlands), Wysoczyzna Drohiczyńska (Drohiczyn Highland): forest between Adamów and Koterka villages, forest section No. 275,
Fraxinus excelsior, May 11, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC). GD-20 – Nizina
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Południowopodlaska (South Podlachian Lowland), Równina Łukowska (Łuków Plain):
Chmielne Nature Reserve, Leśna near Biała Podlaska town, Ulmus sp., May 18, 1962, leg.
B. Ordyczyńska (LBL). GD-95 – Polesie Zachodnie (Western Polesie), Polesie Brzeskie:
Sobibór village near Włodawa town, Quercus sp. 1965, leg. J. Bystrek (LBL). GE-52 – Wyżyna
Lubelska (Lublin Upland), Działy Grabowieckie mountain ranges: Krupe near Krasnystaw
villages, Betula sp., 1975, leg. J. Nowak (LBL). GE-91 – Roztocze, Central Roztocze: Kosobudy
Forest District, Fagus sylvatica, September 25, 1964, leg. B. Warmińska (LBL); Krzywe Forest
District, Jarugi Forest District, Fagus sylvatica, August 25, 1964, leg. B. Warmińska (LBL);
Rudka village, Corylus avellana, September 1960, leg. J. Bystrek (LBL). GF-13 – Roztocze,
Central Roztocze: Ulmus sp., s. coll. (KRAM-L 39043); Hrebenne village, Fagus sylvatica,
March 30, 1957, leg. K. Glanc (KRAM-L 39044). GF-41 – Kotlina Sandomierska (Sandomierz
Basin), Płaskowyż Tarnogrodzki (Tarnogród Plateau): near Oleszyce town, Fagus sylvatica,
1973, leg. J. Bystrek (LBL); Bida village near Oleszyce town, Fagus sylvatica, leg. J. Bystrek
(LBL). GF-80 – Kotlina Sandomierska (Sandomierz Basin), Dolina Dolnego Sanu (Lower
San River Valley): Przemyśl City, Acer sp., 1969, leg. J. Piórecki (LBL). GG-71 – Beskidy Lesiste
Mountains, West Bieszczady Mountains: Bieszczady National Park, Abies alba, June 19, 2002,
leg. P. Czarnota (GPN 2875); Sianki village, Niedźwiedzi Stream, Acer pseudoplatanus, August
25, 1999, leg. R. Kościelniak (KRAP 1886).
Additional material examined. AUSTRIA: August 1890, leg. E. Fries (KRAM-L 3444): ca.
6 km N of Kaprun, Carpinus betulus, April 1, 2005, leg. A. Wieczorek (herb. Wieczorek).
BELARUS: Gomel Region, Gomel District, Chenki forest, 52°19' N, 30°57' E, near F. Skorina
Gomel State University field practice camp, Populus sp., June 20, 2006, leg. A. Tsurykau 01445
(GSU). CZECH REPUBLIC: Southern Moravia, Vyškov District, Moravský kras, Drahanská
vrchovina, Ruprechtov, Malá Haná valley below castle ruins Kuchlov, Acer campestre, April
20, 2007, leg. U. Schiefelbein 2085 (herb. Schiefelbein); Botanischer Tauschverein in Wien
No. 4120, 1854, leg. W. Siegmund (WA); South Moravia, Jihomoravsky kraj, S of Lanžhot,
Polinský vršek, elev. 175 m, edge of a managed floodplain forest, ca. 80 years old, few isolated
trees in an adjacent meadow, Acer campestre, March 31, 2014, leg. M. Kukwa 12364 (UGDAL 12364). DENMARK: Bornholm, Dondalen, forest, Carpinus betulus, July 5, 2014, leg.
A. Wieczorek & A. Łysko (herb. Wieczorek); Ekkodalen, Carpinus betulus, July 5, 2014, leg.
A. Wieczorek (herb. Wieczorek). FINLAND: s. loc., January 5, 1933, s. coll., (WAR.). GREAT
BRITAIN: Fountains Abbey, Acer sp., May 3, 2013, leg. V. John & M. R. D. Seaward (herb.
Seaward). GERMANY: Fyn, Romso Island, Fraxinus excelsior, October 30, 1994, leg. J. Motiejunaite 7893 (BILAS); Mecklenburg-Vorpommern, Kreis Uecker-Randow, Kleppelshagener
Forst, Fagus sylvatica, October 21, 1999, leg. U. Schiefelbein 500 (herb. Schiefelbein); ca. 3 km
SSW of Mönkebude, ehemalige Hoftselle, Acer pseudoplatanus, April 8, 1999, leg. U. Schiefelbein 304 (herb. Schiefelbein); SE VW. Theerofen, Ulmus sp., November 22, 2001, leg. U. Schiefelbein 1241 (herb. Schiefelbein); N Landstraße Jatznick-Rothemühl, ca. 3 km S of Eichhof,
Fagus sylvatica, June 3, 2001, leg. U. Schiefelbein 748 (herb. Schiefelbein); ca. 3 km E of
Storkow, ca. 0.5 km NE of Kotzensee, Ulmus sp., May 15, 2002, leg. U. Schiefelbein 885 (herb.
Schiefelbein); Wald, ca. 1.5 km N of Marienthal, Ulmus sp., December 17, 2001, leg. U. Schiefelbein 939 (herb. Schiefelbein); südlicher Rand des Thursees, ca. 1.2 km N of Pampow, Fagus
sylvatica, April 6, 2002, leg. U. Schiefelbein 991 (herb. Schiefelbein); Landkreis Müritz,
Wäldchen W VW. Troja, ca. 0.2 km W of Troja, Acer platanoides, November 1, 2002, leg.
U. Schiefelbein 1184 (herb. Schiefelbein); Landkreis Ostvorpommern, Insel Usedom, Wald
© The Author(s) 2018
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Wieczorek / The lichen genus Opegrapha s. l. in Poland
zwischen der Ortslage Ahlbeck und der polnischen Grenze, nördl. der Bundesstraße, Fagus
sylvatica, November 3, 2003, leg. U. Schiefelbein 1926 (herb. Schiefelbein); Ückeritz, Naturschutzgebiet “Wockninsee”, NE vom See, am Aussichtsturm, Fagus sylvatica, November 27,
2002, leg. U. Schiefelbein 1321 (herb. Schiefelbein); Greifswalder Oie am Leuchtturm, Sambucus
nigra, November 7, 2002, leg. U. Schiefelbein 1324 (herb. Schiefelbein); Kreis Güstrow, May
10, 1998, leg. U. Schiefelbein 205 (herb. Schiefelbein); Niedersachsen, Landkreis LychowDannenb, W Naturschutzgebiet Breeser Grund, ca. 4 km SSE of Göhrde, ca. 1.5 km S of Gr.
Stern, Fagus sylvatica, October 26, 2003, leg. U. Schiefelbein 1376 (herb. Schiefelbein).
LITHUANIA: Biržų raj., Biržų giria, Spalviškių gja, 235 kvartalas, Fraxinus excelsior, July 7,
2003, leg. S. Skuja 10305 (BILAS); Akmenės raj., Kruopių g-ja, 548 kvartalas, 27/28 sklypas,
Fraxinus excelsior, September 20, 2003, leg. E. Panovienė 10319 (BILAS); Naujosios Akmenės
g-ja, 438 kvartalas, 16 sklypas, ant lapuočio medžio kamieno, September 2003, leg. E. Panovienė
8460 (BILAS); 441 kvartalas, 50 sklypas, ant lapuočio medžio kamieno, September. 2003, leg.
E. Panovienė 8461 (BILAS); Raseinų raj., Viduklės g-ja, 89 kvartalas, 11 sklypas, ant lapuočio
medžio kamieno, November 26, 2003, leg. A. Masaitis 9468 (BILAS); Vilniaus raj., Liubavo
dvaras, Fraxinus excelsior, October 11, 2007, leg. D. Stončius 9316 (BILAS); Kelmės raj.,
Užpelkių miškas, Tytuvėnu g-ja, 133 kvartalas, 14 sklypas, Fraxinus excelsior, January 7, 2007,
leg. D. Stončius 9315 (BILAS); Ukmergės raj., Narkuškių miškas, Giedraičių g-ja, 56 kvartalas,
17 sklypas, Fraxinus excelsior, June 14, 2009, leg. D. Stončius 9317 (BILAS); Asvejos RP,
Molėtų raj., Dubingiu g-ja, 1075 kvartalas, 48 sklypas, Ulmus sp., April 24, 2010, leg. D. Stončius
9622 (BILAS); Labanoras Regional Park, Molėtai District, Stirniai village, between Lakes
Galnakis I and Galnakis II, Acer platanoides, April 4, 2011, leg. D. Stončius 10244 (BILAS);
Elėktrenų sav., Vievio miškas, Vievio g-ja, 153 kvartalas, 16 sklypas, Acer platanoides, November
14, 2006, leg. D. Stončius 9524 (BILAS); Kuršių Nerijos NP, palvės miškas ties II Nidos
įvažiavimu, Betula sp., July 29, 2006, leg. D. Stončius 10267 (BILAS); Širvintų raj., Sailidžioniu
miškas, Širvintų g-ja, 34 kvartalas, 9 sklypas, Quercus robur, September 15, 2006, leg. D. Stončius
8706 (BILAS); Molėtų raj., Šešuolėlių g-ja, 18 kvartalas, 15 skl., Šilo miškas, Fraxinus excelsior,
November 26, 2006, leg. D. Stončius 8281 (BILAS); Sartu RP, Dusetu giria, Vosynos rezervatas,
25 kvartalas, 8 sklypas, Acer platanoides kamieno pammatines dallies, June 10, 2003, leg.
J. Motiejunaite 6578 (BILAS); Molėtų raj., miškelis į V nuo Aleksandriškių km., Giedraičių
gja, 681 kvartalas, 63 sklypas, Ulmus glabra, December 7, 2006, leg. D. Stončius 7791 (BILAS);
Panemunių regioninis parkas, Šakių raj., Ilguvos apyl., Valniaravio rezervatas, Fraxinus excelsior, June 16, 2007, leg. A. Uselienė 9685 (BILAS); Vilniaus raj., Šuzionių g-ja, 74 kvartalas,
5 skl., Šuzionių miškas UTM: 595620-6094584, Quercus robur, October 24, 2012, leg.
D. Stončius10511 (BILAS); Žemaitijos NP., Plungės raj., Plateliai, Fraxinus excelsior, September
14, 2001, leg. J. Motiejunaite 5636 (BILAS); Liepijos miskas, Tilia cordata, May 12, 2002, leg.
J. Motiejunaite 5863 (BILAS); Lazdijų raj., Rinkoto miškas, 1 kvartalas, Dusios ežero krantas,
Quercus robur, April 3, 2001, leg. J. Motiejunaite 5635 (BILAS); Kurtuvėnų RP, Šiaulių raj.,
Vainagiai, Acer platanoides, July 2003, leg. V. Lopeta 7134 (BILAS); Bubių g-ja, 45 kvartalas,
2 sklypas, Fraxinus excelsior, August 27, 2002, leg. V. Lopeta 5813 (BILAS); Ąžuolijos miškas,
Utenos raj, 44 kvartalas, 7 sklypas, Quercus robur, September 8, 2001, leg. I. Prigodina 2916
(WI); 47 kvartalas, 7 sklypas, Utenos raj., Quercus robur, August 22, 2001, leg. I. Prigodina
3133 (WI); Dūkstu ąžuolynas, 3 kvartalas, Neries RP, Vilniaus raj., Quercus robur, June 25,
2002, leg. I. Prigodina 1276 (WI); 10 kvartalas, 3 sklypas, Neries RP, Quercus robur, September
13, 2001, leg. I. Prigodina 2944 (WI); 9 kvartalas, 4 sklypas, Neries RP, Vilniaus raj., Quercus
robur, July 10, 2002, leg. I. Prigodina 1275 (WI); Varnikų miškas, 95 kvartalas, 4 sklypas,
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Tarkų istorinis-nacionalinis parkas, Quercus robur, June 4, 2002, leg. I. Prigodina 1299 (WI);
95 kvartalas, 4 sklypas, Traku istornis nacionalinis parkas, Quercus robur, June 9, 2002, leg.
I. Prigodina 2973 (WI); Palanga, ant klevo kamieno, August 8, 1975, leg. T. Rudzinskaitė 2166
(WI); Juodkrantė, Kuršių Nerija, ant klevo kamieno, August 2, 1971, leg. T. Rudzinskaitė 2456
(WI). NETHERLANDS: Olanda, Ohenby, 1943, leg. O. Almborn. NORWAY: Stryn, forest,
Abies alba, August 28, 1995, leg. A. Wieczorek (herb. Wieczorek). RUSSIA: Republic of
Karelia, Sortavala District, Ladoga Lake, Valaamarchipelago, W of Valaam island, peninsula
between Bolshaya Nikonovskaya Bay and Krasny Gulf, Sorbus aucuparia, August 13, 2004,
leg. U. Schiefelbein 1820 (herb. Schiefelbein); Nizhni Novgorod Oblast, Sharanga region,
Kilemarsky Reserve, linden-spruce forest, Tilia sp., September 27, 1999, leg. A. Shestakova
(NNSU); Arzamas region, near Staray Pustyn village, Biological Station of State University,
Tilia sp., August 24, 1994, leg. P. Suvorov (NNSU); Baikalsky Nature Reserve, Republic of
Buryatia, Khamar-Daban, near of mouth of Verkhnij Khandagaita River, pine(sibirica)spruce-birch forest, Populus sp., July 29, 2002, leg. G. P. Urbanavichus 1990 (KPABGI);
Kalujskie zaseki Reserve, Kaluga Province, Ulyanovsky District, flood plain, limeoak forest,
Quercus sp., July 27, 2004, leg. A. V. Gudovicheva 11989 (KPABGI); Orel region, T. 217,
Znamensky Distr., Orlovskoye Polesie National Park, Krasnikovskoye Forestry, 33 quar.,
broad-leaved forest, bark of Ulmus sp., 53°29.175' N, 35°33.951' E, alt. 196 m, June 12, 2012,
leg. E. E. Muchnik 9048 (OHHI); Ryazan region, T. 112, Kadomsky Distr., right bank of
Moksha River, oak forest, bark of Ulmus, 54°26.156' N, 42°27.710' E, alt. 116 m, July 27, 2010,
leg. E. E. Muchnik & L. A. Konoreva 102 (RUS); Ryazan region, T. 106, Kadomsky Distr.,
Kadomskoye Forestry, 28 quar., mied forest, bark of Quercus, 54°30.561' N, 42°23.399' E, alt.
111 m, July 25, 2010, leg. E. E. Muchnik & L. A. Konoreva 120 (RSU); Bryansk region, T. 378,
Suzemsky Distr., Bryansky les State Nature Reserve, 103 quar., broad-leaved forest, bark of
Fraxinus excelsior, 52°27.286' N, 33°51.155' E, alt. 153 m, August 10, 2015, leg. E. E. Muchnik
(herb. Muchnik). SLOVAKIA: August 5, 1959, leg. Vězda (WAAR). SWEDEN: Populus sp.,
July 13, 1949, leg. E. Smaland, P. Dodershult, ST. Ekershult & A. H. Magnusson (KRAML 34679). UKRAINE: near Kamenets-Podolsk, an baumrinden, 1909, leg. V. Kutak (WRSL).
USA: New York, Carya alba, May 1893, leg. N. Y. Alcove (herb. Seaward e Mus. Bot.
Stockh.).
Exsiccatae seen. Wetmore, Lich. Exs. No. 44, as Opegrapha varia (KRAM); Poelt & Steiner,
Lich. Alp. No. 342, as Opegrapha lichenoides var. chlorina (KRAM); Poelt & Steiner, Lich. Alp.
No. 371 as Opegrapha lichenoides var. chlorina (KRAM); Savic, Lichenotheca Rossica. Decas
3, as Opegrapha lichenoides (BILAS, WI); Lich. Helv. Exs. No. 282 (WRSL).
Arthonia atra (Pers.) A. Schneid.
Specimens examined. AB-24 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam and Wolin
islands; Wolin Island, Wolin National Park, Carpinus betulus, August 15, 1996, leg. A. Wieczorek
(herb. Wieczorek). AB-94 – Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe
(Beech Hills): Puszcza Bukowa (Beech Forest), Szczecin Landscape Park, Carpinus betulus,
August 20, 1997, leg. A. Wieczorek (herb. Wieczorek). BA-88 – Pobrzeże Koszalińskie (Koszalin
Coastland), Równina Słupska (Słupsk Plain): Wodzisław town, Quercus sp., August 28, 1987, leg.
W. Fałtynowicz & J. Miądlikowska (UGDA-L 3901). BA-93 − Pobrzeże Koszalińskie (Koszalin
Coastland), Wybrzeże Słowińskie (Slovincian Coast): Mielno town, Fagus sylvatica, June 8,
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1984, leg. W. Fałtynowicz (UGDA-L 2845). BC-12 – Pojezierze Południowopomorskie (South
Pomeranian Lake District), Równina Drawska (Drawno Plain): Drawieński Park Narodowy
(Drawa National Park), Trangankowe Urwisko Nature Reserve, about 2 km of Zatom village,
Fagus sylvatica, May 6, 2010, leg. U. Schiefelbein 2621 (herb. Schiefelbein). BC-22 – Pojezierze
Południowopomorskie (South Pomeranian Lake District), Równina Drawska (Drawno Plain):
Drawieński Park Narodowy (Drawa National Park), Lake Ostrowiec, Fraxinus sp., August
17, 1989, leg. L. Lipnicki (herb. Lipnicki); Radęcin Nature Reserve, Fagus sylvatica, July 7,
2002, leg. A. Wieczorek (herb. Wieczorek); beech forest, forest section No. 250, Carpinus
betulus, July 7, 2002, leg. A. Wieczorek (herb. Wieczorek). BF-13 – Central Sudetes, Stołowe
Mountains: near Karłów–Kudowa highway, Acer pseudoplatanus, September 25, 1951, leg.
Z. Tobolewski (POZ). CA-38 – Pobrzeże Koszalińskie (Koszalin Coastland)land, Wybrzeże
Słowińskie (Slovincian Coast): Bielawskie Błoto near Silice village, Populus tremula, August
19, 1951, leg. Z. Tobolewski (POZ). CA-58 – Pobrzeże Koszalińskie (Koszalin Coastland),
Pradolina Redy-Łeby (Reda-Łeba Urstromtal): 2 km of Wejherowo town, Fagus sylvatica,
July 1983, leg. A. Zalewska (UGDA-L 2882). CA-69 – Pojezierze Wschodniopomorskie (East
Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): between Rumia
and Zagórze towns, Fagus sylvatica, March 28, 1987, leg. W. Fałtynowicz (UGDA-L 3268).
CA-99 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Dolina Reknicy (Reknica River Valley), Acer sp., December 5,
1976, leg. W. Fałtynowicz (UGDA-L 3405). CB-09 – Pojezierze Wschodniopomorskie (East
Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Zaskoczyn
village, Acer sp., April 5, 1986, leg. W. Fałtynowicz & Z. Tobolewski (UGDA-L 3388). CB-75 –
Pojezierze Południowopomorskie (South Pomeranian Lake District), Dolina Brdy (Brda River
Valley): Piekło Nad Brdą Nature Reserve, Quercus sp., March 30, 1985, leg. L. Lipnicki (TRN).
CC-27 – Pradolina Toruńsko-Eberswaldzka (Toruń-Eberswalder Urstromtal), Kotlina Toruńska
(Toruń Basin): Ostromecko village, Chełmno County, Carpinus betulus, May 22, 1952, leg.
J. Hutorowicz (TRN). DA-85 – Pobrzeże Gdańskie (Gdańsk Coastland), Mierzeja Wiślana
(Vistula Spit): Krynica Morska town, forest section No. 26m, Populus sp., August 7, 1981, leg.
E. Budzbon (UGDA-L 2188). DB-05 – Pobrzeże Gdańskie (Gdańsk Coastland), Wysoczyzna
Elbląska (Elbląg Highland): Elbląg City, Betula sp., June 21, 1978, leg. H. Karwowska (TRN).
DB-53– Pojezierze Południowobałtyckie (South Baltic Lakeland), Pojezierze Iławskie (Iława
Lake District): E of Szadowo village, valley of Liwa River, N bank of the river, hornbeam-linden
forest, Carpinus betulus, April 16, 2006, leg. M. Kukwa 4949 (UGDA-L 13532). DE-57 –
Wyżyna Przedborska (Przedborska Upland), Wzgórza Radomszczańskie (Radomsko Hills):
Kobiele Wielkie Nature Reserve, Quercus robur, July 7, 1971, leg. K. Czyżewska (LOD-L 1643).
DE-68 – Wyżyna Przedborska (Przedborska Upland), Niecka Włoszczowska (Włoszczowa
Basin): Dębowiec Nature Reserve, forest section No. 162, Ulmus glabra, October 30, 1969, leg.
K. Czyżewska (LOD-L 1022). EB-01 – Nizina Staropruska (Old Prussian Lowland), Równina
Ornecka (Orneta Plain): Mingajny village, Salix sp., May 10, 1989, leg. S.Cieśliński (KTC).
EB-22 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn
Lake District): about 1.5 km NNE of Smolajny village, Fraxinus excelsior, August 23, 1993,
leg. S. Cieśliński (KTC). ED-91 – Wzniesienia Południowomazowieckie (South Mazovian
Hills), Równina Piotrkowska (Piotrków Plain): Puszcza Pilicka (Pilica Forest), Konewka
Nature Reserve, forest section No. 153, Quercus petraea, in thermophilous oak forest, October 10, 1974, leg. K. Czyżewska (LOD-L 4497). EE-83 – Wyżyna Kielecka (Kielce Upland),
Świętokrzyskie Mountains: Podzamcze near Chęciny town, forest section No. 176, Quercus
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sp., March 10, 1977, leg. K. Toborowicz (KTC). EG-11 – West Beskids, Gorce Mountains:
in Lasek Żydowski by Polana Jaworzynka in valley of Gorcowy Potok (Gorcowy Stream),
Acer pseudoplatanus, February 12, 1997, leg. P. Czarnota (GPN 1841); in shady gorge above
stream, at field edges in Lubomierz town (housing estate Borki), Salix sp., May 29, 2000, leg.
P. Czarnota (GPN 2311). EG-21 – West Beskids, Gorce Mountains: Turbacz Nature Reserve,
Fagus sylvatica, May 12, 1992, leg. P. Czarnota (GPN 231), September 6, 1959, leg. K. Glanc
(KRAM-L 39395). EG-33 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression),
Pieniny Mountains: by Pieniński Potok (Pieniny Stream), Corylus avellana, August 20, 1955,
leg. Z. Tobolewski (POZ). FB-03 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza Borecka (Borki Forest), forest section No. 195, Fagus
sylvatica, May 21, 1985, leg. S. Cieśliński & Z. Tobolewski (KTC); Puszcza Borecka (Borki
Forest), near Czerwony Dworek village, Fraxinus excelsior, May 21, 1985, leg. Z. Tobolewski
& S. Cieśliński (KTC). FB-56 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze
Ełckie (Ełk Lakeland): Kopijki village near road to Prostki village, Fagus sylvatica, August
31, 1988, leg. S. Cieśliński (KTC). FG-08 – Pogórze Środkowobeskidzkie (Central Beskidian
Foothills), Pogórze Przemyskie (Przemyśl Foothills): Leszczawa Góra village, near cemetery,
Carpinus betulus, July 21, 1984, leg. J. Piórecki (BDPA). FG-69 – Beskidy Lesiste Mountains,
West Bieszczady Mountains: Ustrzyki Górne village, Acer pseudoplatanus, August 5, 1959,
leg. K. Glanc (KRAM-L 39496). GB-31 – Pojezierze Litewskie (Lithuanian Lake District),
Równina Augustowska (Augustów Plain): Puszcza Augustowska (Augustów Primeval Forest),
Starożyn Nature Reserve, forest section No. 195, Fraxinus excelsior, September 17, 1986, leg.
S. Cieśliński (KRAM-L 31823).
Additional material examined. AUSTRIA: Botanischer Tauschverein in Wien, leg. Verseghy
(KRAM-L 381, 388). CZECH REPUBLIC: Deutschbrod in Böhmen, leg. Jul. Svacha (KRAML 390, 396). DENMARK: Damshalte, Hyelmsmark, Populus sp., June 13, 1947, leg. M. Christiansen (POZ); Bornholm, Dondalen, forest, Carpinus betulus, July 5, 2014, leg. A. Wieczorek
& A. Łysko (herb. Wieczorek); Ekkodalen, Carpinus betulus, July 5, 2014, leg. A. Wieczorek
& A. Łysko (herb. Wieczorek). GERMANY: Blankenberg, Hampe, (KRAM-L 27008, 26977);
Balingen, 1950, leg. F. O. Westerberg (WA); Mecklenburg-Vorpommern, Ucker-Randow,
mied forest, October 14, 2001, leg. U. Schiefelbein 743 (herb. Schiefelbein); moist alder forest,
Fraxinus excelsior, March 12, 2000, leg. U. Schiefelbein 497 (herb. Schiefelbein); Straße JatznickRothemühl, ca. 1 km E of Rothemühl, Fagus sylvatica, October 19, 1999, leg. U. Schiefelbein
614 (herb. Schiefelbein); Buchenwald auf dem Butterberg, Fagus sylvatica, April 1, 2001, leg.
U. Schiefelbein 609 (herb. Schiefelbein); Demnitzer Bruch, Eschen-Mischwald, Fraxinus
excelsior, October 22, 2000, leg. U. Schiefelbein 711 (herb. Schiefelbein); Klepelshagen Nature
Reserve on Knüppelbach, Quell-Eschenwald forest, Carpinus betulus, October 20, 1999, leg.
U. Schiefelbein 455 (herb. Schiefelbein); 2.5 km N of Ahlbeck, Eschen-Mischwald, Fraxinus
excelsior, April 24, 1998, leg. U. Schiefelbein 460 (herb. Schiefelbein); Nordwestmecklenburg,
Klützer Winkel, Brooker forest, 1 km NE of Schlossgut Groß Schwanensee, Fagus sylvatica,
October 29, 2010, leg. U. Schiefelbein 2464 (herb. Schiefelbein); Landkreis Ostvorpommern,
Greifswalder Oie, orchard near lamppost, Malus sylvestris, August 11, 2007, leg. U. Schiefelbein
2123 (herb. Schiefelbein); Mecklenburg Vorpommern, Ostvorpommern region, Eschenwald
N Müggenburg, July 30, 1996, leg. U. Schiefelbein 152 (herb. Schiefelbein); Blesewitzer Holz,
1.5 km SE of Sanitz, July 30, 1996, leg. U. Schiefelbein 149 (herb. Schiefelbein); W Spantekow,
Kaiserkoppel, Fraxinus sp., July 17, 1996, leg. U. Schiefelbein 148 (herb. Schiefelbein); ca.
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2 km E of Wockninsee, Carpinus betulus, November 27, 2002, leg. U. Schiefelbein 1296
(herb. Schiefelbein); Koserow, an der Steilküste, NW of Streckelberg Nature Reserve, Fagus
sylvatica, April 2, 2002, leg. U. Schiefelbein 1299 (herb. Schiefelbein); waldgebiet zwischen
Seebad Bansin und Ückeritz, ca. 0.5 km westlich der ehemaligen Selliner Bootstelle, Fraxinus
excelsior, November 14, 2003, leg. U. Schiefelbein 1382 (herb. Schiefelbein). GREAT BRITAIN:
Worcestershire, Alfrick, leg. H. B. Holl (KRAM-L 26998). ITALY: Caldes, Baglietto, (KRAML 26800). IRELAND: Mullet Peninsula, Lighthouse Island, Sambucus sp., June 12, 1996, leg.
M. R. D. Seaward 107490 (herb. Seaward). POLAND: s. loc., March 19, 1864, leg. Strasburger
(WA). RUSSIA: Nizhni Novgorod Oblast, Vetluga region, Klenovik forest reserve, birch
forest, Sorbus sp., August 13, 1999, leg. M. Sharapova (NNSU); Varnavino region, between
Nepogodiha and Reshetiha villages, spruce forest, on young Sorbus sp., August 7, 1999, leg.
M. Sharapova (NNSU); Bogorodsk region, near Shoniha station, coniferous-deciduous forest,
on Populus tremula, August 2, 1998, leg. M. Sharapova (NNSU). SWITZERLAND: Zurich,
October 1877, leg. J. G. Winter (KRAM-L 27014).
Exsiccate seen. Arnold, Lich. Exs. No. 767, as Opegrapha gyrocarpa for. rhododendri (KRAM);
Krypt. Exs. No. 1526, as Opegrapha atra (KRAM); Bot. Mus. Hist Verseghy Lich. Exs. No.
76, as Opegrapha atra (KRAM); Baglietto, Cesati & Notaris, Erb. Critt. Ital. Ser. I No. 203, as
Opegrapha atra (KRAM).
Arthonia calcarea (Turner ex Sm.) Ertz & Diederich
Specimens examined. DG-39 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Pogórze Spisko-Gubałowskie (Spisz-Gubałówka Foothills): Skalice Nowotarskie,
limestone, July 20, 1997, leg. A. Wieczorek (herb. Wieczorek). DG-58 − Łańcuch Tatrzański
Mountains, Western Tatras: Dolina Chochołowska (Chochołów Valley), rock, July 24, 1997,
leg. A. Wieczorek (herb. Wieczorek).
Additional material examined. DENMARK: Jylland, leg. M. Christiansen (LBL); Bornholm:
an Wallen des Schlosses Hamnershuus auf der Insel, 1884, leg. Hellbom (KRAM-L 44331).
GERMANY: Hansted, limestone, July 25, 1942, (BMUC); Schleswig-Holstein, Nordfriesland,
Insel Föhr, Süderende, Kirchof St. Laurentius, concrete, October 9, 2009, leg. U. Schiefelbein
2424 (herb. Schiefelbein); Niedersachsen, Landkreis Leer, Gem. Jemgum, Holtgaste, alte
Kirche am Ortsrand, concrete, October 23, 2004, leg. U. Schiefelbein 1874 (herb. Schiefelbein);
Gem. Weener, Stapelmoor, alte Kirche im Dorfzentrum mit Baumbestand, concrete, October
23, 2004, leg. U. Schiefelbein 1872 (herb. Schiefelbein). GREAT BRITAIN: Herb. C. du Bois
Larbalestier, April 1896, (KRAM-L 27266).
Exsiccate seen. Arnold, Lich. Exs. No.1398, as Opegrapha conferta (KRAM); Anzi, Lich.
Exs. No. 36, as Opegrapha calcarea (KRAM); Anzi, Lich. Exs. No. 37, as Opegrapha calcarea
(KRAM); Zwackh, Lich. Exs. No.1194, as Opegrapha calcarea (KRAM).
Gyrographa gyrocarpa (Flot.) Ertz &Tehler
Specimens examined. AE-78 – Western Sudetes, Karkonosze Mountains: Owcze Skały,
July 30, 2008, leg. M. Kossowska (herb. Kossowska); Twarożnik Mount, October 6, 2007,
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leg. M. Kossowska (herb. Kossowska). AE-88 − Western Sudetes, Karkonosze Mountains:
Łabski Szczyt Mount, July 31, 2008, leg. M. Kossowska (herb. Kossowska). BF-06 – Western
Sudetes, Sowie Mountains: Srebrna Góra (Silver Mount), walls by fortress, rock, April 22,
2004, leg. M. Kukwa 3102 (UGDA-L 10288). BF-14 – Central Sudetes, Stołowe Mountains:
ca. 5 km of Karłów village, August 26, 1954, leg. Z. Tobolewski (POZ); near road from Karłów
to Radków villages, July 17, 1951, leg. Z. Tobolewski (POZ); near road from Pasterka to
Nowy Karłów villages, August 6, 1960, leg. Z. Tobolewski (POZ). BF-38 – Eastern Sudetes,
Góry Złote (Golden Mountains): Bialskie Mountains, Trzy Siostry, August 7, 2003, leg.
K. Szczepańska (herb. Szczepańska). BF-47 – Eastern Sudetes, Masyw Śnieżnika (Śnieżnik
Massif): Bialskie Mountains, Goworek Mount, June 28, 2004, leg. K. Szczepańska (herb.
Szczepańska). DG-13 – West Beskids, Beskid Śląski Mountains: western slope of Barania Góra
(Ram Mountain), sandstone, July 22, 1965, leg. J. Kiszka (KRAP); SW crest of Barania Góra
(Ram Mountain), sandstone, June 8, 1966, leg. J. Kiszka (KRAP); sandstone, June 7, 1966,
leg. J. Kiszka (KRAP); Barania Góra (Ram Mountain), Biała Wisełka Stream, sandstone, June
9, 1966, leg. J. Kiszka (KRAP). DG-17 – West Beskids, Beskid Żywiecki Mountains: Pasmo
Policy (Polica Range), Hala Śmietanowa, October 21, 1965, leg. J. Nowak (KRAM-L 15921).
DG-25 – West Beskids, Beskid Żywiecki Mountains: Pilsko Massif, Skałka Mount, July 16,
1966, leg. J. Nowak (KRAM-L 17040). DG-33 – West Beskids, Beskid Żywiecki Mountains:
Przysłop Wielki Mount, sandstone, August 1, 1962, leg. J. Kiszka (KRAP). EG-21 – West
Beskids, Gorce Mountains: Gorc Mount, S slope, August 23, 1967, leg. K. Glanc (KRAML 39486); Gorce National Park, under glade, N slope of Mostownica village, in Roztoka
Stream catchment area, July 30, 1999, leg. P. Czarnota (GPN); crest under Przysłop Mount,
sandstone, June 9, 1966, leg. J. Kiszka (KRAP); above Kopa gorge, near black tourist trail,
November 4, 1999, leg. B. J. Coppins (GPN 2145). EG-22 – West Beskids, Gorce Mountains:
Lubań Mount, August 9, 1968, leg. K. Glanc (KRAM-L 39485); N slope of Mostownica, July
30, 1999, leg. P. Czarnota (GPN 2244). EG-36 – West Beskids, Beskid Sądecki Mountains:
Jaworzyna Krynicka Mount, August 11, 1989, leg. A. Wieczorek (herb. Wieczorek); Żebracze
Nature Reserve, July 30, 1999, leg. P. Czarnota (GPN 2052); July 5, 2001, leg. P. Czarnota (GPN
2584, 2528); Pasmo Jaworzyny (Jaworzyna Range), Pusta Wielka Mount, August 23, 1967,
leg. M. Olech (KRA). EG-50 – West Tatras: Dolina Gąsienicowa (Gąsienica Valley) by Czarny
Potok (Black Stream) near Czarny Staw Gąsienicowy (lake), among scree in moist place, on
granitic stone, August 29, 2002, leg. A. Flakus 142 (KRAM-L 48297). EG-60 – West Tatras:
Wyżnia Spadowa Przełączka (pass), on granite rock, W aspect, the subnival belt; alt. 2,222 m,
August 20, 2005, leg. A. Flakus 5417 (KRAM-L 52156); valley behind Mnich Mount, on granite, alt. 1,950 m, August 21, 2002, leg. A. Flakus 107 (KRAM-L 48293); Ciemnosmerczyńska
Przełączka (pass), in vertical rock crack, mylonite area, alt. 2,115 m. N aspect slope 900,
August 16, 2003, leg. A. Flakus 1785 (KRAM-L 48343). FG-68 – Beskidy Lesiste Mountains,
West Bieszczady Mountains: Bieszczady National Park, Połonina Wetlińska (alpine meadow),
Roh Mount, rock debris on SW slope, sandstone, September 11, 2004, leg. R. Kościelniak
(KRAP 1870); alpine meadow, Osadzki Wierch Mountain, sandstone, September 10, 2004,
leg. J. Kiszka & R. Kościelniak (KRAP 1872); rock outcrops under mountain hut, sandstone,
September 12, 2007, leg. J. Kiszka & R. Kościelniak (KRAP 965); SE crest under mountain
hut, sandstone, September 9, 2004, leg. J. Kiszka & R. Kościelniak (KRAP 1874); S slope of
Halicz Mount, near tourist trail, sandstone, July 12, 2005, leg. J. Kiszka & R. Kościelniak
(KRAP 930). FG-69 – Beskidy Lesiste Mountains, West Bieszczady Mountains; Bieszczady
National Park, Brzegi Górne village, valley of Prowcza Stream, near a quarry, sandstone,
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July 16, 2005, leg. R. Kościelniak (KRAP 1875). GG-60 – Beskidy Lesiste Mountains, West
Bieszczady Mountains: Bieszczady National Park, Tarnica Mount, sandstone, July 18, 2005,
leg. J. Kiszka (KRAP 1876). GG-70 – Beskidy Lesiste Mountains, West Bieszczady Mountains:
Bieszczady National Park, rock outcrops S of Rozsypaniec Mount, near tourist trail, July 7,
2004, leg. J. Kiszka (KRAP 1869).
Additional material examined. AUSTRIA: 1878, (WRSL). GREAT BRITAIN: Westmorland
(V.C69), Great Asby Scar National Nature Reserve, grid ref. 35/65.10 and 64.10., on acidic
rock, August 30, 2006, leg. B. Krzewicka (KRAM-L 50190). CZECH REPUBLIC: Southern
Moravia, Brno Venkov District, Dolní Loučky-Mezihoří, boulder scree flow at N foot of Pásník
hill, below point Bílá skála, schist, April 19, 2007, leg. U. Schiefelbein 2078 (herb. Schiefelbein); Severo-Český kraj, Lužické hory, CHKO Lužické hory, Spravedlnost, April 27, 2003,
leg. U. Schiefelbein 1364 (herb. Schiefelbein). GERMANY: Niedersachsen, Landkreis Leer,
Gem. Westoverledingen, Großwolde, alte Kirche am Ortsrand, brick, September 22, 2004, leg.
U. Schiefelbein 1857 (herb. Schiefelbein); Gem. Leer, Bingum, alte Kirche am Ortsrand, brick,
October 23, 2004, leg. U. Schiefelbein 1864 (herb. Schiefelbein); Gem. Jemgum, Holtgaste, alte
Kirche am Ortsrand, brick, October 23, 2004, leg. U. Schiefelbein 1873 (herb. Schiefelbein).
HUNGARY: 1867, 1872 (WRSL). ITALY: Auf den Stammchen und Zweigen von Rhodod.
Ferrugin in einem Syenitgerolle am Fusse der Margola unweit Predazzo in Sudtirol, September
17, 1878, leg. Arnold (herb. Seaward).
Opegrapha dolomitica (Arnold) Clauzade & Cl. Roux ex Torrente & Egea
Specimens examined. DE-51 – Wyżyna Woźnicko-Wieluńska (Woźniki-Wieluń Upland),
Wyżyna Wieluńska (Wieluń Upland): Węże Nature Reserve, Góra Zelce (Zelce Mountain), on
vertical limestone rocks in slightly shaded habitat, N exposure, May 30, 1963, leg. J. Nowak
(KRAM-L 12349). DE-52 – Wyżyna Woźnicko-Wieluńska (Woźniki-Wieluń Upland), Wyżyna
Wieluńska (Wieluń Upland): Zalesiaki village near Działoszyn town, on slightly sunny limestone
rocks, June 1, 1963, leg. J. Nowak (KRAM-L 12158). DE-62 – Wyżyna Woźnicko-Wieluńska
(Woźniki-Wieluń Upland), Wyżyna Wieluńska (Wieluń Upland): Kamyk village near Kłobuck
town, on vertical limestone rock, NW exposure, May 21, 1964, leg. J. Nowak (KRAM-L 12273).
DF-58 – Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Wyżyna
Olkuska (Olkusz Upland): Dolina Kobylańska (Kobylańska Valley), limestone, slope E, August
6, 1957, leg. J. Nowak (KRAM-L 4131); between Rudawa and Nielepice villages, of Jurassic
limestone, slope N, October 9, 1971, leg. J. Nowak (KRAM-L 29981); Dolina Sąspowska
(Sąspowska Valley), rocks, October 10, 1956, leg. K. Glanc (KRAM-L 39406). DF-68 – Wyżyna
Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Garb Tenczyński (Tenczynek
Ridge): Kamyk near Nielepice villages, vertical rock of Jurassic limenstone, July 30, 1990, leg.
J. Nowak (KRAM-L 34351); September 30, 1986, leg. J. Nowak (KRAM-L 30447). DF-96 –
Pogórze Zachodniobeskidzkie (West Beskids Foothills), Beskid Mały Mountains: Żurawnica
Mount, sandstone, 1961, leg. J. Nowak (KRAM-L 9661). DG-58 – Łańcuch Tatrzański
Mountains, Western Tatras: Dolina Chochołowska (Chochołów Valley), August 21, 1956, leg.
Z. Tobolewski (POZ); Dolina Białego (Valley of the White), near tourist trail, September 10,
1955, leg. Z. Tobolewski (POZ); Dolina Kościeliska (Kościeliska Valley), near road from Hala
Pisana to Hala Ornak, June 22, 1954, leg. Z. Tobolewski (POZ); Tatras gorge, June 23, 1954, leg.
Z. Tobolewski (POZ). EE-82 − Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
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Chęciny Forest Division, Polichno village, limestone, October 21, 1986, leg. K. Toborowicz
(KTC). EE-83 – Wyżyna Kielecka (Kielce Upland), Dolina Nidy (Nida River Valley): Chęciny
Forest Division, Tokarnia village near Chęciny town, June 12, 1975, leg. K. Toborowicz (KTC);
Zelejowa Mount, N slope, August 14, 1976, leg. K. Toborowicz (KTC). EG-20 – West Beskids,
Gorce Mountains: Goła Góra Mount, N slope, June 27, 1955, leg. Z. Tobolewski (POZ); Nowy
Targ County, Białe Skały, August 19, 1955, leg. Z. Tobolewski (POZ). EG-32 – Obniżenie
Orawsko-Podhalańskie (Orava-Podhale Depression), Pieniny Mountains: Skalice Nowotarskie, rock outcrop above Falsztyn village, limestone, August 12, 1968, leg. J. Kiszka (KRAP
11101). EG-33 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Pieniny
Mountains: Wąwóz Szopczański [Sobczański] (Szopczański Gorge), limestone, August 11,
1967, leg. K. Glanc (KRAM-L 39407); Podskalnia Góra (Podskalnia Mountain), limestone,
August 4, 1955, leg. Z. Tobolewski (POZ); Wąwóz Szopczański [Sobczański] (Szopczański
Gorge), limestone, June 30, 1955, leg. Z. Tobolewski (POZ); limestone, April 6, 1954, leg.
Z. Tobolewski (POZ). EG-34 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression),
Pieniny Mountains: Valley of Krupianka Stream, Wąwóz Homole (Homole Gorge), July 28,
1955, leg. Z. Tobolewski (POZ); right slope of Wąwóz Homole (Homole Gorge), September
16, 1956, leg. Z. Tobolewski (POZ); Pieninki Mountains, Czerszlowe Skałki, September 18,
1969, leg. J. Nowak (LOD).
Additional material examined. GREAT BRITAIN: Westmorland, Great Asby Scar National
Nature Reserve, grid ref. 35/65.10 and 64.10, on limestone, August 30, 2000, leg. B. Krzewicka
(KRAM-L 50189). RUSSIA: Orel region, T. 301, Krasnozorensky Distr., near v. Elagino,
bank of Lubovsha River, limestone rocks in deciduous forest, on limenstone, 52°50.438' N,
37°33.978' E, alt. 182 m, July 9, 2012, leg. E. E. Muchnik 11474 (OHHI). SWITZERLAND:
1876, (WRSL).
Exsiccata seen. Weber, Lich. Exs. No. 198, as. Opegrapha saxicola (KRAM); Poelt & Steiner,
Lich. Alp. No. 331, as Opegrapha saxicola (KRAM).
*Opegrapha geographicola (Arnold) Hafellner
Specimens examined. EG-60 – Łańcuch Tatrzański Mountains, Eastern Tatras: High Tatras,
Rysy Mount, the subnival belt; alt. 2,160 m, on granite rock, on thallus of Rhizocarpon badioatrum, July 20, 2004, leg. A. Flakus 2740/1 (KRAM-L 15542, herb. Flakus).
Opegrapha lithyrga Ach.
Specimens examined. DG-05 – Pogórze Zachodniobeskidzkie (West Beskids Foothills),
Beskid Mały Mountains: Czarne Działy hill near Ślemień village, sandstone, August 6, 1965,
leg. J. Nowak (KRAM-L 5059). DG-34 – West Beskids, Beskid Żywiecki Mountains: Wielka
Racza Massif, Zimna Woda forest, on slopes of Smerkówka Mała Mount, rock, August 14,
1964, leg. J. Nowak (KRAM-L 13909). EG-50 – Łańcuch Tatrzański Mountains, Eastern
Tatras: in forest near glade under the rock, above Dolina Roztoki (Roztoka Stream Valley),
sandstone, October 17, 1966, leg. J. Nowak (KRAM-L 5557).
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Additional material examined. CZECH REPUBLIC: Třebíč in decl. collis drinora hora in
vallem fluvii Jihlanka limestone, 1929, leg. J. Suza (LBJS).
Opegrapha niveoatra (Borrer) J. R. Laundon
Specimens examined. BA-87 – Pobrzeże Koszalińskie (Koszalin Coastland), Równina Słupska
(Słupsk Plain): Wodzisław town, S of Sławno town, Fagus sylvatica, August 28, 1987, leg.
W. Fałtynowicz & J. Miądlikowska (UGDA-L 3653). BA-98 − Pobrzeże Koszalińskie (Koszalin
Coastland), Równina Słupska (Słupsk Plain): Chomiec village near Janiewie village, Quercus
sp., August 27, 1987, leg. W. Fałtynowicz & J. Miądlikowska (UGDA-L 3673). BA-91 – Pobrzeże
Koszalińskie (Koszalin Coastland), Wybrzeże Słowińskie (Slovincian Coast): 5 km E of Ustronie Morskie village, Salix sp., June 15, 1986, leg. W. Fałtynowicz (UGDA-L 2791). BA-97 –
Pobrzeże Koszalińskie (Koszalin Coastland), Równina Słupska (Słupsk Plain): Drzeńsko
village, Fagus sylvatica, August 28, 1987, leg. W. Fałtynowicz & J. Miądlikowska (UGDA-L 3947).
BB-29 – Pojezierze Zachodniopomorskie (West Pomeranian Lake District), Pojezierze Bytowskie (Bytowskie Lake District): Krzeszewo village, Quercus sp., August 29, 1987, leg.
W. Fałtynowicz & J. Miądlikowska (UGDA-L 3657). BC-22 – Pojezierze Południowopomorskie
(South Pomeranian Lake District), Równina Drawska (Drawno Plain): Drawieński Park
Narodowy (Drawa National Park), Wydrzy Głaz Nature Reserve, 3 km N of Moczele village,
Fagus sylvatica, May 8, 2010, leg. U. Schiefelbein 2720 (herb. Schiefelbein). CA-58 – Pojezierze
Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian
Lake District): Reda town, Pieleszewo District, Fagus sylvatica, May 1, 1985, leg. W. Fałtynowicz
(UGDA-L 3435). CA-66 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District),
Pojezierze Kaszubskie (Kashubian Lake District): Porzecze village, S of Strzebielina village,
Fagus sylvatica, July 15, 1985, leg. W. Fałtynowicz (UGDA-L 2872). CA-86 – Pojezierze
Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian
Lake District): near Staniszewskie Zdroje Nature Reserve, Fagus sylvatica, April 4, 1986, leg.
W. Fałtynowicz (UGDA-L 3377). CA-99 – Pojezierze Wschodniopomorskie (East Pomeranian
Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Jar Reknicy Nature Reserve,
Fagus sylvatica, September 19, 1983, leg. W. Fałtynowicz & Z. Tobolewski (UGDA-L 1755).
CB-04 – Pojezierze Zachodniopomorskie (West Pomeranian Lake District), Pojezierze Bytowskie (Bytowskie Lake District): Glinowo Forest District, Fagus sylvatica, April 3, 1986,
leg. W. Fałtynowicz (UGDA-L 3448); Quercus sp., April 3, 1986, leg. W. Fałtynowicz &
Z. Tobolewski (UGDA-L 3378). CB-08 – Pojezierze Wschodniopomorskie (East Pomeranian
Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Przywidz village, Fagus
sylvatica, October 25, 1985, leg. W. Fałtynowicz (UGDA-L 2952). DA-85 – Pobrzeże Gdańskie
(Gdańsk Coastland), Mierzeja Wiślana (Vistula Spit): Mierzeja Wiślana Landscape Park, Buki
Mierzei Wiślanej Nature Reserve, beech forest with small patches of black alder bog forest
around small ponds, on Quercus robur, April 21, 2013, leg. M. Kukwa 12078 (UGDA-L19246).
DF-48 – Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa Upland), Wyżyna
Olkuska (Olkusz Upland): Ojców National Park, Wąwóz Jamki (Jamki Gorge), Acer pseudoplatanus, September 22, 2005, leg. J. Kiszka (KRAP). DG-34 – West Beskids, Beskid Żywiecki
Mountains: Oszast forest reserve, Oszus Mount, slope N, Acer pseudoplatanus, September 4,
1964, leg. J. Nowak (KRAM-L 14871). DG-49 – Obniżenie Orawsko-Podhalańskie (OravaPodhale Depression), Rów Podtatrzański (Podtatrzański Trench): Gubałówka, Abies alba,
September 19, 1964, leg. J. Kiszka (KRAP); tree stump, August 18, 1964, leg. J. Kiszka, (KRAP);
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Abies alba, September 16, 1964, leg. J. Kiszka (KRAP). EB-26 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): ca. 2 km E of Otry village,
Fagus sylvatica, May 8, 1990, leg. S. Cieśliński (KTC). ED-99 – Nizina Środkowomazowiecka
(Central Mazovian Lowland), Równina Kozienicka (Kozienice Plain): Puszcza Kozienicka
(Kozienice Forest), Zagożdżon Nature Reserve, Carpinus betulus, October 12, 1969, leg.
S. Cieśliński (KTC). EE-65 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
Dolina Wilkowska (Wilkowska), forest section No. 266, Alnus glutinosa, September 23, 2000,
leg. A. Donica (KTC). EE-66 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
Świętokrzyski National Park, Miejska Góra, forest section No. 15, Quercus robur, September
5, 1999, leg. A. Donica (KTC); forest section No. 26, September 5, 1999, leg. A. Donica (KTC).
EE-74 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains: Kielce Forest Division,
forest section No. 6, Quercus sp., 1986, leg. Bidzyński (KTC). EE-76 – Wyżyna Kielecka
(Kielce Upland), Świętokrzyskie Mountains: Św. Katarzyna Forest District, forest section No.
138, Sorbus aucuparia, September 25, 2000, leg. A. Donica (KTC); Acer pseudoplatanus,
September 25, 2000, leg. A. Donica (KTC); forest section No. 103, Carpinus betulus, leg.
A. Donica (KTC); forest section No. 142, Acer pseudoplatanus, August 10, 2000, leg. A. Donica
(KTC). EE-77 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains: Świętokrzyski
National Park, Nowa Słupia village, Dębno Forest District, forest section No. 125, Acer
platanoides, July 19, 2001, leg. A. Donica (KTC); Łysa Góra (Bald Mountain), forest section
No. 118, Acer sp., July 4, 2000, leg. A. Donica (KTC); Św Krzyż Forest District, Łysa Góra
(Bald Mountain), forest section No. 117, Fagus sylvatica, July 3, 2000, leg. A. Donica (KTC);
Acer pseudoplatanus, July 3, 2000, leg. A. Donica (KTC); forest section No. 115, Acer pseudoplatanus, leg. A. Donica (KTC). EG-32 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale
Depression), Pieniny Mountains: Pieniny Czorsztyńskie Mountains, Abies alba, September
20, 1993, leg. J. Kiszka (KRAP); Zielone Skałki (Green Rocks), Tilia cordata, May 29, 1970,
leg. J. Kiszka (KRAP 10726, 10678). EG-41 – Obniżenie Orawsko-Podhalańskie (OravaPodhale Depression), Pogórze Spisko-Gubałowskie (Spisz-Gubałówka Foothills): Abies alba,
June 19, 1970, leg. J. Kiszka (KRAP 10940, 10902). FB-13 – Pojezierze Mazurskie (Masurian
Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza Borecka (Borki Forest), Quercus
sp., May 17, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC); Borki Nature Reserve, forest
section No. 20, Tilia cordata, May 19, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC). FG21– Beskidy Środkowe Mountains, Beskid Niski Mountains: Magurski National Park, Hucisko
forest, ca. 2.5 km SSW of Huta Krempska settlement, September 11, 2009, leg. M. Kukwa
7653, K. Szczepańska (UGDA-L 15904). FG-58 – Beskidy Lesiste Mountains, SanockoTurczyńskie Mountains: Bieszczady National Park, Suche Rzeki (part of Zatwarnica village),
NW slope of Wierch Mountain, Fagus sylvatica, September 3, 2001, leg. J. Kiszka (KRAP
1881). FG-68 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Bieszczady National
Park, Dolina Moczarnego (Moczarne Valley), between Lutowy and Tarnica streams, Acer
pseudoplatanus, August 10, 2001, leg. R. Kościelniak (KRAP 1880). FG-69 – Beskidy Lesiste
Mountains, West Bieszczady Mountains: Bieszczady National Park, valley of Rzeczyca Stream,
Fagus sylvatica, July 9, 2003, leg. R. Kościelniak (KRAP 1878); Widełki near Bereżki villages,
Acer pseudoplatanus, July 15, 2001, leg. J. Kiszka (KRAP 1877); valley of Rzeczyca Stream,
Acer pseudoplatanus, July 11, 2009, leg. R. Kościelniak (KRAP 4334). GB-10 – Pojezierze
Litewskie (Lithuanian Lake District), Równina Augustowska (Augustów Plain): Wigry National
Park, Fraxinus excelsior, September 1984, leg. W. Fałtynowicz (UGDA-L 2772). GB-40 –
Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska (Augustów Plain):
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Puszcza Augustowska (Augustów Primeval Forest), forest section No. 153, Quercus sp.,
September 21, 1989, leg. S. Cieśliński (KRAM-L 31963). GB-91 – Nizina Północnopodlaska
(North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza
Knyszyńska (Knyszyn Forest), 1 km NWW of Machnacz village, Carpinus betulus, 1987, leg.
Z. Tobolewski & K. Glanc (KTC). GB-92 – Nizina Północnopodlaska (North Podlachian
Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn
Forest), Budzisk Nature Reserve, Corylus avellana, May 13, 1993, leg. K. Kolanko (University
of Bialystok, Institute of Biology). GC-02 – Nizina Północnopodlaska (North Podlachian
Lowland), Wysoczyzna Białostocka (Białystok Highland): 4 km SW of Sokołda village, near
road Sokołda–Supraśl, Carpinus betulus, August 4, 1994, leg. S. Cieśliński (KTC). GC-12 –
Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna Białostocka (Białystok
Highland): Puszcza Knyszyńska (Knyszyn Forest), Stare Biele Nature Reserve, plots 7, 8,
s.coll. (University of Bialystok, Institute of Biology). GC-45 – Nizina Północnopodlaska
(North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Browsk Forest District, forest section No. 101b, plot 216, Acer platanoides,
1983, leg. S. Cieśliński & Z. Tobolewski (KTC). GC-54 – Nizina Północnopodlaska (North
Podlachian Lowland), Równina Bielska (Bielsk Plain): Hajnówka Forest District, forest section
No. 246b, plot 374, Alnus glutinosa, 1983, leg. S. Cieśliński & Z. Tobolewski (KTC); forest
section No. 333b, plot 154, Fraxinus excelsior, 1983, leg. S. Cieśliński & Z. Tobolewski (KTC).
GC-55 − Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk
Plain): Białowieża National Park; forest section No. 256, Carpinus betulus, 1988, leg. S. Cieśliński,
(KTC); Alnus glutinosa, 1988, leg. S. Cieśliński (KTC); forest section No. 320b/d, Carpinus
betulus, 1985, leg. S. Cieśliński & Z. Tobolewski (KTC); plot 341c, Tilia cordata, 1984, leg.
S. Cieśliński & Z. Tobolewski (KTC); Białowieża, Fraxinus sp., September 14, 1953, leg.
J. Rydzak (LBL); forest section No. 256, on Thelotrema lepadinum growing on Quercus robur,
August 2014, leg. M. Kukwa 12877, A. Łubek (UGDA-L 20426); on bark of snag (Fraxinus
excelsior), October 2014, leg. M. Kukwa 13060, A. Łubek (UGDA-L 20570); Tilia cordata,
August 2014, leg. M. Kukwa 13972, 14102, A. Łubek (UGDA-L 21590, 21688); May 2014,
leg. M. Kukwa 12866, A. Łubek (UGDA-L 20418); Alnus glutinosa, October 2014, leg. M. Kukwa
14199, 14259, A. Łubek (UGDA-L 22068, 22116); August 2014, leg. M. Kukwa 13841, 14047,
A. Łubek (UGDA-L 21506, 21639); Fraxinus excelsior, October 2014, leg. M. Kukwa 13070,
A. Łubek (UGDA-L 20578); Quercus robur, August 2014, leg. M. Kukwa 13872, A. Łubek
(UGDA-L 21526); Carpinus betulus, August 2014, leg. M. Kukwa 13859, A. Łubek (UGDAL 21518); May 2014, leg. M. Kukwa 13802, A. Łubek (UGDA-L 21479). GC-65 − Nizina
Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk Plain): forest section
No. 607b/d, Quercus robur, 1983, leg. S. Cieśliński & Z. Tobolewski (KTC). GC-66 – Nizina
Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza
Białowieska (Białowieża Primeval Forest), Białowieża Forest District, Białowieża Forest Division, forest section No. 429, Tilia cordata, 1981, leg. S. Cieśliński & Z. Tobolewski (KTC).
GC-75 − Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk
Plain): forest section No. 673a, Carpinus betulus, 1982, leg. S. Cieśliński & Z. Tobolewski
(KTC). GG-60 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Bieszczady National
Park, Tarnica Mount, valley of Zwór Stream, Fagus sylvatica, October 2, 2000, leg. J. Kiszka
(KRAP 1879).
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Additional material examined. BELARUS: Puszcza Białowieska (Białowieża Primeval
Forest), Kamienivki village, Acer sp., 2000, leg. S. Cieśliński (KTC). CZECH REPUBLIC:
South Moravia, Jihomoravský kraj, S of Lanžhot, Cahnov-Soutok National Nature Reserve,
the protected area Cahnov, elev. 152 m, Fraxinus angustifolia, April 1, 2014, leg. M. Kukwa
12403 (UGDA-L 20009); Polinský vršek, elev. 175 m, edge of a managed floodplain forest,
ca. 80 years old, few isolated trees in an adjacent meadow, Carpinus betulus, May 31, 2014,
leg. M. Kukwa 12362 (UGDA-L 19967). DENMARK: Bornholm, Dondalen, forest, Carpinus betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek). LITHUANIA:
Kaišiadorių raj., Laukystos žemupys, 500 m aukščiau Jonavos-Žąslių plento, Laukystos slėnio
dešinysis šlaitas, Betula sp., October 23, 2005, leg. J. Motiejūnaitė 8455 (BILAS); Quercus robur,
October 13, 2005, leg. D. Stončius 8387 (BILAS); Vilniaus raj., Taurų g-ja, 46 kv., 26 skl., ant
lapuocio medzio kamieno, June 10, 2003, leg. T. Navsutis 10047 (BILAS); Brižų giria, Alnus
sp., September 16, 1992, leg. J. Motiejūnaitė 1484 (BILAS); Nemuno kilpų regioninis parkas,
Alytaus raj., Punios šilas, 17 kv., 1 skl., Acer platanoides, June 14, 2006, leg. J. Motiejūnaitė 7681
(BILAS); Joniškio raj., Endriškių g-ja, 73, kv., 18 skl., Fraxinus excelsior, September 2003, leg.
Ž. Sinkevičius 8385 (BILAS); 36 kv., 1 skl., Quercus robur, September. 2003, leg. Ž. Sinkevičius
8386 (BILAS); Akmenės raj., Naujosios Akmenės g-ja, 438 kv., 16 skl., ant lapuocio medzio
kamieno, September 2013, leg. E. Panovienė 8485 (BILAS); Neries RP, Trakų raj., Panerių
miškas, Lentvario g-ja, 10 kv., 3 skl., dešinysis Saidės slėnio šlaitas, Quercus robur, October
14, 2006, leg. D. Stončius 9473 (BILAS); Sartų RP, Dusetų giria, Vasynos rezervatas, 25 kv.,
Tilia sp., June 10, 2003, leg. J. Motiejūnaitė, 6793, 6557 (BILAS); Dusetų giria, 17 kv., 4 skl.,
Tilia sp., August 18, 2003, leg. J. Motiejūnaitė 680 (BILAS); Raseinių m. ū., Paliepių g-ja,
7 kv., 10 skl., Quercus robur, September 18, 2003, leg. A. Masaitis 6270 (BILAS); Raseinių raj.,
Padubysio g-ja, 3 Masaitis A. 6 kv. 17 skl., Quercus robur, October 21, 2003, leg. A. Masaitis
6357 (BILAS). GERMANY: Mecklenburg-Vorpommern, Vorpommern-Rügen, Isle of Rügen,
coastal cliff N of Nardewitz, ca. 0.2 km NW of Rugeshus, alt. 5 m Fraxinus excelsior, October 25,
2013, leg. U. Schiefelbein 3791 (herb. Schiefelbein); Landkreis Ostvorpommern, Greifswalder
Oie, Obstbäume am Leuchtturm, Malus sylvestris, August 11, 2007, leg. U. Schiefelbein 2122
(herb. Schiefelbein); Landkreis Uecker-Randow, Brohmer Berge, Kleppelshagener Forst,
Einsberg, ca. 1.5 km N of Klepelshagen, Acer pseudoplatanus, leg. U. Schiefelbein 2340 (herb.
Schiefelbein); Münster in Westfalen, Acer campestre and Fraxinus sp., April 1880, leg. Reiss
(herb. Seaward). RUSSIA: Leningrad region, Berezovye Islands, southern part of the island,
Quercus sp., September 14, 2010, leg. U. Schiefelbein 2513 (herb. Schiefelbein).
Exsiccata seen. Arnold, Lich. Exs. No. 1129, as Opegrapha subsiderella (KRAM); Arnold,
Lich. Exs. No. 855, as. Opegrapha subsiderella (KRAM); Poelt & Steiner, Lich. Alp. No. 355,
as Opegrapha subsiderella (KRAM).
Opegrapha saxicola Ach.
Specimens examined. BF-47 – Eastern Sudetes, Masyw Śnieżnika (Śnieżnik Massif): Bialskie
Mountains, Dolina Kleśnicy (valley of Kleśnica Stream), limestone, August 12, 2001, June
27, 2004, leg. K. Szczepańska (herb. Szczepańska). DE-62 – Wyżyna Woźnicko-Wieluńska
(Woźniki-Wieluń Upland), Wyżyna Wieluńska (Wieluń Upland): Biała village near Kłobuck
town, hill W of Kłobuck, alt. 266 m, on vertical limestone rocks, in ± shaded habitats, May
21, 1964, leg. J. Nowak (KRAM-L 12277). DF-37 − Wyżyna Krakowsko-Częstochowska
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(Kraków-Częstochowa Upland), Wyżyna Olkuska (Olkusz Upland): Zubowa Skała near Olkusz
town, limestone, April 7, 1956, leg. J. Nowak (KRAM-L 4136). DF-48 – Wyżyna KrakowskoCzęstochowska (Kraków-Częstochowa Upland), Wyżyna Olkuska (Olkusz Upland): Ojców
National Park, limestone, June 29, 2005, leg. J. Kiszka (KRAP); by the Pradnik–Ojców road,
limestone, August 16, 1956, leg. J. Nowak (KRAM-L 4129). DF-57 − Wyżyna KrakowskoCzęstochowska (Kraków-Częstochowa Upland); Garb Tenczyński (Tenczynek Ridge): Kamyk
village near Rudawa town, limestone, August 9, 1956, leg. J. Nowak (KRAM-L 4135). DF-68
− Wyżyna Krakowsko-Częstochowska (Kraków-Częstochowa Upland); Dolina Sanki (Sanka
River Valley) near Mników village, limestone, August 1956, leg. J. Nowak (KRAM-L 4130).
DF-96 – Beskid Mały Mountains; Żurawnica Mount near Krzeszów, sandstone, August 26,
1962, leg. J. Nowak (KRAM-L 9661). DG-59 – Łańcuch Tatrzański Mountains, Western
Tatras: Dolina Białego (Valley of the White), September 10, 1955, leg. Z. Tobolewski (POZ);
limestone, June 17, 1998, leg. U. Bielczyk (KRAM-L 44572); Dolina Kościeliska (Kościeliska
Valley), limestone, s. coll. (KRAM-L 18017). EE-77 – Wyżyna Kielecka (Kielce Upland),
Świętokrzyskie Mountains: Świętokrzyski National Park, Chełmowa Góra Forest District, on
rock, September 30, 2001, leg. A. Donica (KTC); Łysogóry, on rock, July 1957, leg. B. Halicz
& S. Kuziel (KTC); Święty Krzyż Mount, on rock, July 1957, leg. B. Halicz & S. Kuziel (KTC).
EG-31 − Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Kotlina OrawskoNowotarska (Orava-Nowy Targ Basin): Kramnica Mount, rock, August 15, 1998, leg. J. Kiszka
(KRAM-L 60043). EG-32 − Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression),
Pieniny Mountains: Macelowy Wąwóz (Macelowy Gorge), limestone, September 3, 1998,
leg. J. Kiszka (KRAM-L 60044). EG-33 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale
Depression), Pieniny Mountains: Podskalnia Góra (Podskalnia Mountain), Pieniński Potok
(Pieniny Stream), without substrate, June 10, 1956, leg. Z. Tobolewski (POZ); Zamkowa Góra
(Zamkowa Mount), limestone, May 5, 1957, leg. J. Nowak (KRAM-L 4124); Pieniny National
Park road from Krościenka to Kras, Biała Skała (White Rock), July 19, 1998, leg. J. Kiszka
(KRAM-L 56518). EG-34 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression),
Pieniny Mountains: Krupianka Mount, Wąwóz Homole (Homole Gorge), on rocky slope near
stream, limestone, June 9, 1956, leg. Z. Tobolewski (POZ).
Additional material examined. AUSTRIA: Tirol, 1875, leg. Sander (WRSL). DENMARK:
1867, s. coll. (WRSL). UKRAINE: National Park “Podilskyi tovtry” Kitaihorod, 15 km SE of
Kamianets Podilskyi, Khmelnitskyi region, Kamianets Podilskyi District, alt. 141 m, canyon
of Tarnava River, SW slope, Silurian limestone, June 24, 2004, leg. M. Kukwa 1851a dupl.
(UGDA-L 9493).
Referenced material examined. Opegrapha rupestris – BELGIUM: Mosan District, vicinity
of Nismes, Roche á Lomme, on Bagliettoa calciseda on limestone, November 8, 2008, leg.
M. Kukwa 7280, 7282, D. Ertz, A. Suija (UGDA-L 14839, 14841).
Opegrapha vermicellifera (Kunze) J. R. Laundon
Specimens examined. AB-32 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam and
Wolin islands: Wolin Island, Wolin National Park, forest section No. 60 (plot 35), forest section No. 125 (plot 47), Fagus sylvatica, 1995, leg. A. Wieczorek (herb. Wieczorek). AB-33 –
Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe (Beech Hills): Puszcza Bukowa
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(Beech Forest), Szczecin Landscape Park, forest section No. 184 (plot 23), forest section No.
178 (plot 175), forest section No. 202 (plot 305), Fagus sylvatica, July 20, 1997, leg. A. Wieczorek (herb. Wieczorek). BB-00 – Pobrzeże Szczecińskie (Szczecin Coastland), Równina Gryficka
(Gryfice Plain): Kołobrzeg town, Quercus sp., June 8, 1986, leg. W. Fałtynowicz (UGDA-L 2785).
BC-22 – Pojezierze Południowopomorskie (South Pomeranian Lake District), Równina
Drawska (Drawno Plain): Drawieński Park Narodowy (Drawa National Park), Radęcin Nature
Reserve, 2001, leg. A. Wieczorek (herb. Wieczorek). CA-36 – Pobrzeże Koszalińskie (Koszalin
Coastland), Wybrzeże Słowińskie (Slovincian Coast): near Białogóra village, Kaszebskie
Liszaje (now Babnica) Nature Reserve, decaying wood, July 29, 1982, leg. W. Fałtynowicz
(UGDA-L 2469). CA-79 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District),
Pojezierze Kaszubskie (Kashubian Lake District): Wysoka village, near Wysockie Lake, Ulmus
sp., July 26, 1973, leg. K. Czyżewska (LOD-L 3810). CC-27 – Pradolina Toruńsko-Eberswaldzka
(Toruń-Eberswalder Urstromtal), Kotlina Toruńska (Toruń Basin): Ostromecko village,
Chełmno County, decaying wood, June 16, 1981, leg. Z. Tobolewski (TRN). DA-85 – Pobrzeże
Gdańskie (Gdańsk Coastland), Mierzeja Wiślana (Vistula Spit): Przebrno village, forest section No. 173, Fagus sylvatica, October 17, 1981, leg. E. Budzbon & Z. Tobolewski (UGDAL 2195). DB-53 – Pojezierze Iławskie (Iława Lake District), Szadowo village, Quercus sp.,
April 12, 2004, leg. M. Kukwa 3047 (UGDA-L-10258); March 28, 2005, leg. M. Kukwa 3832
(UGDA-L-11860); Alnus glutinosa and Ulmus scabra, May 15, 2006, leg. M. Kukwa 5133,
5134 (UGDA-L-12789, 12790). DE-68 – Wyżyna Przedborska (Przedborska Upland), Niecka
Włoszczowska (Włoszczowa Basin): Dębowiec Nature Reserve, forest section No. 161, Ulmus
glabra by stream, August 25, 1970, leg. K. Czyżewska (LOD-L 833); forest section No. 162,
Acer platanoides, August 27, 1970, leg. K. Czyżewska (LOD-L 832). EB-67 – Pojezierze
Mazurskie (Masurian Lake District), Równina Mazurska (Masurian Plain): Kuklanka Nature
Reserve, Acer sp., May 1, 1989, leg. S. Cieśliński (KTC). ED-99 – Nizina Środkowomazowiecka
(Central Mazovian Lowland), Równina Kozienicka (Kozienice Plain): Puszcza Kozienicka
(Kozienice Forest), Zagożdżon Nature Reserve, Carpinus betulus, 2002, leg. S. Cieśliński
(KTC). EE-60 – Wyżyna Przedborska (Przedborska Upland), Wzgórza Łopuszańskie
(Łopuszańskie Hills): Oleszno Nature Reserve, forest section No. 53, Acer pseudoplatanus,
July 19, 1974, leg. K. Czyżewska, (LOD-L 16372). EE-66 – Wyżyna Kielecka (Kielce Upland),
Świętokrzyskie Mountains: Świętokrzyski National Park, Podgórze Forest District, forest
section No. 42/43, Tilia cordata, September 5, 1999, leg. A. Donica (KTC). EE-76 – Wyżyna
Kielecka (Kielce Upland), Świętokrzyskie Mountains: Podgórze Forest District, forest section
No. 110, Quercus robur, September 5, 1999, leg. A. Donica (KTC). EF-62 − Brama Krakowska,
Pomost Krakowski: on E from Kraków City, Forest District No. 30, Quercus spp., November
3, 2000, leg. J. Kiszka (KRAP). EG-50 – Łańcuch Tatrzański Mountains, Western Tatras:
Dolina Olczyska (Olcza Valley), Acer pseudoplatanus, May 12, 1998, leg. U. Bielczyk (KRAML 44575); Dolina Białego (Valley of the White), beside tourist trail near valley mouth, limestone,
September 10, 1955, leg. Z. Tobolewski (POZ). FB-68 – Nizina Północnopodlaska (North
Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin): Biebrza National Park, Czerwone
Bagno Nature Reserve, forest section No. 246, Carpinus betulus, September 24, 1986, leg.
S. Cieśliński (KTC); Grzędy forest, Quercus robur, August 11, 2005, leg. J. Motiejūnaite 8388
(BILAS). FG-19 – Beskidy Lesiste Mountains, Sanocko-Turczyńskie Mountains: Pasmo
Turnicy, Fraxinus excelsior, May 22, 1986, leg. J. Piórecki (BDPA); near Arłamów village,
Fraxinus excelsior, May 22, 1986, leg. J. Kiszka & J. Piórecki (KRAM-L 60057). FG-58 – Beskidy
Lesiste Mountains, West Bieszczady Mountains: near road by Hylaty Stream, Abies alba,
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September 3, 2001, leg. J. Kiszka & R. Kościelniak (KRAP 1891). FG-59 – Beskidy Lesiste
Mountains, West Bieszczady Mountains: between Pszczeliny and Stuposiany villages, on SE
from Stuposiany village, Acer pseudoplatanus, July 29, 1959, leg. K. Glanc (KRAM-L 39484);
between Pszczeliny and Jawor villages, Acer pseudoplatanus, July 29, 1959, leg. K. Glanc
(KRAM-L 39481). FG-69 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Widełki
Mount, N slope, Acer pseudoplatanus, July 3, 1959, leg. K. Glanc (KRAM-L 39482). FG-89
− Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl
Foothills): Hołubla Stream, forest, Acer pseudoplatanus, September 27, 1983, leg. J. Kiszka &
J. Nowak (KRAM-L 60059). GB-20 – Nizina Północnopodlaska (North Podlachian Lowland),
Kotlina Biebrzańska (Biebrza Basin): Puszcza Augustowska (Augustów Primeval Forest),
Płaska Forest District, forest section No. 30, Fraxinus excelsior, November 1, 1958, leg. K. Glanc
(KRAM-L 39483). GB-21 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska (Augustów Plain): Perkuć village, Puszcza Augustowska (Augustów Primeval
Forest), Acer sp., September 18, 1986, leg. S. Cieśliński (KTC). GB-30 – Pojezierze Wschodniosuwalskie (East Suwałki Lake District), Równina Augustowska (Augustów Plain): Puszcza
Augustowska (Augustów Primeval Forest), Płaska Forest District, Fraxinus excelsior, November
1, 1958, leg. K. Glanc (POZ). GB-92 – Nizina Północnopodlaska (North Podlachian Lowland),
Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn Forest), Budzisk
Nature Reserve, forest section No. 109i, Quercus robur, June 11, 1999, leg. K. Czyżewska
(LOD-L 11146); Ulmus sp., August 5, 1994, leg. S. Cieśliński (KTC), 5919 (BILAS); Quercus
robur, June 11, 1999, leg. S. Cieśliński 5920 (BILAS). GC-55 – Nizina Północnopodlaska
(North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Białowieża National Park, Carpinus betulus, 1996, leg. S. Cieśliński (KTC);
forest section No. 256d (plot 90), Fraxinus excelsior, 1982, leg. S. Cieśliński & Z. Tobolewski
(KTC); forest section No. 283 (plot 47), Tilia cordata, 1982, leg. S. Cieśliński & Z. Tobolewski
(KTC); forest section No. 316c (plot 92), Quercus robur, 1982, leg. S. Cieśliński & Z. Tobolewski
(KTC); forest section No. 369a (plot 44), Acer platanoides, 1982, leg. S. Cieśliński & Z. Tobolewski (KTC); forest section No. 399b (plot 54), Quercus robur, 1982, leg. S. Cieśliński &
Z. Tobolewski (KTC); forest section No. 256, Acer platanoides, 1982, leg. S. Cieśliński (KTC);
forest section No. 339b, Tilia cordata, 1992, leg. S.Cieśliński (KTC); forest section No. 256,
Tilia cordata, September 16, 1987, leg. K. Czyżewska (LOD-L 8177); Carpinus betulus, September 16, 1987, leg. K. Czyżewska (LOD-L 9039). GC-64 – Nizina Północnopodlaska (North
Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Hajnówka Forest District, forest section No. 570b (plot 232), Fraxinus
excelsior, 1984, leg. S. Cieśliński (KTC). GC-65 – Nizina Północnopodlaska (North Podlachian
Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest),
Białowieża Forest District, forest section No. 609a (plot 231), Fraxinus excelsior, 1983, leg.
S. Cieśliński & Z. Tobolewski (KTC). GC-66 – Nizina Północnopodlaska (North Podlachian
Lowland), Równina Bielska (Bielsk Plain): Białowieża National Park, forest section No. 340a,
Carpinus betulus, August 12, 2002, leg. P. Czarnota (GPN 2946, 2947); ca. 3 km NNW of
Białowieża, forest section No. 340c, Carpinus betulus, May 11, 2006, leg. U. Schiefelbein 2033
(herb. Schiefelbein); ca. 4 km NNW of Białowieża, forest section No. 314c, Fraxinus excelsior,
May 11, 2006, leg. U. Schiefelbein 2047 (herb. Schiefelbein).
Additional material examined. AUSTRIA: ca. 6 km N of Kaprun, Carpinus betulus, April
1, 2005, leg. A. Wieczorek (herb. Wieczorek). DENMARK: Bornholm, Dondalen, forest,
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Carpinus betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek). GERMANY:
Bayern, Lkr. Weilheim: Alpenvorland Paterzeller Eibenwald, südöstlich von Wessobrunn,
atl. 620 m (MTB 8132/1). An loser Borke der Stûämme einzeln stehender Bergahorne Acer
pseudoplatanus, November 19, 1977, leg. T. Feuerer, H. M. Hertel & E. Krach (KRAM-L 25816);
Mecklenburg-Vorpommern, Kreis Uecker-Randow, Wald westlich der Kirche Schmarsow,
October 14, 2001, leg. U. Schiefelbein 741 (herb. Schiefelbein); Kreis Güstrow, May 10, 1998,
leg. U. Schiefelbein 207 (herb. Schiefelbein); Demnitzer Bruch, ca. 1 km E of Charlottenhorst,
Carpinus betulus, May 24, 2006, leg. U. Schiefelbein 2368 (herb. Schiefelbein). LITHUANIA:
Kėdainių raj., Kėdainių m. ū., Skaistgirio g-ja, 42 kvartalas, 1 sklypas, Acer platanoides, October
3, 2008, leg. Ž. Preikša 8762 (BILAS). SWITZERLAND: Kanton Zug, an einer Eichne im
Walde bei Kloster Frauenthal, September. 1886, leg. Hegetschweiler (herb. Seaward).
Exsiccata seen. Arnold, Lich Exs. No. 1186, as Opegrapha hapaleoides (KRAM); Poelt, Lich.
Alp. No. 126, as Opegrapha fuscella (KRAM); Poelt & Steiner, Lich. Alp. No. 307 (KRAM).
Opegrapha vulgata (Ach.) Ach.
Specimens examined. BC-46 – Pradolina Toruńsko-Eberswaldzka (Toruń-Eberswalder
Urstromtal), Kotlina Gorzowska (Gorzów Basin): Noteć District, near Czarnków village,
Alnus glutinosa, March 12, 1967, leg. K. Glanc (KRAM-L 39101). BF-38 – Eastern Sudetes,
Góry Złote (Golden Mountains): Bialskie Mountains, red tourist trail from Gierałtów to
Czernica villages, Acer sp., September 19, 2003, leg. K. Szczepańska (herb. Szczepańska);
Bialska Pętla road, Acer sp., July 29, 2003, leg. K. Szczepańska (herb. Szczepańska); Dolina
Czarnego Potoku (Black Stream Valley), Acer sp., June 25, 2004, leg. K. Szczepańska (herb.
Szczepańska). BF-46 − Eastern Sudetes, Masyw Śnieżnika (Śnieżnik Massif): Bialskie Mountains, Dolina Wilczki (Wilczka Stream Valley), Acer sp., July 10, 2003, leg. K. Szczepańska
(herb. Szczepańska); Jaworek Mount, Acer sp., August 23, 2003, leg. K. Szczepańska (herb.
Szczepańska); road from Idzików to Biała Woda villages, Acer sp., August 25, 2008, leg.
K. Szczepańska (herb. Szczepańska). BF-47 − Eastern Sudetes, Góry Złote (Golden Mountains):
Dolina Kleśnicy (valley of Kleśnica Stream), Acer sp., June 30, 2001, May 25, 2004, leg.
K. Szczepańska (herb. Szczepańska); Nowa Morawa Nature Reserve, Acer sp., July 7, 2003,
leg. K. Szczepańska (herb. Szczepańska); Dolina Kamienicy (Kamienica River Valley), yellow
tourist trail, Acer sp., August 2, 2003, leg. K. Szczepańska (herb. Szczepańska); tree near
stream, Acer sp., August 2, 2003, leg. K. Szczepańska (herb. Szczepańska). BF-48 – Eastern
Sudetes, Masyw Śnieżnika (Śnieżnik Massif): Bialskie Mountains, forest road, near stream,
Acer sp., August 1, 2003, leg. K. Szczepańska (herb. Szczepańska); Czarna Droga (Black Route),
Fagus sylvatica, July 10, 2003, leg. K. Szczepańska (herb. Szczepańska); valley of the Nowinka
Stream, Acer sp., August 22, 2003, leg. K. Szczepańska (herb. Szczepańska). BF-56 − Eastern
Sudetes, Masyw Śnieżnika (Śnieżnik Massif): road from Jodłowa to Goworów villages, Acer
sp., August 24, 2003, leg. K. Szczepańska (herb. Szczepańska); Dolina Goworówki (Goworówka
Stream Valley), Acer sp., August 24, 2003, leg. K. Szczepańska (herb. Szczepańska). CA-69 –
Pobrzeże Gdańskie (Gdańsk Coastland), Pobrzeże Kaszubskie (Kashubian Coastland): Gdynia
City, Kępa Radłowska Nature Reserve, Quercus sp., August 6, 1983, leg. W. Fałtynowicz
(UGDA-L 1834). CA-86 − Pojezierze Wschodniopomorskie (East Pomeranian Lake District),
Pojezierze Kaszubskie (Kashubian Lake District): Wysoczyzna Staniszewska (Staniszewska
Highland), forest section No. 227, Fagus sylvatica, September 27, 1983, leg. W. Fałtynowicz
© The Author(s) 2018
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(UGDA-L 1761); forest section No. 208, Fagus sylvatica, September 27, 1983, leg. W. Fałtynowicz
(UGDA-L 1781). CA-99 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District),
Pojezierze Kaszubskie (Kashubian Lake District): Jar Reknicy Nature Reserve, Fagus sylvatica,
September 19, 1983, leg. W. Fałtynowicz & E. Budzbon (UGDA-L 1875). CB-16 – Pojezierze
Południowopomorskie (South Pomeranian Lake District), Bory Tucholskie (Tuchola Forest):
Kościerzyna town, near road to Gdynia City, March 24, 1959, leg. Z. Jarocka (TRN). DA-85 –
Pobrzeże Gdańskie (Gdańsk Coastland), Mierzeja Wiślana (Vistula Spit); Przebrno village,
forest section No. 127, Fagus sylvatica, August 6, 1982, leg. E. Budzbon (UGDA-L 2194);
forest section No. 128/129, Quercus sp., August 6, 1981, leg. E. Budzbon (UGDA-L 2191);
decaying wood, July 23, 1982, leg. E. Budzbon (UGDA-L 2193); Krynica Morska town, forest
section No. 122, Acer sp., 1982, leg. E. Budzbon & Z. Tobolewski (UGDA-L 2192). DE-29
− Wyżyna Przedborska (Przedborska Upland), Wzgórza Opoczyńskie (Opoczno Hills):
Puszcza Pilicka (Pilica Forest), Błogie Forest District, Jaksonek Forest District, forest section
No. 374, Quercus robur, associated with Chrysothrix candelaris, in thermophilous oak forest,
October 11, 1974, leg. K. Czyżewska (LOD-L 4520). DE-38 – Wzniesienia
Południowomazowieckie (South Mazovian Hills), Równina Piotrkowska (Piotrków Plain):
Puszcza Pilicka (Pilica Forest), Lubień Forest District, forest section No. 123, Lubień Forest
District, Quercus petraea, June 9, 1971, leg. K. Czyżewska (LOD-L 1648). DE-48 – Wyżyna
Przedborska (Przedborska Upland), Wzgórza Radomszczańskie (Radomsko Hills): Puszcza
Pilicka (Pilica Forest), Bąkowa Góra Forest District, forest section No. 39, Quercus petraea,
in thermophilous oak forest, April 28, 1971, leg. K. Czyżewska (LOD-L 1846). DE-57 – Wyżyna
Przedborska (Przedborska Upland), Wzgórza Radomszczańskie (Radomsko Hills): Puszcza
Pilicka (Pilica Forest), Kobiele Wielkie Nature Reserve, forest section No. 175f, Quercus robur,
associated with Chrysothrix candelaris and Schismatomma pericleum, October 27, 1970, leg.
K. Czyżewska (LOD-L 1647). DF-03 – Pogórze Zachodniobeskidzkie (West Beskids Foothills),
Pogórze Śląskie (Silesian Foothills): Szyndzielnia Mount, Fagus sylvatica, June 25, 1964, leg.
J. Kiszka (KRAP). DF-94 – Pogórze Zachodniobeskidzkie (West Beskids Foothills), Beskid
Mały Mountains: valley stream behind Okrąglak) Acer pseudoplatanus, October 5, 1961, leg.
J. Nowak (KRAM-L 10012), Zwalisko Mount, Fagus sylvatica, May 11, 1960, leg. J. Nowak
(KRAM-L 7289); valley of Cisowy Stream, alt. 600 m, Abies alba, April 12, 1961, leg. J. Nowak
(KRAM-L 7288); valley stream on S Wielki Cisownik Mount, alt. 580 m) Abies alba, April
13, 1961, leg. J. Nowak (KRAM-L 10299, 7287); valley stream on S slope of Czarne Działy
Mount alt. 600 m, Abies alba, April 24, 1961, leg. J. Nowak (KRAM-L 10343); valley stream,
on S slope Potrójna Mount, alt. 630 m, Abies alba, April 14, 1961, leg. J. Nowak (KRAML 10137); Ślemień village by the way to Hucisko, alt. 500 m, Fraxinus excelsior, April 26, 1961,
leg. J. Nowak (KRAM-L 7252). DF-96 – Pogórze Zachodniobeskidzkie (West Beskids Foothills),
Beskid Mały Mountains: Targoszów village, gorge between Zwalisko and Czarna Góra mounts,
alt. 600 m, Abies alba, May 11, 1960, leg. J. Nowak (KRAM-L 6226). DG-12 – West Beskids,
Beskid Śląski Mountains: Barania Góra (Ram Mountain), valley of Biała Wisełka Stream,
Abies alba, October 2, 1975, leg. U. Bielczyk (KRAM-L 44023). DG-16 – West Beskids, Beskid
Żywiecki Mountains: Pasmo Jałowieckie (Jałowieckie Range), Zasepnica Mount, alt. 460 m Acer
pseudoplatanus, September 6, 1965, leg. J. Nowak (KRAM-L 15430); Alnus sp., September 6,
1965, (KRAM-L 15809); Abies alba, September 6, 1965, leg. (KRAM-L 15808); Opaczny Potok
(Opaczny Stream), alt. 800 m, Abies alba, July 14, 1965, leg. J. Nowak (KRAM-L 15779);
Dolina Roztoki (Roztoka Valley), alt. 700 m, Abies alba, August 16, 1965, leg. J. Nowak
(KRAM-L 15768); near Siwcówka village, alt. 580 m, Abies alba, August 16, 1965, leg. J. Nowak
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(KRAM-L 15769); valley of Magurek Mount, Abies alba, July 15, 1965, leg. J. Nowak (KRAML 15788). DG-18 – West Beskids, Beskid Żywiecki Mountains: Pasmo Policy (Polica Range),
valley of Syhlec Stream, alt. 920 m, Abies alba, June 23, 1965, leg. J. Nowak (KRAM-L 37465);
valley of Kamieński Stream, Abies alba, May 26, 1965, leg. J. Nowak (KRAM-L 16211); Sidzina
village, valley of Ciśniawka Stream, Abies alba, May 22, 1965, leg. J. Nowak (KRAM-L 15061).
DG-22 – West Beskids, Beskid Śląski Mountains: Gańczorka Mount, Fagus sylvatica, August
4, 1962, leg. J. Kiszka (KRAP). DG-24 – West Beskids, Beskid Żywiecki Mountains: Pilsko
Massif, Pod Rysianką Nature Reserve, Acer pseudoplatanus, May 23, 1966, leg. J. Nowak
(KRAM-L 17303). DG-25 – West Beskids, Beskid Żywiecki Mountains: Pilsko Massif, valley
stream under Hala Lipowska alp, alt. 800 m, Abies alba, September 6, 1964, leg. J. Nowak
(KRAM-L 14823); valley of Nickulin Stream, Abies alba, August 16, 1964, leg. J. Nowak
(KRAM-L 13952); valley of Straceniec Stream, Abies alba, September 5, 1964, leg. J. Nowak
(KRAM-L 14886); valley of Buczynka Stream, Abies alba, September 17, 1964, leg. J. Nowak
(KRAM-L 14831). DG-26 – West Beskids, Beskid Żywiecki Mountains: Babia Góra Mount,
Markowe Szczawiny alp, Pinus sp., June 24, 1974, leg. U. Bielczyk (KRAM-L 44022). DG-32 –
West Beskids, Beskid Żywiecki Mountains: Wielka Racza Massif, valley of Danielka Stream,
Abies alba, August 8, 1964, (KRAM-L 17213); Beskid Bednarów Mountains, Pinus sp., September 4, 1964, leg. J. Nowak (KRAM-L 14729). DG-33 – West Beskids, Beskid Żywiecki
Mountains: Pilsko Massif, Szeroki Kamieniec place, Abies alba, September 6, 1964, leg.
J. Nowak (KRAM-L 14860); Złatna village, Szeroki Kamieniec place Abies alba, September
6, 1964, leg. J. Nowak (KRAM-L 16456); Pasmo Policy (Polica Range), between Talapka and
Matjasikowa, alt. 650 m, Fagus sylvatica, May 27, 1965, leg. J. Nowak (KRAM-L 16223); valley
of Wężna Stream, alt. 600 m, Abies alba, May 27, 1965, leg. J. Nowak (KRAM-L 17205); Śrubita
Nature Reserve, Abies alba, August 12, 1964, leg. J. Nowak (KRAM-L 14448); valley of
Plaskurówki Stream, Abies alba, August 6, 1964, leg. J. Nowak (KRAM-L 14476); Beskid
Bednarów Mountains, Fagus sylvatica, September 4, 1964, leg. J. Nowak (KRAM-L 14780).
DG-58 − Łańcuch Tatrzański Mountains, Western Tatras: on right hand slope below Polana
Jarząbcza (Jarząbcza Glade), Pinus sp., August 19, 1961, leg. Z. Tobolewski (POZ); Dolina
Chochołowska (Chochołów Valley), near trail end, decaying wood, August 14, 1960, leg.
Z. Tobolewski (POZ); near Ścieżka nad Reglami (trail), above Kamienisty Żleb, Pinus sp.,
August 14, 1960, leg. Z. Tobolewski (POZ). DG-59 – Łańcuch Tatrzański Mountains, Western
Tatras: Dolina Spadowiec (Spadowiec Valley), Fagus sylvatica, April 12, 1999, leg. U. Bielczyk
(KRAM-L 44616); Dolina Strążyska (Strążyska Valley), near trail to Wielka Rówień under
Giewont Mount, Acer pseudoplatanus, August 18, 1959, leg. Z. Tobolewski (POZ); on slope,
Dolina ku Dziurze (Ku Dziurze Valley), Abies alba, September 1959, leg. Z. Tobolewski (POZ).
DG-68 − Łańcuch Tatrzański Mountains, Western Tatras: N slope of Czerwony Wierch
Mountain, Pinus sp., August 18, 1960, leg. Z. Tobolewski (POZ). EA-82 – Nizina Staropruska
(Old Prussian Lowland), Wzniesienia Górowskie (Górowo Hills), ca. 1 km N of Garbniki
village, Tilia cordata, May 8, 1989, leg. S. Cieśliński (KTC); Zamkowa Góra (Zamkowa Mount),
Tilia cordata, May 8, 1989, leg. S. Cieśliński (KTC). EA-83 – Nizina Staropruska (Old Prussian
Lowland), Nizina Sępopolska (Sępopolska Lowland): 1.5 km to Nowa Wieś village, Quercus
robur, May 5, 1989, leg. S. Cieśliński (KTC). EB-13 − Pojezierze Mazurskie (Masurian Lake
District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 2 km SW of Lidzbark Warmiński
town, Alnus glutinosa, September 14, 1989, leg. S. Cieśliński (KTC). EB-22 – Pojezierze
Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District), ca. 1.5 km
NE of Smolajny village, Carpinus betulus, August 23, 1993, leg. S. Cieśliński (KTC): Tilia
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cordata, August 23, 1993, leg. S. Cieśliński (KTC). EB-26 – Pojezierze Mazurskie (Masurian
Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): ca. 2 km E of Otry village,
Fagus sylvatica, May 8, 1990, leg. S. Cieśliński (KTC); 3 km of Bredynki village, Fraxinus
excelsior, May 7, 1990, leg. S. Cieśliński (KTC). EB-32 – Pojezierze Mazurskie (Masurian
Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): Las Warmiński Nature Reserve
on Łyna River, Carpinus betulus, May 28, 1994, leg. S. Cieśliński (KTC, OLTC); Tilia cordata,
August 29, 1993, leg. S. Cieśliński (KTC); ca. 2 km SW of railway station in Cerkiewniki
village, Tilia cordata, September 14, 1989, leg. S. Cieśliński (KTC); 3 km from Dobre Miasto
village, Acer sp., September 14, 1989, leg. S. Cieśliński (KTC). EB-42 – Pojezierze Mazurskie
(Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): Olsztyn City, city
forest, Quercus robur, August 9, 2000, leg. D. Kubiak (OLTC); Carpinus betulus, June 5, 2001,
leg. D. Kubiak (OLTC); Corylus avellana, October 2000, leg. D. Kubiak (OLTC); Acer sp.,
October 21, 2000, leg. D. Kubiak (OLTC); Acer sp., August 3, 2000, leg. D. Kubiak (OLTC);
Aesculus hippocastanum, May 2001, leg. D. Kubiak (OLTC). EB-49 – Pojezierze Mazurskie
(Masurian Lake District), Kraina Wielkich Jezior Mazurskich (Great Masurian Lakes Land):
Masurian Landscape Park, Strzałowo Forest District, forest section No. 99, Quercus robur,
July 1998, leg. D. Kubiak (OLTC). EB-51 − Pojezierze Mazurskie (Masurian Lake District),
Pojezierze Olsztyńskie (Olsztyn Lake District): 3 km SWW of Łupstych village, Quercus robur,
July 21, 1988, leg. S. Cieśliński (KTC); road from Olsztyn City to Gietrzwałd town, ca. 2.5 km
S of Łupstych village, forest section No. 299, Quercus robur, July 21, 1988, leg. S. Cieśliński
(KTC). EB-52 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie
(Olsztyn Lake District): Olsztyn City, housing estate Dajtki, Acer sp., July 24, 2003, leg.
D. Kubiak (OLTC). EB-59 − Pojezierze Mazurskie (Masurian Lake District), Kraina Wielkich
Jezior Mazurskich (Great Masurian Lakes Land): Masurian Landscape Park, Strzałowo Forest
District, forest section No. 99, Quercus robur, July 1998, leg. D. Kubiak (OLTC); Puszcza Piska
(Pisz Forest), Krutynia Nature Reserve, Quercus robur, August 31, 1987, leg. S. Cieśliński
(KTC); Tilia cordata, August 31, 1987, leg. S. Cieśliński (KTC). EB-60 – Pojezierze Mazurskie
(Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 2 km E of railway
station in Stare Jabłonki village, Tilia cordata, September 20, 1994, leg. S. Cieśliński (KTC).
EB-71 − Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn
Lake District): near road, Pawłowo village, Quercus robur, July 20, 1988, leg. S. Cieśliński
(KTC). EB-79 – Pojezierze Mazurskie (Masurian Lake District), Równina Mazurska (Masurian Plain): Puszcza Piska (Pisz Forest), forest section Nos 145/134, ca. 1 km NNW of forester’s lodge Ruczaj, Quercus robur, September 3, 1987, leg. S. Cieśliński (KTC). ED-99 – Nizina
Środkowomazowiecka (Central Mazovian Lowland), Równina Kozienicka (Kozienice Plain):
Puszcza Kozienicka (Kozienice Forest), Zagożdżon Nature Reserve, Carpinus betulus, 2002,
leg. S. Cieśliński (KTC). EE-00 – Wzniesienia Południowomazowieckie (South Mazovian
Hills), Równina Piotrkowska (Piotrków Plain): Puszcza Pilicka (Pilica Forest), Spała Nature
Reserve, forest section No. 286, Acer pseudoplatanus, September 2, 1969, leg. K. Czyżewska
(LOD-L 520). EE-21 – Wyżyna Przedborska (Przedborska Upland), Wzgórza Opoczyńskie
(Opoczno Hills): Białaczów Nature Reserve, forest section No. 101b, Carpinus betulus, in
oak-linden-hornbeam forest, April 20, 2002, leg. K. Czyżewska (LOD-L 11360). EE-55 –
Wyżyna Kielecka (Kielce Upland), Płaskowyż Suchedniowski (Suchedniów Plateau): Bliżyńskie
forests, near Skarżysko-Kamienna town, Świnia Góra Nature Reserve, Quercus petraea, June
6, 1974, leg. K. Czyżewska (LOD-L 3106). EE-65 − Wyżyna Kielecka (Kielce Upland),
Świętokrzyskie Mountains: Pasmo Klonowskie (Klonowskie Range), Bukowa Góra (Bukowa
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Mount), Abies alba, 1980, leg. K. Toborowicz (KTC). EE-86 – Wyżyna Kielecka (Kielce
Upland), Świętokrzyskie Mountains: Cisów Nature Reserve, Quercus sp., August 21, 2000,
leg. Supeł & S. Cieśliński (KTC); Pasmo Orłowińskie (Orłowińskie Range), N slope of Słowiec
Mount, Acer pseudoplatanus, 1981, leg. K. Toborowicz & S. Cieśliński (KTC). EG-01 – West
Beskids, Beskid Wyspowy Mountains: Śnieżnica Mount, Fagus sylvatica, August 11, 1966,
leg. J. Nowak (KRAM-L 4632). EG-10 – West Beskids, Gorce Mountains: Niedźwiedź village,
by Porębianka Stream, Salix sp., June 10, 1999, leg. P. Czarnota (GPN 1954). EG-11 – West
Beskids, Gorce Mountains: N slope of Gorc Porębski glade, Alnus sp., October 4, 1996, leg.
P. Czarnota (GPN 1192); by Gorcowy Potok Stream, N exposure, Fagus sylvatica, October 4,
1996, leg. P. Czarnota (GPN 1159); stream above Stara Huta, valley of Turbacz Stream, SW
exposure, Abies alba, October 26, 1995, leg. P. Czarnota (GPN 569); SW exposure, Fagus
sylvatica, October 26, 1995, leg. P. Czarnota (GPN 391); valley of Gorcowy Potok (Gorcowy
Stream), in its catchment, N exposure, Acer sp., October 4, 1996, leg. P. Czarnota (GPN 1212);
NE slope of Kudłoń Mount, below Adamówka glade, Abies alba, August 25, 1967, leg. K. Glanc
(KRAM-L 39502). EG-20 – West Beskids, Gorce Mountains: under Giecka glade, N exposure,
Fagus sylvatica, November 8, 1994, leg. P. Czarnota (GPN 690); under Obidowiec Mount, NE
exposure, Fagus sylvatica, November 9, 1994, leg. P. Czarnota (GPN 678); under Suhora
Mount, NE exposure, Fagus sylvatica, November 9, 1993, leg. P. Czarnota (GPN 985); plot
25, valley of Olszowy Potok (Olszowy Stream), decaying wood, Abies alba, March 9, 1994,
leg. P. Czarnota (GPN 274); N exposure, Acer sp., March 9, 1994, leg. P. Czarnota (GPN 273);
plot 4, W exposure, Fagus sylvatica, July 5, 1994, leg. P. Czarnota (GPN 986); water gap of
Olszowy Potok (Olszowy Stream), NW exposure, Abies alba, June 26, 1996, leg. P. Czarnota
(GPN 1222); Turbacz Nature Reserve, NW exposure, Fagus sylvatica, June 21, 1994, leg.
P. Czarnota (GPN 768); W exposure, Fagus sylvatica, May 12, 1992, leg. P. Czarnota (GPN
231); slope of Suhora Mount, NE exposure, Fagus sylvatica, July 4, 1994, leg. P. Czarnota
(GPN 591); Pod Suhorą Forest District, Abies alba, May 24, 1995, leg. P. Czarnota (GPN 758);
Suche Buczki Forest District, N exposure, Fagus sylvatica, July 31, 1995, leg. P. Czarnota (GPN
1850); near road in valley of Olszowy Potok (Olszowy Stream), decaying wood, Abies alba,
May 6, 1993, leg. P. Czarnota (GPN 516); under Obidowiec Mount, in Ścisłe Brzyżki Forest
District, in Olszowy Potok (Olszowy Stream) catchment area, Fagus sylvatica, November 9,
1994, leg. P. Czarnota (GPN 695). EG-21 – West Beskids, Gorce Mountains: under Szyja
Turbacza in Dolina Roztoki (Roztoka Stream Valley), E exposure, decaying wood, Fagus
sylvatica, May 26, 1994, leg. P. Czarnota (GPN 1014); Znaki Forest District, N exposure, under
Polana Bieniowe (Bieniowe Glade), Fagus sylvatica, May 26, 1997, leg. P. Czarnota (GPN
1655); Dolina Łopusznej (Łopuszna Valley), plot 115, W exposure, Fagus sylvatica, August
25, 1994, leg. P. Czarnota (GPN 783); Dolina Łopusznej (Łopuszna Valley), S slope of Turbacz
Mount, Abies alba, August 24, 1966, leg. K. Glanc (KRAM-L 39501); Ustępny Stream Valley,
SW exposure, Abies alba, February 20, 1995, leg. P. Czarnota (GPN 237); by Kamienica Stream,
N exposure, Fagus sylvatica, May 15, 1997, leg. P. Czarnota (GPN 1699); SE exposure, Acer
sp., June 11, 1996, November 24, 1992, leg. P. Czarnota (GPN 587); under Przysłop Mount,
S exposure, Abies alba, December 16, 1994, leg. P. Czarnota (GPN 689); under Zrąb on S slope
of Polana Pustak (Pustak Glade), Dolina Kamienicy (Kamienica River Valley), Fagus sylvatica,
June 26, 1997, leg. P. Czarnota (GPN 1642); Polana Stawieniec (Stawieniec Glade), S exposure,
Fagus sylvatica, July 14, 1994, leg. P. Czarnota (GPN 792); in Łopuszanka Stream catchment
area, W exposure, Fagus sylvatica, May 26, 1993, leg. P. Czarnota (GPN 1022); Konina Stream,
under Przełęcz Borek (Borek Pass), N exposure, Acer sp., May 10, 1994, leg. P. Czarnota (GPN
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917); plot 71, under Szyja Turbacza, NE exposure, Fagus sylvatica, June 5, 1994, leg. P. Czarnota
(GPN 994); near blue tourist trail, Dolina Kamienicy (Kamienica River Valley), E exposure,
Abies alba, June 11, 1996, leg. P. Czarnota (GPN 1391); Turbacz Nature Reserve, E exposure,
Fagus sylvatica, May 23, 1996, leg. P. Czarnota (GPN 1230); Nad Huciskiem Forest District,
E exposure, Acer sp., July 11, 1994, leg. P. Czarnota (GPN 383); under Szyja Gorca Porębskiego,
E slope, in Dolina Kamienicy (Kamienica River Valley), Acer pseudoplatanus, December 1,
1997, leg. P. Czarnota (GPN 1838); Pasmo Pewelskie (Pewelskie Range), Wątrobów Potok
(Wątrobów Stream), Abies alba, July 25, 1965, leg. J. Nowak (KRAM-L 15252); Wątrobów
Potok (Wątrobów Stream), Abies alba, leg. J. Nowak (KRAM-L 15263); Dolina Łopusznej
valley, S slope of Turbacz Mount, Abies alba, August 24, 1966, (KRAM-L 39501); Polana pod
Przysłopem, Fagus sylvatica, August 2, 1962, leg. J. Kiszka (KRAP); N slope of Kiczora Mount,
Fagus sylvatica, Abies alba, June 11, 1963, leg. J. Kiszka (KRAP). EG-32 – West Beskids, Gorce
Mountains: valley of Kluszkowianka Stream, Abies alba, September 10, 1967, leg. K. Glanc
(KRAM-L 39498). EG-33 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression),
Pieniny Mountains: Toporzyska village, at forest edge, Abies alba, May 4, 1955, leg. J. Nowak
(KRAM-L 1482); Nowy Targ County, near trail from Wyrobek glade to Zamkowa Góra
(Zamkowa Mount), Abies alba, June 12, 1956, leg. Z. Tobolewski (POZ); Wielka Racza Massif,
Śrubita forest reserve, Fagus sylvatica, August 12, 1964, leg. J. Nowak (KRAM-L 14471);
Pasmo Radziejowej (Radziejowa Range), Abies alba, September 13, 1967, leg. M. Olech (KRA);
valley of Czercz Stream, Abies alba, August 4, 1990, leg. L. Śliwa (KRA); Pasmo Radziejowej
(Radziejowa Range), Baniska Nature Reserve, Fagus sylvatica, July 23, 1990, leg. L. Śliwa
(KRA). EG-34 – West Beskids, Beskid Sądecki Mountains: Pasmo Radziejowej (Radziejowa
Range), Mała Sucha Dolina (Mała Sucha Valley), Abies alba, May 30, 1990, leg. L. Śliwa (KRA);
Dolina Białego (Valley of the White), Abies alba, June 28, 1965, leg. M. Olech (KRA); ca.
300 m from Kamień Św. Kingi, Acer pseudoplatanus, August 18, 1960, leg. J. Nowak (KRAML 9178). EG-36 – West Beskids, Beskid Sądecki Mountains: Żebracze Nature Reserve, Fagus
sylvatica, July 5, 2001, leg. P. Czarnota (GPN 2490). EG-37 – West Beskids, Beskid Sądecki
Mountains: foot of Jaworzynka Mount, Abies alba, August 18, 1960, leg. J. Nowak (KRAML 9186); Pasmo Jaworzyny (Jaworzyna Range), valley of Czercz Stream, Fagus sylvatica, February
21, 1990, leg. L. Śliwa (KRA); valley of Kysców Stream, Abies alba, July 18, 1967, leg. M. Olech
(KRA). EG-50 – Łańcuch Tatrzański Mountains, Eastern Tatras: Dolina Filipka (Filipka
Valley), Fagus sylvatica, May 14, 1998, leg. U. Bielczyk (KRAM-L 44403); near road from
Zakopane town to Morskie Oko Lake, between Przyporniak Stream and Zazadnia glade,
Abies alba, September 21, 1966, leg. Z. Tobolewski (POZ). FA-84 – Pojezierze Litewskie
(Lithuanian Lake District), Puszcza Romincka (Romincka Forest): near road to Jurkiszki
village, Salix sp., September 19, 1992, leg. M. Wojciechowska-Gutowska (University of Bialystok, Institute of Biology); forest section No. 252/253, Fraxinus sp., May 14, 1985, leg.
S. Cieśliński (KTC). FB-13 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Ełckie
(Ełk Lakeland): forest section No. 134, ca. 3 km N of Zakątek Leśny village, Fraxinus sp., May
21, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC); Puszcza Borecka (Borki Forest), forest
section No. 168, near Litygajno Lake, Carpinus betulus, May 21, 1985, leg. S. Cieśliński &
Z. Tobolewski (KTC); forest section No. 104, Carpinus betulus, May 18, 1997, leg. Z. Tobolewski
& S. Ciesliński (KTC); forest section No. 41, Tilia sp., May 20, 1987, leg. Z. Tobolewski &
S. Cieśliński (KTC); forest section No. 48, Acer sp., May 19, 1987, leg. S. Cieśliński & Z. Tobolewski (KTC). FB-58 – Nizina Północnopodlaska (North Podlachian Lowland), Kotlina
Biebrzańska (Biebrza Basin): Czerwone Bagno Nature Reserve, forest section No. 246, Carpinus
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betulus, September 24, 1986, leg. S. Cieśliński (KTC); Tilia cordata, 24 September. 1986, leg.
S. Cieśliński (KTC). FD-45 – Wzniesienia Południowomazowieckie (South Mazovian Hills),
Równina Łukowska (Łuków Plain): Jata Nature Reserve, near Łuków town, Fraxinus sp.,
November 3, 1972, leg. M. Motyka-Zgłobicka (LBL). FG-18 – Pogórze Środkowobeskidzkie
(Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills): crest of Chwaniów
Mount, near tributary of Ropienka Stream, Acer pseudoplatanus, June 3, 1987, leg. Piórecki
(BDPA). FG-39 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Puszcza Bukowa
(Beech Forest), near Wetlina village, by Beskidnik Stream, Acer pseudoplatanus, August 28,
1957, leg. Z. Tobolewski (POZ). FG-68 – Beskidy Lesiste Mountains, West Bieszczady Mountains: by Wielki Lutowy Stream, Puszcza Bukowa (Beech Forest), Abies alba, August 18, 1958,
leg. K. Glanc (KRAM-L 39102); Bieszczady National Park, Moczarne village, Wielki Lutowy
Stream, Acer pseudoplatanus, July 24, 2002, leg. J. Kiszka & R. Kościelniak (KRAP-L 1890);
Dolina Moczarnego (Moczarne Valley), Głuchy Stream under Czerteż Mount, Acer pseudoplatanus, July 5, 2002, leg. J. Kiszka & R. Kościelniak (KRAP-L 1892). GB-31 – Nizina
Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin): Puszcza
Augustowska (Augustów Primeval Forest), Starożyn Nature Reserve, Quercus robur, July 15,
1965, leg. J. Zielińska (WA); forest section No. 95/96, Carpinus betulus, September 21, 1986,
leg. S. Cieśliński (KTC); forest section No. 211/212, Fraxinus excelsior, September 17, 1986,
leg. Z. Tobolewski & S. Cieśliński (KTC). GB-92 – Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn
Forest), Budzisk Nature Reserve, Tilia cordata, August 5, 1994, leg. S. Cieśliński (KTC); Alnus
glutinosa, August 5, 1994, leg. S. Cieśliński (KTC); Tilia cordata, May 13, 1993, leg. K. Kolanko
(University of Bialystok, Institute of Biology); Carpinus betulus, October 21, 1993, leg. K. Kolanko (University of Bialystok, Institute of Biology). GC-02 − Nizina Północnopodlaska
(North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): forest section
No. 72/73, Quercus robur, August 6, 1994, leg. S. Cieśliński (KTC); forest section No. 121/122,
Carpinus betulus, September 8, 1987, leg. K. Toborowicz (KTC). GC-03 – Nizina
Północnopodlaska (North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn Forest), Stare Biele Nature Reserve, Fraxinus excelsior,
1999, leg. S. Cieśliński (KTC); (University of Bialystok, Institute of Biology). GC-44 – Nizina
Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza
Białowieska (Białowieża Primeval Forest), Browsk Forest District, Carpinus betulus, September
13, 1964, leg. J. Rydzak (LBL); Narewka Forest District, Betula sp., September 18, 1964, leg.
J. Rydzak (LBL). GC-54 – Nizina Północnopodlaska (North Podlachian Lowland), Równina
Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest), Zwierzyniec Forest
District, Carpinus betulus, August 25, 1965, leg. J. Rydzak (LBL); forest section No. 257, Betula
sp., 1995, leg. S. Cieśliński (KTC). GC-55 – Nizina Północnopodlaska (North Podlachian
Lowland), Równina Bielska (Bielsk Plain): Białowieża National Park, forest section No. 256,
Pinus sylvestris, 1988, leg. S. Cieśliński (KTC); Corylus avellana, 1988, leg. S. Cieśliński (KTC):
Fraxinus excelsior, 1987, leg. S. Cieśliński (KTC); Tilia cordata, 1988, leg. S. Cieśliński (KTC);
Tilia cordata, June 16, 1988, leg. S. Cieśliński (LOD-L 7479); Carpinus betulus, 1987, 1988,
leg. S. Cieśliński (KTC); forest section No. 340a, Abies sp., August 12, 2002, leg. P. Czarnota
(GPN 3006); forest section No. 343, Carpinus betulus, August 5, 1953, leg. J. Rydzak (LBL).
GC-56 − Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk
Plain): near Dziedzinka village, Carpinus betulus, 1988, leg. S. Cieśliński (KTC). GG-70
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– Beskidy Lesiste Mountains, West Bieszczady Mountains: Bieszczady National Park, Syhłowaty
Stream, Acer pseudoplatanus, July 19, 2001, leg. J. Kiszka & R. Kościelniak (KRAP-L 1889).
Additional material examined. AUSTRIA: ca. 6 km N of Kaprun, Carpinus betulus, April
1, 2005, leg. A. Wieczorek (herb. Wieczorek). BELARUS: Puszcza Białowieska (Białowieża
Primeval Forest), Kamieniwki [Kamenivki], Acer sp., leg. S. Cieśliński (KTC). DENMARK:
Møn, Damsholte, Hjelmmark, Alnus glutinosa, June 15, 1947, leg. M. S. Christiansen (KRAML 29853); Populus sp., June 13, 1947, leg. M. S. Christiansen (BMUC); Populus sp., February
25, 1952, leg. M. S. Christiansen (herb. Seaward); Bornholm, Dondalen, forest, Carpinus
betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek); Ekkodalen, Carpinus
betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek). FRANCE: between
Latrusd Ansergne and Mont-Dore, leg. J. J. Barkman. GERMANY: München, 1957: Allgauer,
Alpen, Bayern, leg. Schroppel, J. Poelt; Mecklenburg-Vorpommern, Kreis Uecker-Randow,
Buchenwald auf dem Butterberg, Fagus sylvatica, September 1, 2001, leg. U. Schiefelbein 712
(herb. Schiefelbein); Kleppelshagener Forst, ca. 1 km S of Gehren, Quercus sp., December 1,
1999, leg. U. Schiefelbein 704 (herb. Schiefelbein); ca. 0.3 km W of Mümmelkensee, Fraxinus
excelsior, December 3, 2002, leg. U. Schiefelbein 1367 (herb. Schiefelbein); Landkreis Ostvorpommern, Ückeritz, Naturschutzgebiet “Wockninsee”, Alnus glutinosa, November 27, 2002,
leg. U. Schiefelbein 1308 (herb. Schiefelbein). GIBRALTAR: N. Lincolnshire, Acer sp., April
4, 1987, leg. M. R. D. Seaward (herb. Seaward); LITHUANIA: Žemaitijos NP., Plungės raj.,
Liepijos miškas, Tilia sp., May 12, 2002, leg. J. Motiejūnaite 5740 (BILAS); Ulmus sp., May
12, 2002, leg. J. Motiejūnaite 5739 (BILAS); Kauno raj., Vytėnų g-ja, 73 kvartalas, 12 sklypas,
Quercus robur, November 3, 2004, leg. D. Stončius 7099 (BILAS); Panemunių RP, Šakių raj.,
Ilguvos apyl., Velniaravio rezervatas, Fraxinus excelsior, June 16, 2007, leg. A. Uselinė 9669
(BILAS); Raseinių raj., Padubysio g-ja, 41 kvartalas, 6 sklypas, Quercus robur, October 21,
2003, leg. A. Masaitis 9472 (BILAS); 42 kvartalas, 7 sklypas, Quercus robur, October 21, 2003,
leg. A. Masaitis 9470 (BILAS); Biržų giria, Spalviškių g-ja, 209 kvartalas, 14 sklypas, ?, June
18, 2003, leg. S. Skuja 9471 (BILAS); Vilniaus raj., Neries RP, Saiės upės slėnis ties Saidžiais,
Fraxinus excelsior, April 21, 2004, leg. J. Motiejūnaite 6466 (BILAS); Sartų RP, Dusetų giria,
Vasynos rezervatas, 25 kvartalas, Fraxinus excelsior, June 10, 2003, leg. J. Motiejūnaite 6560
(BILAS); Joniškio raj., Endriškiu g-ja, 36 kvartalas, 1 sklypas, Quercus robur, September
2003, leg. Ž. Sinkevičius 8462 (BILAS); Liepijos miškas, 37 kvartalas, 4 sklypas Žemaitijos
NP., Quercus robur, September 29, 2001, leg. I. Prigodina 2151 (WI); 19 sklypas, Žemaitijos
NP., Plunges raj., Quercus robur, September 29, 2001, leg. I. Prigodina 1306 (WI); Griekynės
miškas 2.5 km I vakaris Nuo Juodkrantės, Kuršių Nerija, Quercus robur, May 24, 2001,
leg. I. Prigodina 1770 (WI). NORWAY: Stryn village, forest, Abies alba, August 28, 1995,
leg. A. Wieczorek (herb. Wieczorek). RUSSIA: Nizhni Novgorod Oblast, Sharanga region,
Kilemarsky Reserve, Rhe Yuronga River floodplain, spruce-broad-leaved forest, Tilia sp.,
July 1999, leg. S. Bakka (NNSU); dead tree, July 1999, leg. S. Bakka (NNSU); Tonshaevo
region, nature sanctuary, southern taiga forest, areas in Burepolom Forestry qu. 33, 34, 35,
Betula sp., August 2000, leg. A. Shestakova (NNSU). SLOVAKIA: prov. Brezno, Western
Carpathians, Nizke Tatry Mountains, Krpáčovo, ca. 0.4 km SE of hotel Junior, Abies alba,
August 2, 2002, leg. U. Schiefelbein 1152 (herb. Schiefelbein). SWITZERLAND: Kanton
Zug, an Weisstannen bei Bliggenstorf, September 1885, leg. Hegetschweiler (herb. Seaward).
SWEDEN: Västra Götaland, leg. C. Stenholm (WA); Osterskar, Fraxinus sp., June 13, 1947,
leg. M. S. Christiansen, (BMUC).
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Exsiccata seen. Arnold, Lich. Exs. No. 855, as Opegrapha subsiderella (KRAM).
*Phacographa glaucomaria (Nyl.) Hafellner
Specimens examined. EG-60 – Łańcuch Tatrzański Mountains, Eastern Tatras: High Tatras,
Mięguszowiecka Przełęcz Wyżnia, alt. 2,330 m, on thallus of Protoparmelia badia, August 30,
2005, 49°11'12" N, 20°03'48" E, leg. A. Flakus, 5537/1 (herb. Flakus); Mięguszowiecki Szczyt
Czarny, alt. 2,409 m, on granite rock, on thallus of P. badia, August 11, 2003, 49°10'52" N,
20°04'08" E, leg. A. Flakus, s.n. (KRAM-L 52516).
Additional material examined. ITALY: Paneveggio in Sudtirol, Parasitisch auf Lecanora
sordida an Felsklippen der Campiler Schichten auf der Hohe ober dem Rollepasse, July 21,
1887, leg. Arnold (herb. Seaward).
Pseudoschismatomma rufescens (Pers.) Ertz & Tehler
Specimens examined. AB-23 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam and
Wolin islands: Wolin Island, Wolin National Park, Fraxinus sp., November 11, 1996, leg.
A. Wieczorek (herb. Wieczorek); Wolin Island, Warnowo Forest District, Międzyzdroje Forest
District, forest section No. 215, Fagus sylvatica, October 1, 1955, leg. Z. Tobolewski (POZ).
AB-93 – Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe (Beech Hills): Puszcza
Bukowa (Beech Forest), Szczecin Landscape Park, Rozdoły Forest District, near Glinna village,
Fraxinus sp., November 12, 1953, leg. Z. Tobolewski (POZ). AD-35 – Pradolina WarciańskoOdrzańska (Warta-Oder Marginal Valley), Dolina Środkowej Odry (Central Oder Valley):
Retno village, Fagus sylvatica, July 30, 1982, leg. Z. Tobolewski (TRN); Acer sp., July 28, 1985
(TRN). BB-31 – Pobrzeże Szczecińskie (Szczecin Coastland), Równina Gryficka (Gryfice
Plain): E of Sidłowo village, Carpinus betulus, October 18, 1987, leg. W. Fałtynowicz &
J. Miądlikowska (UGDA-L 3747). BF-46 − Eastern Sudetes, Masyw Śnieżnika (Śnieżnik
Massif): Bialskie Mountains, Szklary village, bark, August 25, 2003, leg. K. Szczepańska (herb.
Szczepańska). BF-47 – Eastern Sudetes, Masyw Śnieżnika (Śnieżnik Massif): Bialskie Mountains, forest road, Acer sp., August 1, 2003, leg. K. Szczepańska (herb. Szczepańska); valley of
Kleśnica Stream, Acer sp., April 1, 2001, leg. K. Szczepańska (herb. Szczepańska); Acer sp.,
September 21, 2003, leg. K. Szczepańska (herb. Szczepańska); valley of Równica Stream,
Modrzeńce hill, Acer sp., June 26, 2004, leg. K. Szczepańska (herb. Szczepańska); Dolina
Wilczki (Wilczka Stream Valley), Acer sp., July 10, 2003, leg. K. Szczepańska (herb. Szczepańska).
CA-36 – Pobrzeże Koszalińskie (Koszalin Coastland), Wybrzeże Słowińskie (Slovincian
Coast): Białogóra village, Kaszebskie Liszaje Nature Reserve, Fagus sylvatica, July 29, 1982,
leg. W. Fałtynowicz (UGDA-L 2568). CA-81 – Pojezierze Zachodniopomorskie (West Pomeranian Lake District), Wysoczyzna Polanowska (Polanowska Highland): Dębnica Kaszubska
village, 1978, leg. M. Seaward (herb. Seaward). CA-86 − Pojezierze Wschodniopomorskie
(East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District): 2 km N of
Mirachowo village, Fraxinus sp., April 4, 1986, leg. W. Fałtynowicz & Z. Tobolewski (UGDAL 3408). CA-99 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze
Kaszubskie (Kashubian Lake District): Acer sp., April 14, 1973, leg. W. Fałtynowicz (UGDA);
Jar Reknicy Nature Reserve, ca. 2 km from Kołbudy village, Acer sp., December 5, 1976, leg.
W. Fałtynowicz (UGDA-L 1683); near Reknica River, close to the Kołbudy village, Ulmus sp.,
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December 1, 1974, leg. W. Fałtynowicz (UGDA-L 1853). CB-09 − Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District):
Zaskoczyn village, Aesculus sp., April 5, 1986, leg. W. Fałtynowicz (UGDA-L 3293). CB-53 –
Pojezierze Południowopomorskie (South Pomeranian Lake District), Pojezierze Krajeńskie
(Krajna Lake District): Bory Tucholskie (Tuchola Forest), 5 km from Chojnice village, near
road to Konarzyny village, Quercus sp., August 17, 1980, leg. L. Lipnicki (herb. Lipnicki,
TRN). CB-75 – Pojezierze Południowopomorskie (South Pomeranian Lake District), Równina
Charzykowska (Charzykowy Plain): Bory Tucholskie (Tuchola Forest), near Brda River mouth,
Alnus sp., August 10, 1986, leg. L. Lipnicki (herb. Lipnicki). CC-27 – Pradolina ToruńskoEberswaldzka (Toruń-Eberswalder Urstromtal), Kotlina Toruńska (Toruń Basin): Ostromecko
village, Chełmno County, Carpinus betulus, May 22, 1952, leg. J. Hutorowicz (TRK). CC-80 –
Pojezierze Wielkopolskie (Wielkopolska Lake District), Pojezierze Gnieźnieńskie (Gniezno
Lake District): Zielonka Forest District near Poznań City, forest section No. 3, Salix sp., August
10, 1963, leg. K. Glanc (KRAM-L 39492); forest section No. 111a, Ulmus sp., June 7, 1963,
leg. K. Glanc (KRAM-L 39493). CE-11 – Wał Trzebnicki (Trzebnica Embankment), Wzgórza
Twardogórskie (Twardogóra Hills): Milicz County, Kubryk Forest District, Fraxinus sp., June
27, 1969, leg. K. Glanc (KRAM-L 39490). DA-82 – Pobrzeże Gdańskie (Gdańsk Coastland),
Żuławy Wiślane: near road Gdańsk–Świbno, forest section No. 211, Fraxinus sp., July 29,
1983, leg. W. Fałtynowicz (UGDA-L 2645). DA-85 – Pobrzeże Gdańskie (Gdańsk Coastland),
Mierzeja Wiślana (Vistula Spit): Krynica Morska, forest section No. 21, Salix sp., January 12,
1983, leg. E. Budzbon (UGDA-L 2190). DA-96 – Pobrzeże Gdańskie (Gdańsk Coastland),
Wysoczyzna Elbląska (Elbląg Highland): Kadyny, on Malus domestica, November 30, 2003,
leg. M. Kukwa 2853 (UGDA-L 11071). DB-42 – Pojezierze Południowobałtyckie (South Baltic
Lakeland), Pojezierze Iławskie (Iława Lake District): Kwidzyń town, forest section No. 191,
Malus domestica, April 10, 2004, leg. M. Kukwa 3022 (UGDA-L 10681); Biały Dwór, forest
section No. 244, Malus domestica, s.d., leg. M. Kukwa (UGDA-L 14073). DB-53 – Pojezierze
Południowobałtyckie (South Baltic Lakeland), Pojezierze Iławskie (Iława Lake District): ca.
0.5 km N of Szadowo village, road in mixed forest, Acer platanoides, September 11, 2005, leg.
M. Kukwa 4599 (UGDA-L 12405). DF-48 – Wyżyna Krakowsko-Częstochowska (KrakówCzęstochowa Upland), Wyżyna Olkuska (Olkusz Upland): Ojców village, Wąwóz Korytania
(Korytania Gorge), Carpinus betulus, June 28, 1955, leg. J. Nowak (KRAM-L 2051); Dolina
Sąspowska (Sąspowska Valley), Fagus sylvatica, June 28, 1955, leg. J. Nowak (KRAM-L 4126);
Ojców village, leg. A. Rehmann (KRAM-L 10198). DG-06 − West Beskids, Beskid Żywiecki
Mountains: alt. 500 m, Fraxinus sp., August 16, 1965, leg. J. Nowak (KRAM-L 15776). DG-07
− West Beskids, Beskid Makowski Mountains: alt. 460 m, Acer pseudoplatanus, September
6, 1965, leg. J. Nowak (KRAM-L 15829). DG-15 – West Beskids, Beskid Makowski Mountains:
Pasmo Pewelskie (Pewelskie Range), Janikowa Grapa Mount, alt. 690 m, Fagus sylvatica,
September 17, 1965, leg. J. Nowak (KRAM-L 13511); Kaczanka Stream, alt. 480 m, Fraxinus
sp., s.d., leg. J. Nowak (KRAM-L 17450). DG-16 – West Beskids, Beskid Żywiecki Mountains:
Pasmo Jałowieckie (Jałowieckie Range), Dolina Bystra (Bystra Valley), Acer pseudoplatanus,
July 24, 1965, leg. J. Nowak (KRAM-L 15761). DG-17 – West Beskids, Beskid Żywiecki
Mountains: Pasmo Policy (Polica Range), Osielec village, valley of Baranów Potok (Baranów
Stream), Salix sp., May 23, 1965, (KRAM-L 16213, 16215). DG-24 – West Beskids, Beskid
Żywiecki Mountains: Masyw Pilska (Pilsko Massif), Pod Rysianką Nature Reserve, Fagus
sylvatica, May 25, 1966, leg. J. Nowak (KRAM-L 17271). DG-33 – West Beskids, Beskid
Żywiecki Mountains: Grupa Wielkiej Raczy Mountains, Bednarów Beskid Mount, Fagus
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sylvatica, September 4, 1964, leg. J. Nowak (KRAM-L 14789). EA-80 – Nizina Staropruska
(Old Prussian Lowland), Wzniesienia Górowskie (Górowo Hills): 2 km SW of Wyszkowo
village, Fraxinus sp., May 9, 1989, leg. S. Cieśliński (KTC). EA-82 – Nizina Staropruska (Old
Prussian Lowland), Wzniesienia Górowskie (Górowo Hills): E slope of Zamkowa Góra
(Zamkowa Mount), Fraxinus sp., May 8, 1989, leg. S. Cieśliński (KTC). EB-04 – Nizina
Staropruska (Old Prussian Lowland), Nizina Sępopolska (Sępopolska Lowland): ca. 2 km
N of Galiny village, Fraxinus sp., May 7, 1989, leg. S. Cieśliński (KTC). EB-06 – Nizina
Staropruska (Old Prussian Lowland), Nizina Sępopolska (Sępopolska Lowland): NW of
Sporwiny village, Ulmus sp., May 6, 1989, leg. S. Cieśliński (KTC). EB-07 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): ca. 1.5 km
NW of Grudniki village, near road Reszel–Korsze, Fraxinus sp., May 4, 1989, leg. S. Cieśliński
(KTC). EB-08 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie
(Olsztyn Lake District): ca. 0.5 km S of Dębiany village, Fraxinus sp., May 6, 1989, leg.
S. Cieśliński (KTC). EB-15 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze
Olsztyńskie (Olsztyn Lake District): ca. 2.5 km SEE of Bisztynek town, Populus sp., May 7,
1989, leg. S. Cieśliński (KTC). EB-19 – Pojezierze Mazurskie (Masurian Lake District), Kraina
Wielkich Jezior Mazurskich (Great Masurian Lakes Land): Gierłoż village, Populus tremula,
September 3, 1988, leg. S. Cieśliński (KTC). EB-22 – Pojezierze Mazurskie (Masurian Lake
District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 4 km NNE of Smolajny village,
Quercus sp., August 23, 1993, leg. S. Cieśliński (KTC); ca. 3.5 km of Dobre Miasto, Fraxinus
sp., September 14, 1989, leg. S. Cieśliński (KTC). EB-36 – Pojezierze Mazurskie (Masurian
Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): ca. 2 km ENE of Stryjewo
village, Fraxinus sp., May 8, 1990, leg. S. Cieśliński (KTC); 1.5 km SE of Oterki village, Acer
sp., May 8, 1990, leg. S. Cieśliński (KTC). EB-52 – Pojezierze Mazurskie (Masurian Lake
District), Pojezierze Olsztyńskie (Olsztyn Lake District): Olsztyn City, Kortowo, Populus sp.,
October 15, 2003, leg. D. Kubiak (OLTC). EB-58 – Pojezierze Mazurskie (Masurian Lake
District), Pojezierze Mrągowskie (Mrągowo Lake District): Szklarnia village, ca. 4.5 km NW
of Piecki village, Populus tremula, May 1, 1989. leg. S. Cieśliński (KTC). EB-59 – Pojezierze
Mazurskie (Masurian Lake District), Równina Mazurska (Masurian Plain): Puszcza Piska
(Pisz Forest), ca. 1.5 km E of Krutyń village, Acer sp., September 2, 1987, leg. S. Cieśliński
(KTC). EB-63 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie
(Olsztyn Lake District), Las Warmiński Nature Reserve, forest section No. 252, Acer sp., May
2001, leg. D. Kubiak (OLTC); Nowe Ranuki Forest District, Las Warmiński Nature Reserve,
Populus sp., May 18, 2001, leg. D. Kubiak (OLTC). EB-67 – Pojezierze Mazurskie (Masurian
Lake District), Równina Mazurska (Masurian Plain): Kuklanka Range, near road Marksady–
Babięta, Fraxinus sp., May 1, 1989, leg. S. Cieśliński (KTC). EB-77 – Pojezierze Mazurskie
(Masurian Lake District), Równina Mazurska (Masurian Plain): ca. 1 km N of Cajki Nowe
village, near road Szczytno–Rozłogi, Fraxinus sp., May 4, 1990, leg. S. Cieśliński (KTC).
ED-09 – Nizina Środkowomazowiecka (Central Mazovian Lowland), Kotlina Warszawska
(Warsaw Basin), Równina Wołomińska (Wołomin Plain): Dębina Nature Reserve, Ulmus sp.,
August 1955, leg. J. Zielińska (WA). ED-12 – Nizina Środkowomazowiecka (Central Mazovian
Lowland), Kotlina Warszawska (Warsaw Basin): Kampinos National Park, near park border,
Quercus sp., September 4, 1962, leg. J. Zielińska (WA); near Gajówka Dębowskie village, Salix
sp., September 1, 1962, leg. J. Zielińska (WA). EE-54 – Wyżyna Kielecka (Kielce Upland),
Płaskowyż Suchedniowski (Suchedniów Plateau): Lasy Bliżyńskie (Bliżyńskie Forests), near
Skarżysko-Kamienna town, Świnia Góra Nature Reserve, Carpinus betulus, May 4, 1959, leg.
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J. Nowak (KRAM-L 2311, 5401). EG-20 – West Beskids, Gorce Mountains: Kowaniec Stream,
Alnus sp., July 21, 1971, leg. K. Glanc (KRAM-L 11832). EG-21 – West Beskids, Gorce
Mountains: under Gorc Porębski glade, near Plot 273, by valley of Gorcowy Potok (Gorcowy
Stream), in Dolina Kamienicy (Kamienica River Valley), Fagus sylvatica, October 4, 1996,
leg. P. Czarnota (GPN 1181/94); Nad Huciskiem Forest District, on E slope of Gorc Porębski
glade, in Dolina Kamienicy (Kamienica River Valley), Acer pseudoplatanus, July 11, 1999,
leg. P. Czarnota (GPN 773/94); S slope of Kudłoń Mount, Acer pseudoplatanus, August 28,
1961, leg. K. Glanc (KRAM-L 39492); S slope of Kudłoń Mount, near route down to Kamienica
Stream, Acer pseudoplatanus, August 28, 1961, leg. K. Glanc (KRAM-L 39488). EG-28 –
Central Beskids, Beskid Niski Mountains: Polana Mount, between Zdynia and Ropa rivers,
Quercus sp., leg. M. Olech (KRA). EG-32 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale
Depression), Pieniny Mountains: Czorsztyn village, Salix sp., August 5, 1968, leg. J. Kiszka
(KRAP); August 10, 1968, leg. J. Kiszka (KRAP). EG-33 – Obniżenie Orawsko-Podhalańskie
(Orava-Podhale Depression), Pieniny Mountains: near trail from Polana Wyrobek to Zamkowa
Góra (Zamkowa Mount), Abies alba, August 8, 1954, leg. Z. Tobolewski (POZ); Pieniński
Potok (Pieniny Stream) valley, near Burzyna glade tourist trail, Fagus sylvatica, June 10, 1956,
leg. Z. Tobolewski (POZ); between Wyrobek glade and Zamkowa Góra (Zamkowa Mount),
Corylus avellana, June 12, 1956, leg. Z. Tobolewski (POZ); by the road from Korścienko to
Zamkowa Góra, Abies alba, May 4, 1955, leg. J. Nowak (KRAM-L 2360). EG-36 – West
Beskids, Beskid Sądecki Mountains: Żebracze Nature Reserve, in its central part, Fagus sylvatica, July 5, 2001, leg. P. Czarnota (GPN 2551). FA-85 – Pojezierze Litewskie (Lithuanian
Lake District), Puszcza Romincka (Romincka Forest): forest section No. 88, Fraxinus sp.,
May 16, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC); Boczki Nature Reserve, forest section
No. 86, Fraxinus sp., May 16, 1985, leg. S. Cieśliński & Z. Tobolewski (KTC); forest section
No. 88, Fraxinus sp., May 16, 1985, leg. S. Cieśliński (KRAM-L 31921). FA-86 – Pojezierze
Litewskie (Lithuanian Lake District), Puszcza Romincka (Romincka Forest): ca. 1 km N of
Bludzie Małe village, forest section No. 273, Fraxinus sp., May 13, 1985, leg. Z. Tobolewski
& S. Cieśliński (KTC); ca. 2.5 km N of Czerwona Struga Nature Reserve, Fraxinus sp., May
14, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC); Fraxinus sp., May 14, 1985, leg. S. Cieśliński
(KRAM-L 31916). FA-87 – Pojezierze Litewskie (Lithuanian Lake District), Puszcza Romnicka
(Romincka Forest): 2.5 km SSW of Żytkiejmy village, Acer sp., May 15, 1985, leg. Z. Tobolewski
& S. Cieśliński (KTC); ca. 1.5 km of Żytkiejmy village, near road to Gołdap town, Acer sp.,
May 25, 1985, leg. S. Cieśliński & Z. Tobolewski (KTC). FA-92 – Pojezierze Mazurskie
(Masurian Lake District), Kraina Węgorapy: ca. 0.5 km S of Lisy village, Fraxinus sp., May
18, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC). FA-97 – Pojezierze Litewskie (Lithuanian
Lake District), Pojezierze Wschodniosuwalskie (East Suwałki Lake District): NW edge of
Hańcza Lake, Salix sp., May 18, 1985, leg. S. Cieśliński & Z. Tobolewski (KTC). FB-00 –
Pojezierze Mazurskie (Masurian Lake District), Kraina Wielkich Jezior Mazurskich (Great
Masurian Lakes Land): Mokre Nature Reserve near Mamry Lake, Fraxinus sp., September 4,
1988, leg. S. Cieśliński (KTC). FB-03 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza Borecka (Borki Forest), forest section No. 195, near
Czerwony Dwór village, Fraxinus sp., May 21, 1985, leg. Z. Tobolewski & S. Cieśliński (KTC);
forest section No. 135, ca. 3 km N of Leśny Zakątek, Fraxinus sp., May 21, 1985, leg. Z. Tobolewski (KTC), ca. 3 km N of Zakątek Leśny, forest section No. 134, Fraxinus sp., May 21,
1985, leg. Z. Tobolewski & S. Cieśliński (KTC). FB-13 − Pojezierze Mazurskie (Masurian
Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza Borecka (Borki Forest), near Litygajno
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Lake, forest section No. 168, Fraxinus sp., May 21, 1985, leg. S. Cieśliński & Z. Tobolewski
(KTC). FB-19 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska
(Augustów Plain): Wigry National Park, near Białe Lake, forest section No. 224, Salix sp.,
September 11, 1984, leg. Z. Tobolewski (UGDA-L 2723). FB-50 – Pojezierze Mazurskie
(Masurian Lake District), Kraina Wielkich Jezior Mazurskich (Great Masurian Lakes Land):
Smolak Duży bog near Popielno and Wierzba villages, on dry stalks of sedge (Carex), August
8, 1978, leg. J. Nowak (KRAM-L 25005). FB-58 – Nizina Północnopodlaska (North Podlachian
Lowland), Kotlina Biebrzańska (Biebrza Basin): Czerwone Bagno Nature Reserve, forest
section No. 246, Fraxinus sp., September 24, 1986, leg. S. Cieśliński (KTC). FB-77 – Nizina
Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin):
Biebrzański National Park, E of Płochowo village, the edge of small deciduous forest by the
road, on Salix sp., April 30, 2005, leg. M. Kukwa 3939 (UGDA-L 11495). FC-05 – Nizina
Północnopodlaska (North Podlachian Lowland), Wysoczyzna Kolneńska (Kolneńska Highland): edge of Męczki village, Salix sp., August 22, 1990, leg. S. Cieśliński (KTC). FD-08 –
Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna Drohiczyńska (Drohiczyn
Highland): ca. 1.5 km W of Zajęczniki village, Salix sp., 9 August.1987, leg. S Cieśliński &
Z. Tobolewski (KTC). FD-56 – Wzniesienia Południowomazowieckie (South Mazovian Hills),
Równina Łukowska (Łuków Plain): Jata Nature Reserve near Łuków town, Alnus sp., May
30, 1972, leg. M. Motyka-Zgłobicka (LBL); Alnus sp., November 3, 1973, leg. M. MotykaZgłobicka & B. Żabińska (LBL). FD-68 – Polesie Zachodnie (Western Polesie), Zaklęsłość
Łomaska (Łomaska Depression): Omelno Nature Reserve, Quercus sp., August 3, 1982, leg.
B. Ordyczyńska (LBL). FD-83 – Nizina Południowopodlaska (South Podlachian Lowland),
Wysoczyzna Żelechowska (Żelechowska Highland): Ryki village by Zalesianka River, Salix
sp., August 18, 1957, leg. B. Wielgosz (LBL); Acer sp., August 18, 1957, leg. B. Wielgosz (LBL);
levee between pond and Ryki River, Alnus glutinosa, August 18, 1957, leg. B. Wielgosz (LBL).
FE-07 – Nizina Południowopodlaska (South Podlachian Lowland), Wysoczyzna Lubartowska
(Lubartów Highland): Lubartów County, Kozłówka Forest District, Nasutów Forest District,
forest section No. 52, Quercus sp., September 12, 1962, leg. L. Jędrzejewska (LBL); Dąbrówka
Forest District, Kozłówka Forest District, Alnus glutinosa, August 28, 1961, leg. L. Jędrzejewska
(LBL); Quercus sp., August 28, 1961, leg. L. Jędrzejewska (LBL); Quercus sp., September 15,
1962, leg. L. Jędrzejewska (LBL). FE-53 – Wyżyna Lubelska (Lublin Upland), Wzniesienia
Urzędowskie (Urzędów Hills): Wólka Kolczyńska village near Józefów on Vistula River, Betula
sp., May 6, 1975, leg. Z. Flisińska (LBL). FE-90 – Wyżyna Kielecka (Kielce Upland), Wyżyna
Sandomierska (Sandomierz Upland): Żyznów Forest District, near Klimontów village, forest
section No. 71, Acer pseudoplatanus, 1985, leg. Dziubek (KTC). FF-36 – Kotlina Sandomierska
(Sandomierz Basin), Płaskowyż Kolbuszowski (Kolbuszowa Plateau): Leżajsk County, Puszcza
Sandomierska (Sandomierz Forest), Jelna Forest District, Alnus sp., July 25, 1963, leg.
B. Ordyczyńska (LBL); FF-37 − Kotlina Sandomierska (Sandomierz Basin), Płaskowyż
Kolbuszowski (Kolbuszowa Plateau): Jelna Forest District, forest section No. 113, Quercus
sp., 1963, leg. B. Ordyczyńska (LBL); forest section No. 104, Betula sp., 1963, leg. B. Ordyczyńska
(LBL); Marynin Forest District, forest section No. 149, Alnus glutinosa, July 25, 1963, leg.
B. Ordyczyńska (LBL); Julin Forest District, forest section No. 241, Quercus sp., 1963, leg.
B. Ordyczyńska (LBL); Brzóza Królewska Forest District, forest section No. 210, Quercus sp.,
1963, leg. B. Ordyczyńska (LBL). FF-99 – Pogórze Środkowobeskidzkie (Central Beskidian
Foothills), Pogórze Dynowskie (Dynowskie Foothills): Krasiczyn village, on the San River,
Salix spp., leg. J. Kiszka & J. Piórecki (KRAM-L 60014). FG-07 − Pogórze Środkowobeskidzkie
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(Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills): Leszczawka village,
near the manor house No. 2068, Salix sp., July 21, 1984, leg. J. Kiszka & J. Piórecki (KRAML 60017). FG-08 − Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze
Przemyskie (Przemyśl Foothills): Leszczawa Górna village No. 2045, Malus domestica, July
21, 1984, leg. J. Kiszka & J. Pióreki (KRAM-L 60021). FG-09 − Pogórze Środkowobeskidzkie
(Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills): Turnica valley,
Fraxinus excelsior, July 27, 1984, leg. J. Kiszka & J. Piórecki (KRAM-L 60026). FG-16 − Góry
Sanocko-Turczańskie (Sanok-Turka Mountains), West Bieszczady Mountains: Wujskie village,
Fraxinus excelsior, July 23, 1984, leg. J. Kiszka & J. Piórecki (KRAM-L 60020). FG-17 − Pogórze
Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills):
Kreców village, Tilia sp., June 5, 1986, leg. J. Kiszka & J. Piórecki (KRAM-L 60022). FG-18 –
Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl
Foothills): crest of Chwaniów Mount, Acer pseudoplatanus, June 3, 1987, leg. J. Piórecki
(BDPA), Wojtkowa village, Fraxinus excelsior, July 23, 1984, leg. J. Kiszka & J. Piórecki
(KRAM-L 60015) FG-19 − Góry Sanocko-Turczańskie (Sanok-Turka Mountains), West
Bieszczady Mountains: hummock under Arłamów village, Salix spp., May 22, 1986, leg.
J. Kiszka & J. Piórecki (KRAM-L 60011). FG-27 – Pogórze Środkowobeskidzkie (Central
Beskidian Foothills), Pogórze Bukowskie (Bukowskie Foothills); Słonne Mountains, Manasterzec village, Fraxinus excelsior, July 8, 1988, leg. J. Kiszka & J. Piórecki (KRAM-L 60016).
FG-28 − Góry Sanocko-Turczańskie (Sanok-Turka Mountains), West Bieszczady Mountains:
Serednica village, No. 103, Fraxinus excelsior, May 4, 1990, leg. J. Kiszka (KRAM-L 60025).
FG-47 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Bieszczady National Park,
ruins of Orthodo church in Bukowiec village, Fraxinus excelsior, January 1, 1997, leg.
R. Kościelniak (KRAP 1883). FG-48 − Beskidy Lesiste Mountains, West Bieszczady Mountains:
the road between Studenne village and Tworylne village, Populus tremula, August 8, 1998,
leg. J. Kiszka (KRAM-L 60040). FG-58 − Beskidy Lesiste Mountains, West Bieszczady Mountains: Berdo village, Salix spp., May 6, 1990, leg. J. Kiszka & J. Piórecki (KRAP-L 60029).
FG-59 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Nasiczne village, by Prowcza
Stream, Acer pseudoplatanus, July 16, 2005, leg. J. Kiszka & R. Kościelniak (KRAP 908);
Fraxinus sp., July 16, 2005, leg. J. Kiszka & R. Kościelniak (KRAP 01884). FG-68 – Beskidy
Lesiste Mountains, West Bieszczady Mountains: Wetlina village, glade near stream, Fraxinus
sp., August 8, 2001, leg. J. Kiszka (KRAP 01882). FG-69 – Beskidy Lesiste Mountains, West
Bieszczady Mountains: near valley of Rzeczyca Stream, Alnus sp., July 7, 1959, leg. K. Glanc
(KRAM-L 39491), Widelki village, No 2036, Acer pseudoplatanus, July 27, 1984, leg. J. Kiszka
& J. Piórecki (KRAM-L 60013). FG-79 – Beskidy Lesiste Mountains, West Bieszczady Mountains: Bieszczady National Park, Wołosate village, Przełęcz Żydowski Beskid (Żydowski Beskid
Pass), Acer pseudoplatanus, July 8, 2004, leg. J. Kiszka (KRAP 909). GB-01 – Nizina
Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin): forest
section No. 20, near road Augustów–Sejny, Salix sp., September 19, 1986, leg. S. Cieśliński
(KTC). GB-10 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska
(Augustów Plain): Wigry National Park, Quercus robur, September 15, 1984, leg. H. Wójciak
(LBL); forest section No. 102, Carpinus betulus, September 1984, leg. K. Górzyńska &
S. Cieśliński (UGDA-L 2710). GB-31 – Nizina Północnopodlaska (North Podlachian Lowland),
Kotlina Biebrzańska (Biebrza Basin): Puszcza Augustowska (Augustów Primeval Forest),
Starożyn Nature Reserve, forest section Nos 211, 212, Fraxinus excelsior, September 17, 1986,
leg. S. Cieśliński (KTC); forest section No. 195, Fraxinus sp., September 17, 1986, leg.
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S. Cieśliński (KRAM-L 31823); Acer sp., July 17, 1965, leg. B. Żabińska (WA). GB-40 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska (Augustów Plain):
Puszcza Augustowska (Augustów Primeval Forest), ca. 1 km NE of Balinka village, forest
section Nos 95, 96, 69, 70, Fraxinus excelsior, September 21, 1986, leg. S. Cieśliński (KTC);
forest section Nos 51, 24, Fraxinus excelsior, September 13, 1986, leg. S. Cieśliński (KRAML 31973); Kozi Rynek Nature Reserve, Fraxinus excelsior, September 22, 1986, leg. S. Cieśliński
(KTC). GB-92 – Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna
Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn Forest), Budzisk Nature
Reserve, Fraxinus excelsior, August 5, 1994, leg. S. Cieśliński (KTC). GC-01 – Nizina
Północnopodlaska (North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Krasne village, near road Supraśl–Białystok, Salix sp., August 3, 1994, leg. S. Cieśliński
(KTC). GC-11 – Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna
Białostocka (Białystok Highland): ca. 3.5 km E of Ciasne village, Fraxinus sp., June 30, 1991,
leg. S. Cieśliński (KTC). GC-20 – Nizina Północnopodlaska (North Podlachian Lowland),
Wysoczyzna Białostocka (Białystok Highland): 0.5 km SE of Niewodnica Korycka village,
Salix sp., August 26, 1991, leg. S. Cieśliński (KTC). GC-35 – Nizina Północnopodlaska (North
Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland): Puszcza Białowieska
(Białowieża Primeval Forest), Browsk Forest District, Browsk Forest Division, forest section
No. 259a, Alnus sp., leg. S. Cieśliński & Z. Tobolewski (KTC); forest section Nos 747b, 748a,
Fraxinus excelsior, 1983, leg. S. Cieśliński & Z. Tobolewski (KTC). GC-55 – Nizina
Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk Plain): Białowieża
National Park, forest section No. 256, Tilia cordata, October 18, 1989, leg. S. Cieśliński (KTC);
forest section No. 256, Quercus robur, 1990, leg. S. Cieśliński (KTC); forest section No. 256,
Fraxinus excelsior, October 16, 1989, leg. S. Cieśliński (KTC); forest section No. 340a, Fraxinus
excelsior, 1982, leg. S. Cieśliński 7 Z. Tobolewski (KTC); forest section No. 256, Carpinus
betulus, August 2014, leg. M. Kukwa 13232, A. Łubek (UGDA-L 20700); May 2015, leg.
M. Kukwa 1511, A. Łubek (UGDA-L 22427). GC-64 – Nizina Północnopodlaska (North
Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Hajnówka Forest District, Leśna Forest Division, forest section No. 575a,
Berezowo Forest District, Fraxinus sp., 1982, leg. Z. Tobolewski & S. Cieśliński (KTC);
Hajnówka Forest Division, forest section No. 389d, Fraxinus sp., 1983, leg. Z. Tobolewski &
S. Cieśliński (KTC); Hajnówka Forest Division, forest section No. 307d, Fraxinus excelsior,
1983, leg. Z. Tobolewski & S. Cieśliński (KTC); Starzyna Forest Division, forest section Nos
629b, 630a, Perebel River, Fraxinus excelsior, 1982, leg. S. Cieśliński & Z. Tobolewski (KTC).
GC-90 – Wysoczyzny Podlasko-Białoruskie (Podlachian-Belarusian Highlands), Wysoczyzna
Drohiczyńska (Drohiczyn Highland): Sokole Nature Reserve, Carpinus betulus, August 16,
1992, leg. S. Cieśliński (KTC). GD-20 – Nizina Południowopodlaska (South Podlachian
Lowland), Równina Łukowska (Łuków Plain): Chmielinne Nature Reserve, Alnus glutinosa,
May 18, 1962, leg. B. Ordyczyńska (LBL); Leśna near Biała Podlaska town, Alnus glutinosa,
May 18, 1962, leg. B. Ordyczyńska (LBL). GD-90 − Polesie Zachodnie (Western Polesie),
Zaklęsłość Sosnowicka (Sosnowica Depression): Białka village near Parczew town, Quercus
sp., October 30, 1986, leg. Z. Szczepańska (LBL). GD-91 – Polesie Zachodnie (Western
Polesie), Zaklęsłość Sosnowicka (Sosnowica Depression): Sosnowica Forest District, Bohutyn
village, Betula sp., May 1973, leg. H. Siegieda (LBL). GE-91 – Roztocze, Central Roztocze:
Hrebenne village, Acer pseudoplatanus, March 30, 1957, leg. K. Glanc (KRAM-L 39487);
Lublin Province, Zamość County, Kosobudy Forest District, Czerkies Forest District, Fagus
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sylvatica, June 30, 1952, leg. Z. Tobolewski (POZ); Kosobudy near Zamość town, Słupy Forest
District, Alnus sp., August 25, 1964, leg. B. Warmińska (LBL); Alnus sp., August 26, 1964, leg.
B. Warmińska (LBL); Roztoczański National Park, SE of Guciów village, Nart Nature Reserve,
beech forest, Tilia cordata, September 17, 2015, leg. M. Kukwa 17525 (UGDA-L 22020).
GE-92 – Roztocze, Central Roztocze: Lublin Province, Zamość County, Kosobudy Forest
District, Wólka Forest District, Quercus sp., August 24, 1964, leg. B. Warmińska (LBL).
GF-01 – Kotlina Sandomierska (Sandomierz Basin), Równina Biłgorajska (Biłgorajska Plain):
Szum Nature Reserve near Józefów, Alnus sp., October 16, 1970, leg. B. Żabińska (LBL).
GF-12 – Roztocze, Central Roztocze: Susiec village, by Tanew River, Alnus sp., May 2, 1964,
leg. J. Krygler (LBL).
Additional material examined. AUSTRIA: Botanischer Tauschverein in Wien, 1833,
(KRAM-L 384); leg. F. Hazslinszky (KRAM-L 395); Nenlengbach, 1940, (KRAM-L 386);
ca. 6 km N of Kaprun, Carpinus betulus, April 1, 2005, leg. A. Wieczorek (herb. Wieczorek).
BELARUS: Gomel Region, Gomel District, Korenevka forest, 52°22' N, 31°02' E, Parnassius
mnemosyne microreserve, Quercus sp., 24 October. 2007, leg. A. Tsurykau 01443 (GSU);
Gomel District, city of Gomel, Sverdlova Str. near hospital, Alnus glutinosa, April 15, 2007,
leg. A. Tsurykau 01444 (GSU). CZECH REPUBLIC: Deutschbrod in Böhmen leg. Jul. Svacha
(KRAM-L 387); Rychlebskė Horý, N of Lipová láznĕ, Jeskynĕ na Pomezí, limestone outcrops
covered with beech forest with sycamore, Fraxinus excelsior, April 24, 2004, leg. M. Kukwa
3206 (UGDA-L 10430). DENMARK: Bornholm, Dondalen, forest, Carpinus betulus, July 5,
2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek); Ekkodalen, Carpinus betulus, July 5,
2014, Bornholm, Dondalen, forest, Carpinus betulus, July 5, 2014, leg. A. Wieczorek (herb.
Wieczorek). ESTONIA: Saaremaa Island, Saare Country, Torgu Community, Viieristi Nature
Reserve, Ulmus sp., September 20, 2008, leg. J. Motiejūnaite 8608 (BILAS). GERMANY:
Mecklenburg-Vorpommern, Rostock, Wustrow Peninsula, parkland of former manor,
Fraxinus excelsior, October 20, 2014, leg. U. Schiefelbein 3694 (herb. Schiefelbein); October
20, 2012, leg. U. Schiefelbein 3687 (herb. Schiefelbein); Landkreis Uecker-Randow, Straße
Ueckermünde-Heinrichshof, ca. 0.3 km E Kreuzung mit der Zarow, Quercus sp., May 3,
2002, leg. U. Schiefelbein 945 (herb. Schiefelbein); Ueckertal, Eschenwald, ca. 1 km NW of
Schmarsow, Fraxinus excelsior, October 14, 2001, leg. U. Schiefelbein 1002 (herb. Schiefelbein);
N Borken, Eichen-Ulmen-Hartholz-Auenwald, Fraxinus excelsior, leg. U. Schiefelbein 1242
(herb. Schiefelbein); an der Straße Jatznick-Rothemühl, ca. 1 km E of Rothemühl, Fagus
sylvatica, March 6, 2001, leg. U. Schiefelbein 623 (herb. Schiefelbein); Demnitzer Bruch,
Fraxinus excelsior, October 22, 2000, leg. U. Schiefelbein 610 (herb. Schiefelbein); Landkreis
Ostvorpommern, Insel Usedom, Wald zwischen der Ortslage Ahlbeck und der polnischen
Grenz, nördl. der Bundesstraße, Fagus sylvatica, December 19, 2003, leg. U. Schiefelbein 1928
(herb. Schiefelbein); Waldgebiet zwischen Seebad Bansin und Ückeritz, ca. 0.5 km westlich der
ehemaligen Selliner Bootstelle, Fraxinus excelsior, November 14, 2003, leg. U. Schiefelbein 1383
(herb. Schiefelbein). LITHUANIA: Nemunas loops Regional Park, Prienai District, Balbieriškis
forest, forest area No. 50, Fagus sylvatica, September 28, 2002, leg. P. Czarnota (GPN 3141);
Molėtu raj., Giedraičių g-ja, 59 kvartalas, 11 sklypas, Zdoniškių miškas, Fraxinus excelsior,
November 23, 2006, leg. D. Stončius 8280 (BILAS); Molėtu raj., Sadauskų miškas, pietinis
Grabuosto ežero šlaitas, Acer platanoides, November 6, 2005, leg.D. Stončius 8459 (BILAS);
Akmenės raj., Naujosios Akmenės g-ja, 49 kvartalas, 20 sklypas, Alnus incana, September
2003, leg. E. Panovienė 8458 (BILAS); 438 kvartalas, 16 sklypas, ant lapuocio medzio kamieno,
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September 2003, leg. E. Panovienė 8457 (BILAS); Mažeikių raj., Ranavo g-ja, 272 kvartalas,
29 sklypas, ant lapuocio medzio kamieno, September 2003, leg. E. Panovienė 8456 (BILAS);
Tralus raj., Alnus glutinosa, June 27, 1997, leg. J. Motiejūnaite 2569 (BILAS); ANP, Utenos raj.,
Fraxinus excelsior, May 15, 2001, leg. J. Motiejūnaite 5592 (BILAS); Ukmergės raj., Želvos g-ja,
27 kvartalas, 9 sklypas, Laumėnų miškas, Fraxinus excelsior, March 11, 2009, leg.D. Stončius
9203 (BILAS); Kaisiadorių raj., Fraxinus excelsior, September 18, 1996, leg. J. Motiejūnaite 3571
(BILAS); Traku raj., Aukitadvario apylinkis, Tilia sp., June 27, 1997, leg. J. Motiejūnaite 2517
(BILAS); Biržų raj., Fraxinus excelsior, September 16, 1992, leg. J. Motiejūnaite 1560 (BILAS).
LATVIA: Kemeri National Park, Antinciems, Kanieru Lake, Fraxinus excelsior, September 28,
1996, leg. J. Motiejūnaite 5741 (BILAS); Juodkrantė, Kuršių Nerija, Acer platanoides, August
11, 1975, leg. T. Rudzinskaitė 2169 (WI); Biržų giria, 1963 kvartalas, Biržų raj, ant jauno
lapuocio medzio kamieno, May 13, 2006, leg. I. Prigodina 4048 (WI); Dükštų ąžuolynas,
9 kvartalas, 4 sklypas Neries RP, Vilniaus raj., Quercus robur, June 5, 2002, leg. I. Prigodina
1304 (WI).Talsi District, Šķēde Forest Research Station, Mežmāja, Tilia cordata, September
23, 2014, leg. M. Kukwa LV12955 (UGDA-L 19855). NORWAY: Stryn, forest, Abies alba,
August 28, 1995, leg. A. Wieczorek (herb. Wieczorek). RUSSIA: Kalujskie zaseki Reserve,
Kaluga Province, Ulyanovsky District, 2 km NW of Nogaja village, S plot of reserve, forest,
Acer sp., June 22, 2004, leg. A. V. Gudovicheva 862 (KPABGI); 4.5 km NW of Nogaja village,
S plot of reserve, Acer sp., June 22, 2004, leg. A. V. udovicheva 865 (KPABGI); Tula Province,
Shchyokinsky District, museum Yasnaya Polyana, section Podkapustnik, Betula sp., August 1,
2001, leg. A. V. Gudovicheva 1243 (KPABGI); Orlovskoye region, T. 216, Znamensky Distr.,
Orlovskoye Polesie National Park, Krasnikovskoye Forestry, 33 quar., broad-leaved forest,
bark of Fraxinus, N 53°29.142, E 35°33.750, 198 m alt., June 12, 2012, leg. E. E. Muchnik
9636 (OHHI); Orel region, T. 288, Shablykinsky Distr., near v. Belousovka, Shablykinsky
forest, mied forest, bark of Populus tremula, 53°53.177' N, 35°17.268' E, 226 m alt., July 25,
2013, leg. E. E. Muchnik 10601 (OHHI); Orel region, T. 287, Shablykinsky Distr., near v.
Lesnichestvo, Shablykinsky forest, broad-leaved forest, bark of Sorbus aucuparia, 52°54.199'
N, 35°10.892' E, 166 m alt., July 24, 2013, leg. E. E. Muchnik 10600 (OHHI); Bryansk region,
T. 378, Suzemsky Distr., Bryansky Les State Nature Reserve, 103 quae., broad-leaved forest,
bark od Acer, 52°27.286' N, 33°51.155' E, 153 m alt., August 10, 2015, leg. E. E. Muchnik (herb.
Muchnik). UKRAINE: Lysa Hora, 2 km E of Vilshanytsya, Lviv region, Zolochir District,
49°48' N, 24°43' E, Fraxinus sp., June 5, 2000, leg. U. Bielczyk & J. Kiszka (KRAM-L 45153);
Khmelnitskyi region, Kamianets Podilskyi District, National Park Podilskyi Tovtry, Privorita
near Makiv, Hora Karmeluka forest, 223 m, 48°47'27" N, 26°38'41" E, Carpinus betulus, June
25, 2003, leg. A. Kowalewska (KRAM-L 49103, 49065); Salix sp., June 25, 2006, leg. M. Kukwa
(KRAM-L 48776); An Waldbäumer 1876, leg. W. Krieger (WA).
Exsiccata seen. Erbar. Crittogam Ital. Exs. No. 430, as Opegrapha herpetica (KRAM); Wartmann & Schenk, Schweiz. Krypt. Exs. No. 675, as Opegrapha herpetica for. fuscata (KRAM);
Lichen. Helvec Exs. Schaer. et Hepp. No. 550 (WA).
Zwackhia sorediifera (P. James) Ertz
Specimens examined. AB-23 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam and Wolin
islands: Wolin Island, Wolin National Park, Fraxinus sp. September 22, 2014, leg. A. Wieczorek
(herb. Wieczorek). AB-94 – Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe
© The Author(s) 2018
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(Beech Hills): Puszcza Bukowa (Beech Forest), Szczecin Landscape Park, Carpinus betulus,
August 21, 2014, leg. A. Wieczorek (herb. Wieczorek).
Additional material examined. ESTONIA: Saaremaa, Torgu Commune, Viieristi Nature
Reserve, forest section No. 500, black alder forest with spruces and pines, Alnus glutinosa,
September 20, 2008, leg. M. Kukwa 7259 (UGDA-L 14809). LATVIA: Talsi District, between
Šķēde and Mežmāja, mied humid forest, Quercus robur, September 24, 2014, leg. M. Kukwa
LV12974 (UGDA-L 19874, KRAM-L 67700); Abies alba, September 24, 2014, leg. M. Kukwa
LV12979 (UGDA-L 19879).
Zwackhia viridis (Ach.) Poetsch & Schied.
Specimens examined. AB-15 – Pobrzeże Szczecińskie (Szczecin Coastland), Wybrzeże
Trzebiatowskie (Trzebiatowskie Coast): Łukęcin village, Quercus sp., June 13, 1986, leg.
W. Fałtynowicz (UGDA-L 2793). AB-23 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam
and Wolin islands: Wolin Island, Wolin National Park, forest section Nos. 7, 14, 18, 52, 58,
60, 70, 83, 92, 95, 107, Fagus sylvatica, Quercus sp., 1996, leg. A. Wieczorek (herb. Wieczorek).
AB-24 – Pobrzeże Szczecińskie (Szczecin Coastland), Uznam and Wolin islands: Wolin Island,
Kołczewo village, Fraxinus excelsior, 1996, leg. A. Wieczorek (herb. Wieczorek). AB-93 –
Pobrzeże Szczecińskie (Szczecin Coastland), Wzgórza Bukowe (Beech Hills): Puszcza Bukowa
(Beech Forest), Szczecin Landscape Park, forest section Nos 246, 284, Fagus sylvatica, July
15, 1998, leg. A. Wieczorek (herb. Wieczorek). AB-94 – Pobrzeże Szczecińskie (Szczecin
Coastland), Wzgórza Bukowe (Beech Hills): Puszcza Bukowa (Beech Forest), Szczecin
Landscape Park, Źródliskowa Buczyna Nature Reserve, forest section Nos. 232, 227, Fagus
sylvatica, July 12, 1997, leg. A. Wieczorek (herb. Wieczorek); Fagus sylvatica, forest section
Nos. 240, 248, June 13, 1999, leg. A. Wieczorek (herb. Wieczorek). BA-97 – Pobrzeże
Koszalińskie (Koszalin Coastland), Równina Słupska (Słupsk Plain): Drzeńsko Młyn village
Fagus sylvatica, August 28, 1987, leg. W. Fałtynowicz & J. Miądlikowska (UGDA-L 3889).
BA-98 – Pobrzeże Koszalińskie (Koszalin Coastland), Równina Słupska (Słupsk Plain):
Janiewickie Bagno Nature Reserve near Sławno town, Fagus sylvatica, August 26, 1987, leg.
W. Fałtynowicz & J. Miądlikowska (UGDA-L 3496). BC-22 – Pojezierze Południowopomorskie
(South Pomeranian Lake District), Równina Drawska (Drawno Plain): Drawieński Park
Narodowy (Drawa National Park), Ostrowiec Lake, Tilia sp., August 18, 1989, leg. L. Lipnicki
& J. Kiszka (herb. Lipnicki). BF-47 – Eastern Sudetes, Masyw Śnieżnika (Śnieżnik Massif):
valley of Kalenica Stream, leg. K. Szczepańska & M. Kossowska (herb. Szczepańska); valley
of Bogoryja Stream, Acer sp., August 27, 2003, leg. K. Szczepańska (herb. Szczepańska).
CA-61 – Pobrzeże Koszalińskie (Koszalin Coastland), Wysoczyzna Damnicka (Damnica
Highland): Damnice village, Carpinus betulus, November 20, 1987, leg. W. Fałtynowicz &
J. Miądlikowska (UGDA-L 3938). CA-66 – Pojezierze Wschodniopomorskie (East Pomeranian
Lake District), Pojezierze Kaszubskie (Kashubian Lake District): Porzecze hamlet, between
Lębork and Wejherowo towns, Fagus sylvatica, July 23, 1985, leg. W. Fałtynowicz (KRAML 21767, 21882). CA-68 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District),
Pojezierze Kaszubskie (Kashubian Lake District): between Rumia town and Zagórze, Quercus
sp., August 23, 1987, leg. W. Fałtynowicz (UGDA-L 3475). CA-98 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District):
Jar Rzeki Raduni Nature Reserve, Carpinus betulus, September 19, 1983, leg. W. Fałtynowicz
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(UGDA-L 1760); November 1, 1973, leg. W. Fałtynowicz (UGDA-L 1771); Fraxinus excelsior,
December 5, 1976, leg. W. Fałtynowicz (UGDA-L 1860); Fagus sylvatica, September 19, 1983,
leg. W. Fałtynowicz (UGDA-L 3015); Acer sp., April 14, 1973, leg. W. Fałtynowicz (UGDAL 3252). CA-99 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze
Kaszubskie (Kashubian Lake District): ca. 1.5 km of Kolbudy village, Acer sp., April 14, 1973,
leg. W. Fałtynowicz (UGDA-L 1748). CB-04 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Bory Tucholskie (Tuchola Forest): SW edge of Mausz Lake, Fagus
sylvatica, April 3, 1986, leg. W. Fałtynowicz (UGDA-L 3356). CB-56 – Pojezierze
Południowopomorskie (South Pomeranian Lake District), Bory Tucholskie (Tuchola Forest):
Ustronie and Twarożnica villages, Carpinus betulus, September 18, 1974, leg. L. Lipnicki
(herb. Lipnicki). CB-85 – Pojezierze Południowopomorskie (South Pomeranian Lake District),
Dolina Brdy (Brda River Valley): Piekło Nature Reserve by Brda River, Carpinus betulus,
March 30, 1985, leg. L. Lipnicki (TRN). CC-80 – Pojezierze Wielkopolskie (Wielkopolska
Lake District), Pojezierze Gnieźnieńskie (Gniezno Lake District): Zielonka Forest District
near Poznań City, forest section No. 4, 8, 73, Tilia cordata, June 17, 1961, leg. K. Glanc
(KRAM-L 39382); forest section No. 73, Acer platanoides, June 17, 1961, leg. K. Glanc (KRAML 39383); forest section No. 73, Acer sp., September 15, 1961, leg. K. Glanc (KRAM-L 39391);
forest section No. 8r, Tilia sp., October 29, 1960, leg. K. Glanc (KRAM-L 39393); forest section
No. 73, Tilia cordata, September 15, 1961, leg. K. Glanc (KRAM-L 39403); forest section No.
4j, Acer pseudoplatanus, October 28, 1960, leg. K. Glanc (KRAM-L 39404). CE-00 – Obniżenie
Milicko-Głogowskie (Milicz-Głogów Depression), Kotlina Milicka (Milicz Basin): Milicz
County, Kubryk Forest District, Carpinus betulus, June 27, 1969, leg. K. Glanc (KRAML 39389). DA-70 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pobrzeże
Kaszubskie (Kashubian Coastland), Sopot town, forest near Reja Str., Fagus sylvatica, March
8, 1959, leg. W. Fałtynowicz (UGDA-L 3450). DA-85 – Pobrzeże Gdańskie (Gdańsk Coastland),
Mierzeja Wiślana (Vistula Spit): Przebrno village, forest section No. 173, Carpinus betulus,
August 6, 1982, leg. E. Budzbon & W. Fałtynowicz (UGDA-L 2196); Fagus sylvatica, August
15, 1982, leg. E. Budzbon & Z. Tobolewski (UGDA-L 2197). DB-36 – Pojezierze Wschodniopomorskie (East Pomeranian Lake District), Pojezierze Kaszubskie (Kashubian Lake District),
Porzecze village, forest section No. 261, Acer sp., July 15, 1985, leg. W. Fałtynowicz (UGDAL 2868); Porzecze village, Łeba River, Fagus sylvatica, July 23, 1985, leg. W. Fałtynowicz
(UGDA-L 2888). DB-42 – Pojezierze Południowobałtyckie (South Baltic Lakeland), Pojezierze
Iławskie (Iława Lake District): Kwidyn town, forest section No. 190, Dolina Postolińskiej
Strugi (Postolińska Struga Valley), Tilia cordata, April 21, 2003, leg. M. Kukwa 1787 (UGDAL 13466). DE-68 – Wyżyna Przedborska (Przedborska Upland), Niecka Włoszczowska
(Włoszczowa Basin): Dębowiec Forest District, forest section No. 165, Ulmus glabra, associated
with Chrysothrix candelaris and O. vermicellifera, September 21, 1971, leg. K. Czyżewska
(LOD-L 1023). DG-02 – West Beskids, Beskid Śląski Mountains: Brennica Stream, Abies alba,
August 20, 1965, leg. J. Kiszka (KRAP). DG-03 – West Beskids, Beskid Śląski Mountains:
Skrzyczne village, Fagus sylvatica, September 4, 1964, leg. J. Kiszka (KRAP). DG-06 – West
Beskids, Beskid Makowski Mountains: Pasmo Jałowieckie (Jałowieckie Range), Stryszawa,
Siwcówka hamlet, Abies alba, August 16, 1965, leg. J. Nowak (KRAM-L 15766). DG-12 – West
Beskids, Beskid Śląski Mountains: Barania Góra (Ram Mountain), valley of Biała Wisełka
Stream, Fagus sylvatica, October 2, 1975, leg. U. Bielczyk (KRAM-L 43944); alp below Przysłop
Mountain, Fagus sylvatica, August 2, 1962, leg. J. Kiszka (KRAP); Skrzyczne Mount, S-slope,
Fagus sylvatica, September 4, 1964, leg. J. Kiszka (KRAM-L 19354). DG-13– West Beskids,
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Beskid Śląski Mountains: Malinowska Skała Mount, Acer pseudoplatanus, September 6, 1962,
leg. J. Kiszka (KRAP). DG-25 – West Beskids, Beskid Żywiecki Mountains: Pilsko Massif,
valley of Straceniec Stream, Acer pseudoplatanus, September 5, 1964, leg. J. Nowak (KRAML 14816, 14817). DG-26 – West Beskids, Beskid Żywiecki Mountains, Babia Góra National
Park, s.d., leg. U. Bielczyk (POZ). DG-38 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale
Depression), Kotlina Orawsko-Nowotarska (Orava-Nowy Targ Basin): Piekielnik Stream,
Fagus sylvatica, May 30, 1970, leg. J. Kiszka (KRAP 10817); Abies alba, June 20, 1970, leg.
J. Kiszka (KRAP 10838). DG-49 – Obniżenie Orawsko-Podhalańskie (Orava-Podhale Depression), Rów Podtatrzański (Podtatrzański Trench): Gubałówka, Picea abies, August 18, 1964,
leg. J. Kiszka (KRAP); Bystrzyk Valley, Picea abies, July 24, 1963, leg. J. Kiszka (KRAP).
EA-82 – Nizina Staropruska (Old Prussian Lowland), Wzniesienia Górowskie (Górowo Hills),
ca. 1 km N of Garbniki village, Tilia sp., May 8, 1989, leg. S. Cieśliński, (KTC). EB-22 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District):
ca. 1.5 km NE of Smolajny village, Tilia sp., August 23, 1993, leg. S. Cieśliński (KTC); Carpinus
betulus, August 23, 1993, leg. S. Cieśliński (KTC); ca. 3 km NNE of Smolajny village, Acer
platanoides, August 23, 1993, leg. S. Cieśliński (KTC); ca. 4 km NNE of Smolajny village,
Fraxinus excelsior, August 23, 1993, leg. S. Cieśliński (KTC). EB-26 – Pojezierze Mazurskie
(Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): ca. 2 km E of
Otry village, Fagus sylvatica, May 8, 1990, leg. S. Cieśliński (KTC). EB-27 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District): W edge
of Legińskie Lake, Carpinus betulus, May 3, 1989, leg. S. Cieśliński (KTC). EB-31 – Pojezierze
Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 2.5 km
SW of Różynka village, Fraxinus excelsior, September 14, 1989, leg. S. Cieśliński (KTC).
EB-32 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn
Lake District): Las Warmiński Nature Reserve by Łyna River, forest section No. 139, Fraxinus
excelsior, July 19, 1988, leg. S. Cieśliński (KTC); Carpinus betulus, May 28, 1994, leg. S. Cieśliński
(KTC); Fraxinus excelsior, May 28, 1994, leg. S. Cieśliński (KTC); ca. 2 km SW of railway
station in Cerkiewniki village, Fagus sylvatica, September 14, 1989, leg. S. Cieśliński (KTC);
Acer sp., September 14, 1989, leg. S. Cieśliński (KTC). EB-36 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Mrągowskie (Mrągowo Lake District), ca. 0.5 km NE of Dębowo
Nature Reserve, Fagus sylvatica, August 28, 1993, leg. S. Cieśliński (KTC). EB-42 – Pojezierze
Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): Olsztyn
City, city forest, Acer sp., October 21, 2000, leg. D. Kubiak (OLTC); Carpinus betulus, October
21, 2000, leg. D. Kubiak (OLTC). EB-59 – Pojezierze Mazurskie (Masurian Lake District),
Pojezierze Olsztyńskie (Olsztyn Lake District): Strzałowo Nature Reserve, Tilia cordata,
September 1, 1987, leg. S. Cieśliński (KTC); Puszcza Piska (Pisz Forest), Krutynia Nature
Reserve, Acer sp., August 31, 1981, leg. S. Cieśliński (KTC); Carpinus betulus, August 31,
1987, leg. S. Cieśliński (KTC). EB-60 – Pojezierze Mazurskie (Masurian Lake District), Pojezierze Olsztyńskie (Olsztyn Lake District): ca. 2 km E of Stare Jabłonki railway station, Tilia
cordata, September 20, 1994, leg. S. Cieśliński (KTC). ED-09– Nizina Środkowomazowiecka
(Central Mazovian Lowland), Kotlina Warszawska (Warsaw Basin), Równina Wołomińska
(Wołomin Plain): Dębina Nature Reserve, Carpinus betulus, September 1955, leg. J. Zielińska
(WA). ED-99 – Nizina Środkowomazowiecka (Central Mazovian Lowland), Równina
Kozienicka (Kozienice Plain): Puszcza Kozienicka (Kozienice Forest), Zagożdżon Nature
Reserve, Carpinus betulus, July 25, 1968, leg. S. Cieśliński (KTC). EE-00 – Wzniesienia
Południowomazowieckie (South Mazovian Hills), Równina Piotrkowska (Piotrków Plain):
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Puszcza Pilicka (Pilica Forest), Spała Nature Reserve, forest section No. 286, Carpinus betulus,
in oak-linden-hornbeam forest, April 28, 1969, leg. K. Czyżewska (LOD-L 525). EE-55 –
Wyżyna Kielecka (Kielce Upland), Płaskowyż Suchedniowski (Suchedniów Plateau): Suchedniów
Forest District, Dalejów Nature Reserve, Acer pseudoplatanus, July 1989, leg. Pańczyk &
K. Toborowicz (KTC). EE-64 – Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
Gruchawka near Kielce City, Sufraganiec Nature Reserve, Abies alba, October 22, 1972, leg.
K. Toborowicz (KTC). EE-86– Wyżyna Kielecka (Kielce Upland), Świętokrzyskie Mountains:
Stołowa Góra (Stołowa Mount), Carpinus betulus, July 12, 1964, leg. S. Cieśliński (KTC).
EG-11 – West Beskids, Gorce Mountains: Kudłoń Mount, near Adamówka village, Fagus
sylvatica, August 24, 1968, leg. K. Glanc (KRAM-L 39400). EG-20 – West Beskids, Gorce
Mountains: under Obidowiec Mount, Ścisłe Brzyzki Forest District, in Olszowy Potok (Olszowy
Stream) catchment area, Fagus sylvatica, November 9, 1994, leg. P. Czarnota (GPN 679);
under Obidowiec glade, in valley of Olszowy Potok (Olszowy Stream), Fagus sylvatica, July
4, 1994, leg. P. Czarnota (GPN 659); near Paciepnica Stream, leg. P. Czarnota (GPN 989);
W slope of Suhora Mount, Abies alba, September 10, 1959, leg. K. Glanc (KRAM-L 39384);
Fagus sylvatica, September 10, 1959, leg. K. Glanc (KRAM-L 39399). EG-21 – West Beskids,
Gorce Mountains: Turbacz Mount S slope, Kliczka forest, Fagus sylvatica, August 24, 1966,
leg. K. Glanc (KRAM-L 39405); Gorc Mount, W slope, alt. 900 m, Fagus sylvatica, September
08, 1959, leg. K. Glanc (KRAM-L 35859); Dolina Łopusznej (Łopuszna Valley), Fagus sylvatica,
August 23, 1965, leg. K. Glanc (KRAM-L 39401); under peak of W slope of Turbaczyk Mount,
Fagus sylvatica, September 6, 1964, leg. K. Glanc (KRAM-L 39392); Turbacz Nature Reserve,
on N slope of Turbacz Mount, Olszowy Potok (Olszowy Stream), Fagus sylvatica, June 27,
1966, leg. K. Glanc (KRAM-L 39388); in Łopuszanka Stream, leg. P. Czarnota (GPN 1018);
Turbacz Nature Reserve, in valley of Olszowy Potok (Olszowy Stream), Fagus sylvatica, June
23, 1994, leg. P. Czarnota (GPN 598); Hucisko Forest District, on W slope of Gorc Porębski
glade, Dolina Kamienicy (Kamienica River Valley), Acer pseudoplatanusJuly 11, 1994, leg.
P. Czarnota (GPN 772); Znaki Forest District, Dolina Kamienicy (Kamienica River Valley),
leg. P. Czarnota (GPN 1684); Turbacz Nature Reserve, near blue tourist trail, under Polana
Średnie (Średnie Glade), in valley of Turbacz Stream, Fagus sylvatica, June 18, 1996, leg.
P. Czarnota (GPN 1003, 1099); Szerokie Forest District, on S slope of Gorc glade in Dolina
Kamienicy (Kamienica River Valley), under Gonadrowe Polanki, Fagus sylvatica, July 14,
1994, leg. P. Czarnota (GPN 759); Turbacz Mount, valley of Turbacz Stream, under Polana
Średnie (Średnie Glade), Fagus sylvatica, May 23, 1996, leg. P. Czarnota (GPN 1296, GPN
1231); Turbacz Nature Reserve, under Polana Średnie (Średnie Glade), in valley of Olszowy
Potok (Olszowy Stream), Fagus sylvatica, June 23, 1994, leg. P. Czarnota (GPN 639); Turbacz
Mount, Łocha Forest District, in valley of Olszowy Potok (Olszowy Stream), leg. P. Czarnota
(GPN 1113, 1048); near blue tourist trail, near Kamienica Stream, leg. P. Czarnota (GPN 965,
968, 1391); near Wspólny Potok (Wspólny Stream), in Dolina Kamienicy (Kamienica River
Valley), Fagus sylvatica, June 9, 1997, leg. P. Czarnota (GPN 1647); near Roztoka Stream, leg.
P. Czarnota (GPN 812, 814); Turbacz Nature Reserve, in Dolina Roztoki (Roztoka Stream
Valley), leg. P. Czarnota (GPN 1557, 1014); at the hiking trail on Turbacz Mount, Fagus sylvatica, September 13, 1959, leg. K. Glanc (KRAM-L 39378); Lepietnica Forest District, Fagus
sylvatica, September 12, 1964, leg. K. Glanc (KRAM-L 39390); S slope of Turbacz Mount,
Dolina Łopusznej valley, Fagus sylvatica, August 24, 1966, leg. K. Glanc (KRAM-L 39402).
EG-22 – West Beskids, Gorce Mountains: Lubań Mount, S slope, valley of Skalny Stream,
from Lubań Mount to Grywałd village, Fagus sylvatica, May 10, 1959, leg. K. Glanc
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(KRAM-L 39386, 39379); Lubań Mount, slope N, from Tokarnia village on Lubań Mount,
Fagus sylvatica, September 4, 1960, leg. K. Glanc (KRAM-L 39381); valley of Forędówka
Stream, Fagus sylvatica, April 2, 1959, leg. K. Glanc (KRAM-L 39398). EG-33 – Obniżenie
Orawsko-Podhalańskie (Orava-Podhale Depression), Pieniny Mountains: Sokola Perć Mount,
Fagus sylvatica, July 1, 1955, leg. Z. Tobolewski (POZ); Czertezik Mount, Abies alba Fagus
sylvatica, September 14, 1957, leg. K. Glanc (KRAM-L 39375). EG-34 – West Beskids, Beskid
Sądecki Mountains: Pasmo Radziejowej (Radziejowa Range), nature reserve above Kotelniczy
Potok (Kotelniczy Stream), Fagus sylvatica, September 9, 1976, leg. U. Bielczyk (KRAML 42068); Baniska Nature Reserve, Fagus sylvatica, September 25, 1967, leg. M. Olech (KRA);
July 23, 1990, leg. L. Śliwa (KRA); valley of Potok Kozłecki (Kozłecki Stream), Fagus sylvatica,
June 15, 1965, leg. M. Olech (KRA); valley of Mała Roztoka Stream, Fagus sylvatica, August
24, 1968, leg. M. Olech (KRA). EG-35 – West Beskids, Beskid Sądecki Mountains: valley of
Czercz Stream, Fagus sylvatica, August 4, 1990, leg. L. Śliwa (KRA). EG-36 – West Beskids,
Beskid Sądecki Mountains: Pasmo Jaworzyny (Jaworzyna Range), valley of Potok Łososiowski
(Łososiowski Stream), Fagus sylvatica, October 14, 1967, leg. M. Olech (KRA); Kopiec Mount,
S slope, Fagus sylvatica, September 9, 1967, leg. M. Olech (KRA); Jaworzyna Krynicka Mount,
Fagus sylvatica, July 12, 1995, leg. A. Wieczorek (herb. Wieczorek). EG-37 – West Beskids,
Beskid Sądecki Mountains: valley of Kryściów Stream, Abies alba, July 18, 1967, leg. M. Olech
(KRA). EG-46 – West Beskids, Beskid Sądecki Mountains: Muszyna town, Fagus sylvatica,
August 12, 1953, leg. J. Rydzak (LBL). FA-85 – Pojezierze Litewskie (Lithuanian Lake District),
Puszcza Romincka (Romincka Forest): forest section No. 91, Fraxinus sp. May 16, 1985, leg.
S. Cieśliński (KRAM-L 31842; KTC). FB-00 – Pojezierze Mazurskie (Masurian Lake District),
Kraina Wielkich Jezior Mazurskich (Great Masurian Lakes Land): Mokre village, forest section No. 281, Quercus sp., 3 September. 1989, leg. S. Cieśliński (KTC). FB-03 – Pojezierze
Mazurskie (Masurian Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza Borecka (Borki
Forest), nature reserve, Carpinus betulus, May 14, 1958, leg. K. Glanc (KRAM-L 39394, 39397);
Puszcza Borecka (Borki Forest), near Czerwony Dwór village, forest section No. 195, Fraxinus
sp., May 21, 1985, leg. S. Cieśliński (KTC); Puszcza Borecka (Borki Forest), forest section
Nos 10, 205/206, 211/212, Fraxinus sp., May 21, 1985, leg. S. Cieśliński (KTC); forest section
No. 10, Carpinus betulus, May 19, 1987, leg. Z. Tobolewski & S. Cieśliński (KTC). FB-13 –
Pojezierze Mazurskie (Masurian Lake District), Pojezierze Ełckie (Ełk Lakeland): Puszcza
Borecka (Borki Forest), Borki Nature Reserve, Tilia sp., May 19, 1987, leg. S. Cieśliński (KTC);
forest section No. 172/173, Acer sp., May 20, 1987, leg. S. Cieśliński (KTC). FB-58 – Nizina
Północnopodlaska (North Podlachian Lowland), Kotlina Biebrzańska (Biebrza Basin): Czerwone Bagno Nature Reserve, forest section No. 246, Tilia cordata, 24 September. 1986, leg.
S. Cieśliński (KTC). FD-45 – Nizina Południowopodlaska (South Podlachian Lowland),
Równina Łukowska (Łuków Plain): Jata Nature Reserve near Łuków town, Alnus glutinosa,
June 23, 1971, leg. M. Motyka-Zgłobicka (LBL). FD-68 – Polesie Zachodnie (Western Polesie),
Zaklęsłość Łomaska (Łomaska Depression): Omelno Nature Reserve, Quercus sp., May 15,
1973, leg. B. Ordyczyńska (UMCS). FE-25 – Wyżyna Lubelska (Lublin Upland), Płaskowyż
Nałęczowski (Nałęczów Plateau): Wojciechowski Las near Nałęczów town, 1975, leg. Z. Flisińska
(KTC). FE-55 – Wyżyna Lubelska (Lublin Upland), Wzniesienia Urzędowskie (Urzędów
Hills): Urzędów town, Carpinus betulus, 1975, leg. Z. Flisińska (UMCS). FE-90 – Wyżyna
Kielecka (Kielce Upland), Wyżyna Sandomierska (Sandomierz Upland): Żyznów near Klimontów village, forest section No. 72, Acer pseudoplatanus, 1984, leg. S. Cieśliński & Z. Toborowicz (KTC). FF-89 − Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze
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Dynowskie (Dynowskie Foothills): Holubla Stream, Fraxinus excelsior, September 27, 1983,
leg. J. Kiszka & J. Piórecki (KRAM-L 60065). FG-08 – Pogórze Środkowobeskidzkie (Central
Beskidian Foothills), Pogórze Przemyskie (Przemyśl Foothills): valley of Holubla Stream,
Fraxinus sp., 27 September. 1983, leg. J. Kiszka & J. Piórecki (KRAM-L 30502). FG-18 –
Pogórze Środkowobeskidzkie (Central Beskidian Foothills), Pogórze Przemyskie (Przemyśl
Foothills): crest of Chwaniów Mount, Acer pseudoplatanus, June 3, 1987, leg. J. Piórecki
(BDAP). FG-19 – Beskidy Lesiste Mountains, Góry Sanocko-Turczańskie (Sanok-Turka
Mountains): Jamna Gora village, forest No. 2292, Acer pseudoplatanus, July 27, 1984, Leg.
J. Kiszka & J. Piórecki (KRAM-L 60068). FG-59 – Beskidy Lesiste Mountains, West Bieszczady
Mountains: Nasiczne village, Fagus sylvatica, July 30, 1959, leg. K. Glanc (KRAM-L 39396);
Pszczeliny village, Abies alba, July 29, 1959, leg. K. Glanc (KRAM-L 39387). FG-68 – Beskidy
Lesiste Mountains, West Bieszczady Mountains: Puszcza Bukowa (Beech Forest), by Wielki
Lutowy Stream, Fagus sylvatica, August 18, 1958, leg. K. Glanc (KRAM-L 39377); Fagus
sylvatica, August 23, 1957, leg. Z. Tobolewski, (POZ); Abies alba, August 28, 1957, leg. K. Glanc
(KRAM-L 39374). GB-10 – Pojezierze Litewskie (Lithuanian Lake District), Równina Augustowska (Augustów Plain): Wigry National Park, ca. 2 km of Mikołajewo village, Pinus sp.,
September. 1984, leg. K. Toborowicz (UGDA-L 4068). GB-30 – Pojezierze Wschodniosuwalskie
(East Suwałki Lake District), Równina Augustowska (Augustów Plain): Puszcza Augustowska
(Augustów Primeval Forest), forest section Nos. 51, 24, Fraxinus sp., September 13, 1989,
leg. S. Cieśliński & Z. Tobolewski (KTC). GB-31 – Pojezierze Litewskie (Lithuanian Lake
District), Równina Augustowska (Augustów Plain): Puszcza Augustowska (Augustów Primeval
Forest), Starożyn Nature Reserve, Carpinus betulus, July 15, 1965, leg. J. Zielińska (WA).
GB-32 – Pojezierze Wschodniosuwalskie, Równina Augustowska (Augustów Plain): ca. 2 km
S of Gruszki village, forest section No. 94, Fraxinus excelsior, 13 September. 1989, leg.
S. Cieśliński & Z. Tobolewski (KTC). GB-40 – Nizina Północnopodlaska (North Podlachian
Lowland), Kotlina Biebrzańska (Biebrza Basin): Puszcza Augustowska (Augustów Primeval
Forest), forest section No. 51, Łozy Forest District, Fraxinus excelsior, September 13, 1989,
leg. S. Cieśliński & Z. Tobolewski (KTC). GB-91 – Nizina Północnopodlaska (North Podlachian Lowland), Wysoczyzna Białostocka (Białystok Highland), Puszcza Knyszyńska (Knyszyn
Forest), ca. 1 km NWW of Machnacz village, forest section No. 130, Carpinus betulus, September 1997, leg. K. Glanc, Z. Tobolewski & M. Murati (KTC); Quercus sp., September 8,
1987, leg. Z. Tobolewski & K. Glanc (KTC); Brzozowy Gród Nature Reserve, Carpinus betulus,
1981, leg. Małyszko (LBL). GB-92 – Nizina Północnopodlaska (North Podlachian Lowland),
Wysoczyzna Białostocka (Białystok Highland): Puszcza Knyszyńska (Knyszyn Forest), Budzisk
Nature Reserve, Fraxinus excelsior, August 5, 1994, leg. S. Cieśliński (KTC); Carpinus betulus,
August 5, 1994, leg. S. Cieśliński (KTC); Carpinus betulus, 1980, leg. J. Bystrek (LBL); Tilia
cordata, May 13, 1993, leg. K. Kolanko (University of Bialystok, Institute of Biology); Carpinus
betulus, Acer sp., October 21, 1993, leg. K. Kolanko (University of Bialystok, Institute of Biology); Acer sp., (LBL); Carpinus betulus, leg. J. Rydzak (LBL); forest area 1, Fraxinus excelsior,
June 11, 1999, leg. S. Cieśliński (BILAS 5918). GC-53 – Nizina Północnopodlaska (North
Podlachian Lowland), Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża
Primeval Forest), Hajnówka Forest District, forest section No. 474, 2003, leg. S. Cieśliński
(KTC); forest section No. 360b, Fraxinus excelsior, 1983, leg. S. Cieśliński (KTC): forest section
No. 247b, Carpinus betulus, 1983, leg. S. Cieśliński (KTC); forest section No. 359d, Fraxinus
excelsior, 1983, leg. S. Cieśliński (KTC); forest section No. 663d, Carpinus betulus, 1983, leg.
S. Cieśliński (KTC); Zwierzyniec Forest District, forest section No. 392b (plot 133), Fraxinus
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excelsior, 1982, leg. S. Cieśliński (KTC); Fraxinus excelsior, August 22, 1965, leg. J. Rydzak
(LBL). GC-55 – Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska
(Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest), Białowieża National Park,
Fraxinus excelsior, September 14, 1953, August 7, 1954, leg. J. Rydzak (UMCS); Carpinus
betulus, August 5, 1954, leg. J. Rydzak (UMCS); Acer pseudoplatanus, July 31, 1963, leg.
J. Rydzak (UMCS); forest section No. 314, Betula sp., September 15, 1953, leg. J. Rydzak
(UMCS); forest section No. 399, Carpinus betulus, September 8, 1953, leg. J. Rydzak (UMCS);
forest section No. 314, near Orłówka Stream, Fraxinus excelsior, October 28, 2003, leg.
K. Czyżewska (LOD-L 11834); forest section No. 256, Tilia cordata, June 16, 1988, leg.
S. Cieśliński (LOD-L 777), leg. S. Cieśliński & K. Czyżewska (KTC); June 15, 1988, leg.
S. Cieśliński & K. Czyżewska (KTC); October 12, 1988, October 18, 1989, October 19, 1989,
leg. S. Cieśliński (KTC); Carpinus betulus, June 17, 1988, leg. S. Cieśliński (KTC); Carpinus
betulus, October 16, 1989, leg. S. Cieśliński (KTC); Fraxinus sp., October 15, 1988, September
16, 1989, October 17, 1989, October 18, 1989, leg. S. Cieśliński (KTC); Acer platanoides,
October 18, 1989, leg. S. Cieśliński (KTC); forest section No. 398a/d, leg. S. Cieśliński (KTC);
Carpinus betulus, May 2014, leg. M. Kukwa 12679, 13805, 12819, 12759, A. Łubek (UGDAL 20295, 20382, 20350, 21481); August 2014, leg. M. Kukwa 13958, A. Łubek (UGDA-L 21579);
May 2015, leg. M. Kukwa 15672, A. Łubek (UGDA-L 22251); Fraxinus excelsior, May 2014,
leg. M. Kukwa 12690, A. Łubek (UGDA-L 20302); October 2014, leg. M. Kukwa 13061,
A. Łubek (UGDA-L 20571). GC-65 – Nizina Północnopodlaska (North Podlachian Lowland),
Równina Bielska (Bielsk Plain): Puszcza Białowieska (Białowieża Primeval Forest), Browsk
Forest District, Carpinus betulus, September 13, 1964, leg. J. Rydzak 224 (UMCS); Browsk
Forest District, (plot 162), Fraxinus excelsior, 1983, leg. S. Cieśliński (KTC); forest section
No. 101b (plot 217), Fraxinus sp., 1983, leg. S. Cieśliński (KTC); forest section No. 160d, leg.
S. Cieśliński (KTC); Białowieża Forest District, forest section No. 474, Carpinus betulus, 1981,
leg. S. Cieśliński & Z. Tobolewski (KTC). GC-66 – Nizina Północnopodlaska (North Podlachian Lowland), Równina Bielska (Bielsk Plain): Białowieża National Park, forest section No.
340a, 2002, leg. P. Czarnota (GPN 2955). GC-91 – Nizina Północnopodlaska (North Podlachian
Lowland), Wysoczyzna Drohiczyńska (Drohiczyn Highland): Nurzec village, Carpinus betulus,
May 7, 1993, leg. S. Cieśliński (KTC). GD-01 – Wysoczyzny Podlasko-Białoruskie (PodlachianBelarusian Highlands), Wysoczyzna Drohiczyńska (Drohiczyn Highland): forest between
Adamowo and Złotorka villages, Fraxinus sp., May 1, 1987, May 11, 1987, leg. S. Cieśliński
& Z. Tobolewski (KTC). GE-91 – Roztocze, Central Roztocze: near Kosobudy village, Fagus
sylvatica, 1962, leg. J. Bystrek (UMCS). GG-71 – Beskidy Lesiste Mountains, West Bieszczady
Mountains: Bieszczady National Park, Sianki village, forest section No. 78i, Acer sp., June 19,
2002, leg. P. Czarnota (GPN 2860).
Additional material examined. AUSTRIA: ca. 6 km N of Kaprun, Carpinus betulus, April 1,
2005, leg. A. Wieczorek (herb. Wieczorek). BELARUS: Gomel Region, Zhitkovichi District,
Pripyatsky National Park, Ozerany Forestry, 28 q., 52°01' N, 27°56' E, in ash forest, on Fraxinus
excelsior, June 23, 1971, leg. O. Shakhrai 01489 (GSU). CZECH REPUBLIC: Moravia, N of
Rychtážov village, Ne of Ošlouch Mount, Vápenný žleb valley, Carpinus betulus, April 21,
2007, leg. M. Kukwa 5546a (UGDA-L 15037). DENMARK: Bornholm, Dondalen, forest,
Carpinus betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek); Ekkodalen,
Carpinus betulus, July 5, 2014, leg. A. Wieczorek & A. Łysko (herb. Wieczorek); South Jutland, Hoptrup, near Haderslev, Carpinus, September 11, 1969, leg. M. S. Christiansen (herb.
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Seaward). GERMANY: Oldenburg, An Fraxinus bei Helle, No. 711, leg. H. Sandstede (WA);
Mecklenburg-Vorpommern, Vorpommern-Rügen, Vordarß, Ahrenshooper Holz, northern
part, alt. 2 m, September 8, 2013, leg. U. Schiefelbein 3580 (herb. Schiefelbein); Landkreis
Müritz, Wäldchen W Troja, ca. 0.2 km W of Troja, Acer platanoides, November 1, 2002, leg.
U. Schiefelbein 1186 (herb. Schiefelbein); Landkreis Ostvorpommern, ca. 0.3 km W of Mümmelkensee, Quercus sp., December 3, 2002, leg. U. Schiefelbein 1317 (herb. Schiefelbein); Kreis
Uecker-Randow, ca. 1 km N of Rödershorst, Eichenwald auf entwässertem Moor, Quercus
petraea, May 20, 1999, leg. U. Schiefelbein 293 (herb. Schiefelbein); An glattrindigen Eschen
in der Neehagen unweit Helle bei Zwischenahn in Oldenburg, 1890, leg. H. Sandstede (herb.
Seaward). LITHUANIA: Kėdainių raj., Gelnų miskas, Labūnavos g-ja, 112 kvartalas, 10 sklypas,
Quercus robur, June 10, 2008, leg. D. Stončius 9525 (BILAS); Nemuno kilpų regioninis parkas,
Alytaus raj., Punios šilas, 17 kvartalas, 19 sklypas, Fraxinus excelsior, August 9, 2006, leg.
J. Motiejūnaite 7682 (BILAS); 42 kvartalas, 17 sklypas, Carpinus betulus, August 10, 2006,
leg. J. Motiejūnaite 7683 (BILAS); Radviliškio raj., Baisogalos g-ja, 83 kvartalas, Fraxinus
excelsior, September 30, 2013, leg. V. Stukonis 10512 (BILAS); Aukštelkų g-ja, 62 kvartalas,
2 sklypas, Fraxinus excelsior, September 21, 2013, leg. V. Stukonis 10447 (BILAS). NORWAY:
Stryn, forest, Abies alba, August 28, 1995, leg. A. Wieczorek (herb. Wieczorek). RUSSIA: Tula
Province, Shchyokinsky District, museum Yasnaya Polana, section Afonina Grove, Betula
sp., August 1, 2001, leg. A. V. Gudovicheva 1247 (KPABGI). SWEDEN: Ostergotland, Sund,
Pinus sp., August 27, 1987, leg. E. Kozioł (WRSL).
Exsiccata seen. Migula, Kryptogamae, Germaniae, Austriae et Helvetiae Exs. No. 1
(KRAM).
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