Uholka Primeval Forest in the Ukrainian Carpathians – A Keynote Area for Diversity of Forest Lichens in Europe

Abstract

Malíček, J., Palice, Z., Acton, A., Berger, F., Bouda, F., Sanderson, N. & Vondrák, J. 2018. Uholka primeval forest in the Ukrainian Carpathians — a keynote area for diversity of forest lichens in Europe. — Herzogia 31: 140–171.
One of the largest European primeval forests, Uholka-Shyrokyi Luh in the Ukrainian Carpathians, has received increased attention in recent years. In spring 2015 we explored the lichen biota in the southern part of the reserve. Species richness of epiphytic and epixylic lichens presented in this paper far exceeds all numbers achieved in other Central European old-growth forests. In total, 370 lichenized and lichen-allied fungi and 30 lichenicolous fungi were recorded. We focused on forest lichens on organic substrata, inorganic substrata were largely ignored. Species composition in the Uholka forest includes many rare taxa and typical old-growth forest species: e.g. Cetrelia spp., Gyalecta spp., Leptogium saturninum, Lobaria pulmonaria, Ricasolia amplissima, Sclerophora farinacea, S. pallida, Thelopsis flaveola and T. rubella. Opegrapha fumosa, Pyrenula chlorospila and P. dermatodes represent oceanic species that are very rare outside western Europe. Biatora longispora, Calicium montanum, Menegazzia subsimilis, Micarea perparvula, Ochrolechia trochophora, Pyxine sorediata, Ramonia luteola and Thelotrema suecicum are examples of phytogeographically remarkable or generally very rare lichens. Thirty lichenized and ten lichenicolous fungi are new to Ukraine, including Biatora bacidioides and Pertusaria macounii not previously reported from Europe.

Full text

140 Herzogia 31 (1) Teil 1, 2018: 140 –171
Uholka primeval forest in the Ukrainian Carpathians –
a keynote area for diversity of forest lichens in Europe
Malíek*Palice, Andy Acton, Franz BergerBouda, Neil
Sanderson & Jan Vondrák
Abstract: Malíek, J., Palice, Z., Acton, A., Berger, F., Bouda, F., Sanderson, N. & Vondrák, J. 2018. 

31: 140 –171.
   
         



Cetrelia spp., Gyalecta
spp., Leptogium saturninum, Lobaria pulmonaria, Ricasolia amplissima, Sclerophora farinacea, S. pallida, Thelopsis
flaveola and T. rubella. Opegrapha fumosa, Pyrenula chlorospila and P. dermatodes  
Biatora longispora, Calicium montanum, Menegazzia subsimilis, Micarea per-
parvula, Ochrolechia trochophora, Pyxine sorediata, Ramonia luteola and Thelotrema suecicum

Biatora bacidioides and Pertusaria macounii
Zusammenfassung: Malíek, J., Palice, Z., Acton, A., Berger, F., Bouda, F., Sanderson, N. & Vondrák, J.
2018. 
Herzogia 31: 140 –171.




        
    
CetreliaGyalecta Arten, Leptogium saturninum, Lobaria pulmona-
ria, Ricasolia amplissima, Sclerophora farinacea, S. pallida, Thelopsis flaveola T. rubella. Opegrapha fumosa,
Pyrenula chlorospilaP. dermatodes
Biatora longispora, Calicium montanum, Menegazzia subsimilis, Micarea perparvula, Ochrolechia trochophora,
Pyxine sorediata, Ramonia luteola Thelotrema suecicum  
Biatora
bacidioidesPertusaria macounii
Key words:   Biatora bacidioides      
Pertusaria macounii


Malíek   141
Introduction

 Mace et al. 2012, Gao   
Loo 

         



Vondrák   
Malíek & PaliceVondrák et al. 2016, Groner 2016).


    Dymytrova    


ymytrova et

Methods
Surveyed area








    




            
  

Commarmot
      

 




142 Herzogia 31 (1) Teil 1, 2018

Fagus sylvatica




Fagus sylvatica

Acer platanoides, A. pseudoplatanus, Carpinus betulus, Tilia platy-
phyllos and Ulmus glabra
   


Fig. 1.
Field work







Vondrák

Malíek   


      

 Lithothelium, Naetrocymbe, Ramonia, Thelocarpon
Chaenothecopsis, Microcalicium, Mycocalicium, Stenocybe
Selva
(Anisomeridium, Arthonia, Naetrocymbe
     

   


List of localities
1       
  Fagus sylvatica and Carpinus betulus  

2 

3 

4 Corylus avellana, Carpinus betulus 

5 

6        

7 
8  

Species identification and molecular barcoding
       

Orange



Johnson



Bacidia albogranulosa and Japewia sp.). Both of
Vondrák et al.
Vondrák et al. 2016).

144 Herzogia 31 (1) Teil 1, 2018
Fig. 2.Fagus sylvatica

Fig. 3.


Malíek   
Results

          

    
      
Agonimia flabelliformis, Alyxoria ochro-
cheila, Arthonia biatoricola, A. helvola, Bacidina etayana, Biatora bacidioides, B. longis-
pora, B. mendax, B. pontica, Calicium montanum, Chalara lobariae, Dactylospora homo-
clinella, Gyalecta croatica, Gyalideopsis helvetica, Lecanora exspersa, Lecidella subviridis,
Lithothelium hyalosporum, Menegazzia subsimilis, Merismatium heterophractum, Micarea
anterior, M. deminuta, M. perparvula, M. soralifera, Normandina acroglypta, Opegrapha
fumosa, O. thelotrematis, Pertusaria macounii, Phoma lobariae, Pseudosagedia byssophila,
Psoroglaena stigonemoides, Pyrenula chlorospila, Rhymbocarpus neglectus, Rinodina malan-
gica, Scoliciosporum schadeanum, Sphinctrina leucopoda, Taeniolella toruloides, Verrucaria
hegetschweileri, Vezdaea retigera, Xenonectriella septemseptata and Xylographa trunciseda.
Biatora radicicola, Lecanora stanislai, L. substerilis
Printzen et al. 2016,
Guzow-Krzemiska et al. 2017, Malíek et al. 2017).
Comments on rare, poorly known and phytogeographically remarkable species
Bacidia albogranulosa ined.
Lepraria in its morpho
    Acer 
 Fagus

Vondrák et al. 2016). Bacidia albogranulosa is rela
ted to B. rubella
Vouchers:
Biatora bacidioides
Biatora beckhausii


Printzen & Tønsberg
B. chrysantha or B. efflorescens

B. fallax, B. kodiakensis and B. printzenii (Printzen &
Tønsbergrintzen 
Biatora bacidioidesPicea orientalis forests
Printzen & Tønsberg
Fagus and Carpinus

Vouchers:
Biatora longispora 
B. helvola
Biatora longispora displays


146 Herzogia 31 (1) Teil 1, 2018

      
Printzen & Otte Gasparyan & Sipman 2016, Printzen
Fagus in river valleys.
Vouchers:
Japewia sp.
   Placynthiella dasaea  
Placynthielladasaea (Vondrák 

Japewia


Vouchers:
Micarea perparvula 
M. perparvula         
   

Coppins
this Micarea

Voucher:Additional record:
Pinus,

Ochrolechia trochophora
   
 
  
Kukwa
Tilia platyphyllos
O. pallescens.
Voucher:
Opegrapha fumosa

Opegrapha fumosa
  
(Pentecost & James
Fagus at a very


Voucher:
Pertusaria macounii
Pertusaria pertusaSedifolia
              
ibben
Pertusaria in Thailand (Jariangprasert
Buaruang et al. 2017). Pertusaria macounii
Carpinus
Fagus

Malíek   147
Sedifolia
P. macounii 
morphotype of P. pertusa.
Voucher:Additional record:
                 
 Fagus  

Pyrenula spp.
Pyrenula
P. laevigata, P. nitida and P. niditella  P.
chlorospila, P. coryli and P. dermatodesCarpinus betulus and Acer pseu-
doplatanus. Pyrenula coryli
Vzda & Liška 1999, Bielczyk et al. 2004, Wirth
Pyrenula dermatodes
Orangeksner P. chlorospi-
la imis
Orange
Pyxine sorediata
    
Nádvorník
1947, see the map in Masson
(Masson  
and 1940 (Lynge , Nádvorník 1947).
Voucher:
Lichen communities and keynote habitats for nature conservation

Cetrelia mona-
chorum, C. olivetorum, Heterodermia speciosa, Lobaria pulmonaria, Menegazzia terebrata,
Normandina pulchella, Sclerophora farinacea and Thelopsis rubella

  





Leptogium
saturninum, Nephroma parile, N. resupinatum, Peltigera collina, Ricasolia amplissima. These
   
         


in the Alps (e.g. HintereggerCaloplaca sorocarpa, Lecanora exspersa and
Rinodina malangica. Catillaria erysiboides, Frutidella furfuracea, Lecanora subintricata,
Micarea globulosella and Xylographa trunciseda.    
 

148 Herzogia 31 (1) Teil 1, 2018
Fig. 4. Coniocarpon cinnabarinumCarpinus betulus

Fig. 5. Gyalecta truncigena

Malíek   149
Fig. 6. Nephroma parile
Fig. 7. Sclerophora farinacea   

Acarospora
fuscata, Aspicilia caesiocinerea, Lecanora polytropa, Lecidella carpathica, Porpidia macro-
carpa, Protoparmeliopsis muralis


 Herzogia 31 (1) Teil 1, 2018

        

Fagus 

Hypotrachyna afrorevoluta, Parmotrema

as Arthonia, Graphis, Opegrapha and Pyrenula
Carpinus and Fagus Arthonia atra, A. helvola, A. ruana, Coniocarpon cin-
nabarinum, Inoderma byssaceum and Pyrenula nitidellaBiatora bacidioides,
B. longispora, B. vernalis, Coenogonium luteum, Lecanora cinereofusca, Megalaria laureri,
Menegazzia subsimilis, Pyrenula laevigata, Opegrapha fumosa, Pyxine sorediata and Thelotrema
suecicum
Micarea, among them
M. anterior, M. cinerea, M. deminuta, M. nigella and M. perparvula.

       
       Quercus petraea agg., Tilia
platyphyllos, Taxus baccata and Acer platanoides
  Caloplaca herbidella, Candelariella reflexa,
Gyalecta croatica, Melaspilea proximella, Ochrolechia arborea, O. trochophora, Pannaria
conoplea, Pertusaria flavida, Pseudosagedia byssophila and Verrucaria breussii

Discussion
Our lichen diversity survey vs. previous research in the locality
Dymytrova
  2

Lecanora strobilina, Ramonia luteola, Rinodina
capensis, Tetramelas chloroleucus, Usnea wasmuthii and Wadeana dendrographa

 
  Dymytrova       
the area (e.g. Buellia schaereri, Lecanora strobilina, Mycobilimbia carneoalbida, Pertusaria
pustulata, Rinodina conradii, Usnea ceratina, Wadeana dendrographa and Xanthomendoza
ulophyllodes Arthonia vinosa
A. didyma or A. spadicea), Calicium abietinum 
 C. glaucellum in the list), Haematomma ochroleucum  
Lecanora thysanophoraLecanora allophana
L. glabrata), L. impudens
Lecanora – L. exspersa or L. substerilis), Pyrrhospora quernea     
Lecanora expallens or Lecidella subviridis

reported in DymytrovaLecanora subrugosa Nyl. is regarded as a synonym of L.

Malíek   
argentata (Lumbsch & Feige 1996, Malíek Porina hibernica
P. pseu-
dohibernica (TretiachCandelaria con-
color, Mycobilimbia pilularis, Naetrocymbe rhyponta (as Arthopyrenia rhyponta), Peltigera
degenii, Reichlingia leopoldii, Rinodina pyrina, Tetramelas insignis (as Buellia insignis),
Thelocarpon laureri, Usnea dasopoga and U. wasmuthii.
Comparison of Uholka to other Central European old-growth forests

Dymytrova
    

   
Cieliski
& Tobolewski


Czyewska et al. 2001, SparriusKukwa et al. 2008, Łubek & Jaroszewicz

         

Groner
   

Table 1.        

Locality
(country)
Area
(ha) No. of epiphytic and
epixylic lichens All lichenized
and lichen-allied
fungi
References
 4729   this paper
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  189 202 Vondrák et al. (2016)



Vondrák  


 Herzogia 31 (1) Teil 1, 2018
Malíek, Palice &
Vondrák
Substrate preferences
  
  Hobi
Fagus sylvatica 
Acer pseudoplatanus, A. platanoides,
Carpinus betulus, Fraxinus excelsior and Ulmus glabra
Carpinus and Tilia
Fagus sylvaticaDymytrova
et al. 2014, Malíek & PaliceVondrákHofmeister et al. 2016).
Table 2.

Substrate preferences of epiphytic and epixylic species
Acer platanoides

Acer pseudoplatanus
107
Carpinus betulus

Fraxinus excelsior
82
Fagus sylvatica

Tilia sp.
116
Ulmus glabra


88
Acknowledgements
    
Pertusaria macouniiRinodina



References
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
Bilovitz, 
     136:
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23: 1–91.
Commarmot, B., Brändli, U.-B., Hamor, F. & Lavnyy, V.         

Cieliski, S. & Tobolewski, Z.         
                
1: 1–116.
Czyewska, K., Motiejnait, J. & Cieliski, S.
36
Dibben, M. J.   Pertusaria    
5: 1–162.

Malíek   
Dietrich, M.  101:
167–182.
Dymytrova, L., Nadyeina, O., Naumovych, A., Keller, C. & Scheidegger, C.
26
Dymytrova, L., Nadyeina, O., Hobi, M. L. & Scheidegger, C. 

23
Gao, T., Hedblom, M., Emilsson, T. & Nielse, A. B.
330
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281: 1– 68.
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22
Guzow-Krzemiska, B., Łubek, A., Malíek, J., Tønsberg, T., Oset, M. & Kukwa, M. 2017. Lecanora stanislai,
329: 201–211.
Hinteregger,RhododendronRh. ferrugineumRh. hirsutum) der Ostalpen
Biatora55
Hobi, M. L., Commarmot, B. & Bugmann,    
26
Hofmeister, J., Hošek, J., Malíek, J., Palice, Z., Syrovátková, L., Steinová, J. & ernajová,
Fagus sylvatica     
25
Jariangprasert,Pertusaria 
7
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36
Kondratyuk, S. Y. & Coppins, B. J. 
8: 149 –191.
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  
         
6
Kukwa,Ochrolechia
Kukwa, M., Schiefelbein, U., Czarnota, P., Halda, J., Kubiak, D., Palice, Z. & Naczk, A. 2008. Notes on some
41: 1–11.
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95
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
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27
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
Mace, G. M., Norris, K. & Fitter, A. H. 
27: 19 –26.
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49: 
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26
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56
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 Herzogia 31 (1) Teil 1, 2018
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
34: 619 – 626.
Vondrák, J., Malíek, J., Šoun, J. & Pouska, V. 
28: 104 –126.


Addresses of the authors


             




         






Malíek   
Appendix 1.DymytrovaAcer plata-
noides, AP – Acer pseudoplatanusCorylus avellanaCarpinus betulusFraxinus excelsiorFagus sylvatica, QU – QuercusSorbus aucuparia
Sambucus nigraTilia Ulmus glabra
Hafellner & Türk (2016) and Wirth
Dymytrova

species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Absconditella lignicola 11111  
Acarospora fuscata 1  
Acrocordia gemmata 11 11 11  
Agonimia allobata 11 11 1   
Agonimia borysthenica 
 1 1  ZP2
*Agonimia flabelliformis 1 1 log ZP
Agonimia opuntiella 1 log AA
Agonimia repleta 1 1 1 1   
Agonimia tristicula 1111 1 
(often on mosses)  
*Alyxoria ochrocheila 1 1  
Alyxoria varia 11111   
Amandinea punctata 1 1 1 1   
Anaptychia ciliaris 1 1 1  
Anisomeridium biforme 1 1 1   
Anisomeridium macrocarpum 1 1  
Anisomeridium polypori 1 1 1 1 


Arthonia atra 1  ZP2
Arthonia didyma 11111 1   
*Arthonia helvola 1 1  AA2, FB, FrB, ZP
Arthonia mediella 1 1 1  
Arthonia punctiformis 1  AA
Arthonia radiata 111111   

 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Arthonia ruana 1 1 1   
Arthonia spadicea 1 1 1 1  
Arthonia vinosa 
Arthothellium spectabile 1 1 1 1  
Arthrorhaphis grisea 1 Baeomyces
rufus)ZP
Aspicilia caesiocinerea 1 1  
Bacidia albogranulosa ined. 1 1  
Bacidia arceutina 1  FB
Bacidia circumspecta 1111 1 1 
Bacidia fraxinea 1  
Bacidia incompta 1 1 1  
Bacidia laurocerasi 1  ZP
Bacidia pycnidiata  1 log FrB
Bacidia rosella 1 1 1   
Bacidia rubella 1111 1  ZP 
Bacidia subincompta 11111  
Bacidia vermifera 1 1 1  
Bacidina delicata 1  
*Bacidina etayana 1  FrB
Bacidina mendax  1 1  
Bacidina phacodes 1 1 1 1   
Bacidina sulphurella 11111  
Baeomyces rufus 11  
Biatora albohyalina 1  ZP
*Biatora bacidioides  1 1 1  
Biatora beckhausii 1 1  ZP2
Biatora chrysantha 1 1 1  
Biatora efflorescens 1 1 1 1   
Biatora globulosa 1 1 1 1  
*Biatora longispora  1 1 1  

Malíek   
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
*Biatora mendax 1 1 1 1  
Biatora ocelliformis 1 1 1 1  
*Biatora pontica 1 1 1 1 1  
Biatora radicicola  1 1  
Biatora vernalis 1 1 1 1  
ZP7 
Biatoridium monasteriense 111 1 1   
Bilimbia sabuletorum 1 1 1   
Bryoria fuscescens 1 1   
Bryostigma apateticum 11111  
Buellia disciformis 111111 1  
Buellia erubescens 1  
Buellia griseovirens 11111 1 

 
Buellia schaereri 
Calicium abietinum 
Calicium glaucellum 1 1 QU, snag 
*Calicium montanum 1 snag (QU) 
Calicium salicinum 1 1 1  
Calicium trabinellum 1 snag (QU) 
Caloplaca cerina 1 1  
Caloplaca cerinelloides 11  AA
Caloplaca herbidella 1 1  
Caloplaca lucifuga 1  
Caloplaca monacensis 1 1  
Caloplaca obscurella 11  AA, ZP2
Caloplaca sorocarpa 1 1  
Caloplaca stillicidiorum 1 1 1  FrB 
Caloplaca substerilis 1  ZP
Caloplaca turkuensis 1  
Candelaria concolor 

 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Candelariella efflorescens 1111 1  
Candelariella reflexa s.str. 1 1  
Candelariella vitellina 1  
Candelariella xanthostigma 1111 1   
Catillaria erysiboides 1 log 
Catillaria nigroclavata 11 11 1  
Catinaria atropurpurea 11  
Cetrelia cetrarioides 1   
Cetrelia chicitae 1 1 1  
Cetrelia monachorum 1111111  
Cetrelia olivetorum 11111   
Chaenotheca brachypoda 1 1 1 1 
Chaenotheca furfuracea 1 1 1 1 
Chaenotheca gracilenta 1 1 1  
Chaenotheca trichialis 1 1 1 
Chaenotheca xyloxena 1  
Chaenothecopsis debilis 1 1  
Chaenothecopsis pusilla 1 1 1 snag 
Cheiromycina petri 1  ZP
Chromatochlamys muscorum 1 1  
Cladonia chlorophaea agg. 1 1 1 1  
Cladonia coniocraea 1111 1   
Cladonia fimbriata 11111  
Cladonia macilenta 1 
Cladonia pyxidata 1 1  
Cladonia subulata 1 1  
Cliostomum griffithii 1  FrB*
Coenogonium luteum 1 1   
Coenogonium pineti 11111   
Collema flaccidum 111111   
Collema nigrescens 1 1 1  

Malíek   
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Coniocarpon cinnabarinum 1 1 1  
Cresporhaphis wienkampii 1  
Cryptodiscus gloeocapsa 1 
Dictyocatenulata alba 1 1 1   
Diploschistes muscorum 1 1 Cladonia 
Evernia divaricata 1 QU
Evernia prunastri 1111 1  
Fellhanera bouteillei 11 
Fellhanera gyrophorica 1  
Flavoparmelia caperata 11111  ZP, FrB 
Frutidella furfuracea 1   
Fuscidea arboricola 1 1 1 1  
Fuscidea cyathoides 11  
Graphis scripta 11111  AA, ZP9 
*Gyalecta croatica  1  
Gyalecta flotowii 1 1 1 1   
Gyalecta herculina 
 111 111  
Gyalecta truncigena 1 1 1 1   
Gyalecta ulmi 1 QU 
*Gyalideopsis helvetica 1  ZP
Haematomma ochroleucum 
Halecania viridescens 11  
Hazslinszkya gibberulosa 111 1 1   
Heterodermia speciosa 11111   
Hypocenomyce scalaris 1 
Hypogymnia farinacea 1 1 1 
Hypogymnia physodes 11111  ZP 
Hypogymnia tubulosa 11111  
Hypotrachyna afrorevoluta 11 11 1  
Hypotrachyna revoluta 1   

160 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Imshaugia aleurites 1 snag (QU) FrB, ZP
Inoderma byssaceum  11  
Japewia sp. 1 1  
Lathagrium auriforme 1 1  
Lecania croatica 11111 Lonicera 
Lecania cyrtella 1  FrB, ZP
Lecania cyrtellina 1 1 1  
Lecania naegelii 1  FB
Lecanora albella 1 1  
Lecanora albellula 1  AA, ZP
Lecanora allophana 
Lecanora anopta 1 log ZP2*
Lecanora argentata 11111 1   
Lecanora carpinea s.str. 1 1 1   
Lecanora chlarotera 1 1 1   
Lecanora cinereofusca 1 1 1  
Lecanora ecorticata  s.l. 1 1  
Lecanora expallens 1111 1  
*Lecanora exspersa 1 1  
Lecanora glabrata 1111111   
Lecanora impudens 
Lecanora intricata 1  
Lecanora intumescens 1 1 1 1   
Lecanora leptyrodes 11111   
Lecanora phaeostigma 1  
Lecanora polytropa 1 1 1   
Lecanora pulicaris 11111   
Lecanora saligna 1 1 1  
Lecanora stanislai 
 1  
Lecanora strobilina 

Malíek   161
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Lecanora subintricata 1 snag 
Lecanora substerilis  11 11 1  
Lecanora symmicta 1 1 1   
Lecanora thysanophora 11111  
Lecanora varia 1 1 1  
Lecidea erythrophaea 1 1 1 1  
Lecidea turgidula 1  FrB, ZP2
Lecidella carpathica 1 1  
Lecidella elaeochroma L. achristotera)1111111   
Lecidella flavosorediata 1111 1  
*Lecidella subviridis s.l. 1 1 1 1  
Lepra albescens 11111   
Lepra amara 11111 1   
Lepraria eburnea 1  ZP*
Lepraria elobata 1  
Lepraria finkii 111111 

 
Lepraria incana 1 1 1 
Lepraria membranacea 1 
Lepraria rigidula 11111  
Lepraria vouauxii 11111 1  
Leproplaca chrysodeta 1 1 1 1  
Leptogium cyanescens 1 1 1  
Leptogium saturninum 11 111   
*Lithothelium hyalosporum 1  
Lobaria pulmonaria 1 1111  
Lopadium disciforme 1 1 1 1   
Megalaria laureri 1 1 1   
Melanelixia glabra 1 1 1  AA 
Melanelixia glabratula 11111   
Melanelixia subargentifera 1  

162 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Melanelixia subaurifera 1111 1   
Melanohalea elegantula 1111 1   
Melanohalea exasperata 1 1 
Melanohalea exasperatula 1 1 1 1  
Melaspileella proximella 1 1  
*Menegazzia subsimilis 11  
Menegazzia terebrata 11111   
*Micarea anterior 1 log ZP
Micarea botryoides 1 1 snag ZP2
Micarea byssacea 1 1 log AA2
Micarea cinerea 1 log ZP
*Micarea deminuta 1 log FrB, ZP
Micarea denigrata 1 log 
Micarea globulosella 1  
Micarea lithinella 1  ZP
Micarea melaena 1 snag (QU) 
Micarea micrococca 11111  
Micarea misella 1 1 1 1  
Micarea nigella 1  AA, ZP
Micarea peliocarpa 1  AA
*Micarea perparvula  1  ZP
Micarea prasina 1111 1  AA, ZP*
*Micarea soralifera 
 1 1 log FrB*, ZP*
Microcalicium arenarium 1  ZP
Multiclavula mucida 1 1 1 log  
Mycobilimbia carneoalbida 
Mycobilimbia epixanthoides 1 1   
Mycobilimbia pilularis 
Mycobilimbia tetramera 1 1  
Mycocalicium subtile 1 1 1 1 1 1 snag 

Malíek   
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Myriolecis sambuci 1 1   
Naetrocymbe punctiformis 1 1 1   
Naetrocymbe rhyponta 
Nephroma parile 1 1   
Nephroma resupinatum 1 1   
Nephromopsis laureri 1 1  
*Normandina acroglypta 1 1 1  
Normandina pulchella 1111111  ZP
Ochrolechia alboflavescens 1  
Ochrolechia androgyna agg. 1111 1   
Ochrolechia arborea 1 1 1  
Ochrolechia microstictoides 1 snag 
Ochrolechia pallescens 1 1 1 1   
Ochrolechia szatalaensis 11  
Ochrolechia trochophora 1  
Ochrolechia turneri 1 1 1  
*Opegrapha fumosa  1  ZP*
Opegrapha niveoatra 1 1 1 1  
Opegrapha trochodes 1 1 1 1  
Opegrapha vermicellifera 1 1 1 1   
Pannaria conoplea 1  
Parmelia saxatilis  P. ernstiae, P. serrana)11111   
Parmelia submontana 1 1 1 1   
Parmelia sulcata 11111  ZP2 
Parmeliella triptophylla 1111 1   
Parmelina pastillifera 1 1 1 1 1  ZP 
Parmelina tiliacea 1 1 1 1  ZP2 
Parmeliopsis ambigua 1 1 1 1  
Parmeliopsis hyperopta 1 1  
Parmotrema arnoldii 1 1  
Parmoterma crinitum 1 1  

164 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Parmotrema perlatum 1 1 1  
Peltigera collina 1 1 1   
Peltigera degenii 
Peltigera praetextata 111111 

 
Pertusaria coccodes 1 1 1  
Pertusaria constricta 1 1 1   
Pertusaria coronata 1 1 1 1   
Pertusaria flavida 1 1  
Pertusaria leioplaca 11111 1   
*Pertusaria macounii  1  
Pertusaria pertusa 1111 1 1  
Pertusaria pustulata 
Pertusaria pupillaris 1 1 1 1  
Pertusaria trachythallina 1 1 1  
Phaeophyscia chloantha 1 1 1  FB
Phaeophyscia endophoenicea 1111 1   
Phaeophyscia nigricans 1 
Phaeophyscia orbicularis 11111  ZP 
Phaeophyscia pusilloides 1  FrB
Phlyctis agelaea 1 1 1  
Phlyctis argena 11111   
Physcia adscendens 1 1 1 1 1   
Physcia aipolia 1 1 1  
Physcia dubia 1 
Physcia stellaris 1 1 1  
Physcia tenella 11111 
Physconia detersa 1  FrB, ZP 
Physconia distorta 1 1 1  
Physconia enteroxantha 1 1  
Physconia grisea 1 
Physconia perisidiosa 1 1 1   

Malíek   
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Piccolia ochrophora 11111 1   
Placynthiella dasaea 1 1 log, snag FrB*, ZP*
Placynthiella icmalea 1 1 1 1  
Platismatia glauca 1111 1  
Pleurosticta acetabulum 1 
Polycauliona polycarpa 1 1 
Porina leptalea 1 1 1 1 1   
Porina pseudohibernica 11111 1   
Porpidia macrocarpa P. nigrocruenta) 1 1  
Protoparmeliopsis muralis 1  
Pseudevernia furfuracea 11111  
Pseudosagedia aenea 11111   
*Pseudosagedia byssophila 1  
Pseudoschismatomma rufescens 1111 1   
Psoroglaena abscondita 1 1 1 log 
Psoroglaena dictyospora 1 1 snag, log 
*Psoroglaena stigonemoides 1  ZP
Punctelia jeckeri 1 1 1 
Punctelia subrudecta 11 11 1  
Pycnora sorophora 1 snag (QU)
*Pyrenula chlorospila Arnold 1 AP 
Pyrenula coryli 1 1  
Pyrenula dermatodes  1 1  
Pyrenula laevigata 1 1 1   
Pyrenula nitida 11111 1  ZP4 
Pyrenula nitidella 1 1  
Pyrrhospora quernea 
Pyxine sorediata 1  
Ramalina farinacea 11111  ZP2 
Ramalina fastigiata 1 1 1   
Ramalina fraxinea 1  

166 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Ramalina pollinaria agg. 1111 1  ZP* 
Ramonia interjecta 1 1  
Ramonia luteola 11   
Reichlingia leopoldii 
Rhizocarpon polycarpum 1  
Ricasolia amplissima 1 1  
Rinodina albana 1  
Rinodina capensis 1   
Rinodina conradii 
Rinodina efflorescens 1111111  
Rinodina griseosoralifera 1 1 1  
*Rinodina malangica 1  
Rinodina orculata 1  
Rinodina pyrina 
Rinodina sophodes 1 1 1 1   
Rinodina subparieta  111 1 1  
Rinodina trevisanii 1  ZP
Ropalospora viridis 1 1 1 1   
Sclerophora farinacea 1 1 1 1  
Sclerophora pallida 1 1 1 1   
Scoliciosporum chlorococcum 11   
Scoliciosporum sarothamni 1111 1  
*Scoliciosporum schadeanum 1 1  ZP4
Scoliciosporum umbrinum 1111 1   
Scytinium lichenoides 11  ZP 
Scytinium pulvinatum 1 1 1 1  
Scytinium teretiusculum 11111 1  
Steinia geophana 1 1 1 1 1 log, snag 
Stenocybe pullatula 1  AA 
Strangospora pinicola 1 1  
Strigula glabra 11  

Malíek   167
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Strigula stigmatella 1 1 1 1   
Tephromela atra 1  
Tetramelas chloroleucus 1   
Tetramelas insignis 
Thelocarpon epibolum 1 1 1 log 
Thelocarpon laureri 
Thelocarpon lichenicola 1 1 log 
Thelopsis flaveola 11  
Thelopsis rubella 11111 1   
Thelotrema lepadinum 1 1 1   
Thelotrema suecicum 1 1 1  
Toensbergia leucococca 1  
Trapelia corticola 1 log 
Trapeliopsis flexuosa 1 1 1 1   
Trapeliopsis granulosa 1 log
Trapeliopsis pseudogranulosa 1  AA
Trapeliopsis viridescens 1 log 
Tuckermannopsis chlorophylla 1 1  
Usnea barbata 11  
Usnea ceratina 
Usnea dasopoga 
Usnea hirta 1  
Usnea perplexans 1   
Usnea subfloridana 1 1   
Usnea wasmuthii 
Varicellaria hemisphaerica 1 1 1 1  FrB, ZP
Verrucaria breussii 1 QU ZP
*Verrucaria hegetschweileri  1  
Verrucaria viridigrana 11  
*Vezdaea retigera 1  ZP
Violella fucata 11111   

168 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Vulpicida pinastri 1 1  
Wadeana dendrographa  
Xanthomendoza fulva 1 1  
Xanthomendoza ulophyllodes 
Xanthoria parietina 1 1 1  
*Xylographa trunciseda 1 log 
Zwackhia viridis 1 1 1 1   
lichenicolous fungi
Abrothallus bertianus De Not. 1 1 1 Melanelixia glabratula 
*Arthonia biatoricola  1 1 Lecania croatica, Biatora
chrysantha)ZP2
Briancoppinsia cytospora 
 1 1 Parmelia FrB2
*Chalara lobariae  1 Lobaria pulmonaria FrB
*Dactylospora homoclinella (Nyl.) Hafellner 1 Lecanora sp. FrB
Dactylospora lobariella (Nyl.) Hafellner 1 Lobaria pulmonaria FrB
Dactylospora parasitica 1 Pertusaria FrB
Homostegia piggotii  1 Parmelia saxatilis FrB
Intralichen christiansenii 
 1 Lecanora leptyrodes FrB
Intralichen lichenum
 1 Lobaria pulmonaria FrB
Lichenochora weillii  1 Physconia AA
Lichenoconium erodens  1 Parmelia saxatilis FrB
Lichenodiplis lecanorae
 1 Lecanora FrB
Lichenostigma maureri Hafellner 1 Parmelia saxatilis FrB
*Merismatium heterophractum 1 Biatora sp. FrB
*Opegrapha thelotrematis  1 Thelotrema lepadinum FrB
Phaeopyxis punctum 
 1 Cladonia  AA
*Phoma lobariae  1 Lobaria pulmonaria FrB

Malíek   169
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
*Rhymbocarpus neglectus 
 1 Lepraria FrB
*Sphinctrina leucopoda Nyl. 1 Pertusaria pertusa 
Sphinctrina turbinata (Pers.) De Not. 1 Pertusaria leioplaca FrB
Stigmidium microspilum  1 Graphis scripta
Taeniolella punctata  1 1 1 Graphis scripta 
*Taeniolella toruloides  11 Thelotrema lepadinum FrB2
Tremella lichenicola  1 1 Violella fucata FrB
Tremella lobariacearum 1 1 Lobaria pulmonaria FrB
Trichonectria anisospora
 1 Hypogymnia physodes FrB
Unguiculariopsis acrocordiae
 1 1 1 Acrocordia gemmata 
Vouauxiella lichenicola  1 1 Lecanora pulicaris FrB, ZP
*Xenonectriella septemseptata
den Boom 1 Melanelixia glabratula FrB
other non-lichenized fungi
Bertia moriformis (Tode) De Not. 1 log ZP
Capronia moravica 
 1 log AA
Cryptocoryneum condensatum 
 1  ZP
Cryptodiscus foveolaris  1 1 1  AA, FrB2, ZP2
Cryptodiscus pallidus  1 log AA
Cryptodiscus pini 
 1  FrB
Durella connivens 1 log ZP
Durella melanochlora  1 log ZP
Exarmidium inclusum (Pers.) Aptroot 1 1 log 
Glonium lineare (Fr.) De Not. 1 log, snag AA
Hyalotrochophora lignatilis 
 1 log FrB

170 Herzogia 31 (1) Teil 1, 2018
species 1 2 3 4 5 6 7 8 substrate vouchers Dymytrova
et al. (2013)
Kirschsteiniothelia aethiops  1  ZP
Kirschsteiniothelia recessa 
 1  
Lophiostoma corticola 
 1  ZP
Mniaecia jungermanniae 1 log 
Mycowinteria muriformis Aptroot 1 log AA
Perididiothelia fuliguncta 1 1 1  AA
Stictis radiata 1 1 1  

Malíek   171
Appendix 2.
Species Voucher nrITS mtSSU
Biatora bacidioides ZP 19221  
Biatora bacidioides   
Biatora bacidioides  –
Biatora efflorescens   
Biatora longispora   
Biatora pontica   
Bryostigma apateticum   
Caloplaca monacensis   
Caloplaca sorocarpa   –
Caloplaca substerilis ZP 19680 – 
Caloplaca turkuensis   –
Japewia sp.   
Lecanora  anopta ZP 20047 – 
Lecanora exspersa   
Lecanora exspersa   –
Lecanora intumescens   
Lecanora pulicaris  –
Lecanora stanislai  –
Lecanora substerilis  –
Lecanora substerilis  –
Lecanora substerilis   
Lecanora thysanophora  –
Lecidella subviridis s.l.  –
Lecidella subviridis s.l.  –
Lecidella subviridis s.l.  –
Lepra amara ZP 19422  –
Melaspileella proximella   
Melaspileella proximella   –
Mycobilimbia epixanthoides   