<8) MycoKeys MycoKeys 124: 193-226 (2025) DOI: 10.3897/mycokeys.124.162802 Research Article Unexpected species richness of the lichen genus Protoblastenia (Lecanorales, Psoraceae) in Finland Juha Pykala'™®, Leena Myllys2© 1 Nature Solutions, Finnish Environmental Institute, Latokartanonkaari 11, Fl-00790 Helsinki, Finland 2 Botany and Mycology Unit, Finnish Museum of Natural History, P. 0. Box 7, Fl-00014 University of Helsinki, Helsinki, Finland Corresponding author: Juha Pykala (juha.pykala@syke. fi) OPEN Qaceess Academic editor: Thorsten Lumbsch Received: 22 June 2025 Accepted: 3 September 2025 Published: 29 October 2025 Citation: Pykala J, Myllys L (2025) Unexpected species richness of the lichen genus Protoblastenia (Lecanorales, Psoraceae) in Finland. Mycokeys 124: 193-226. https://doi. org/10.3897/mycokeys.124.162802 Copyright: © Juha Pykala & Leena Myllys. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract The taxonomy of Protoblastenia in Finland was studied, based on morphology and mo- lecular data (nulTS rDNA sequences). Twenty species were recognised, with sixteen spe- cies being newly described here: P arupii sp. nov., P borealis sp. nov., PR compressa sp. nov., P dolomitica sp. nov., P ekmanii sp. nov., P fennoarctica sp. nov., P minuta sp. nov., P oulankaensis sp. nov., P pseudocompressa sp. nov., P pseudoterricola sp. nov., P remo- ta sp. nov., P. rikkinenii sp. nov., P saanaensis sp. nov., P. timdalii sp. nov., P violacea sp. nov. and P westbergii sp. nov. All species are confined to calcareous rocks, except P terri- cola which also grows on calcareous soil. The calcareous fells in Enontekio in NW Finland were identified as hot spots of Protoblastenia diversity. Nine newly-described species (P arupii, P fennoarctica, P minuta, P pseudoterricola, P. rikkinenii, P saanaensis, P. timdalii, P violacea and P. westbergii) are restricted to this area in Finland. Several of the species are semi-cryptic. On average, they may have minor morphological differences, but many specimens cannot be identified, based on morphology only. Protoblastenia rikkinenii is reported from Norway and P oulankaensis from Italy, based on GenBank sequences. Full descriptions and a preliminary key of Protoblastenia in Finland are provided. Key words: Calcareous rocks, DNA barcoding, Italy, ITS, lichenised fungi, new species, Norway, phylogeny Introduction Protoblastenia (Zahlbr.) J. Steiner is a lichenised genus in the family Psoraceae. The genus includes ca. 15-25 accepted, mainly calcicolous, species (Castello and Nimis 1995; Kainz 2004; Kainz and Rambold 2004). Most species occur on calcareous rocks and a few on soil over calcareous rock. Species of Protoblastenia have anthraquinones in the apothecia, causing pre- dominantly orange-coloured discs. Other typical features are predominantly convex apothecia and pale, non-septate medium-sized spores (Kainz and Rambold 2004). Protoblastenia differs from the closely-related genus Psora Hoffm. in the absence or weakly developed upper cortex in the thallus and the absence of calcium oxalate crystals in the hypothecium (Timdal 1987; Kainz and Rambold 2004). Based on the phylogenetic studies of Kainz and Rambold (2004), Ekman and Blaalid (2011) and Miadlikowska et al. (2014), Protoblastenia forms a monophyletic group. 193 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Kainz and Rambold (2004) studied the phylogeny and taxonomy of Protoblaste- nia in Central Europe. They ITS-sequenced 11 species: Protoblastenia aurata Poelt & Vézda, P. calva (Dicks.) Zahbr., P calvella Kainz & Rambold, P cyclospora (Hepp ex Korb.) Poelt, P incrustans (DC.) J. Steiner, P laeta (Poelt) Kainz & Rambold, P lilacina Poelt & Vézda, P pseudoincrustans ined., P rupestris (Scop.) J. Steiner, P siebenhaariana (KOrb.) J. Steiner and P terricola (Anzi) Lynge. Three European species, Protoblastenia coniasis (A. Massal.) Poelt (Poelt 1957), P geitleri Zahlbr. (Zahlbruckner 1936) and P. szaferi J. Nowak (Nowak 1974), were not sequenced. So far, seven species of Protoblastenia are known in Finland: P calva, P calvella, P incrustans, P lilacina, P rupestris, PR siebenhaariana and P terricola (Stenroos et al. 2016; Pykala 2023). However, during the DNA barcoding of the Finnish li- chen biota, it became apparent that the genus is much in need of revision. Here, we present the results of the revised taxonomy and show that the number of species in Finland is 20, almost three times more than previously reported. Material and methods Taxon sampling This study is based on material collected by the first author (JP) during the years 1990-2022 in Finland. The specimens are stored in H. Morphology Apothecia and thalli were hand-sectioned with razor blades. The sections were examined and measured in tap water. Asci and ascospores were also studied in squash preparations of apothecia mounted in water. Spores were measured using 10% potassium hydroxide (KOH). Additionally, the thickness of the hypothecium was examined by cutting the apothecia into two pieces and studying the pieces using a binocular microscope. The range of spore size is indicated as arithmetic mean and standard deviation. Minimum and maximum values are given in parentheses. The size of the apothecia (in diameter) is given in surface view. DNA extraction and sequencing Total genomic DNA was extracted from the apothecia (1-3) of 1 — 31-year-old herbarium specimens. We used two different techniques for extraction and sequencing. Most Finnish samples were sequenced as part of the “Finnish Barcode of Life” re- search project conducted from 2012-2023 (FinBOL, https://finbol.org/). The samples were placed on 96-well microplates and sent to the Canadian Centre for DNA Barcoding (CCDB). CCDB’s standard protocols (documentation available at http://ccdb.ca/resources) were used for extraction, PCR and se- quencing. The primers ITS1 and ITS4 (White et al. 1990) were used both for PCR and sequencing of the nuclear ribosomal ITS region. The barcode sequences and their trace files along with all relevant collection data and photographs of the voucher specimens were uploaded to the Barcode of Life Data Systems (BOLD, https://www.boldsystems.org) database. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 194 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland DNA of the 31 specimens (Pykala 5963, 8547, 29704, 33037, 34043, 35585, 39693, 39851, 39987, 41511b, 43105, 43194, 53252, 43340, 43524, 43605, 43624, 43855, 43910, 44115, 44130, 44177, 44275, 44314, 44431, 44438, 44876, 46413, 57501, 58392, 58457) was extracted using DNeasy® Blood & Tissue kits by Qiagen following the protocol described in Myllys et al. (2011). PCR reactions were prepared using PuReTaq Ready-To-Go PCR beads (GE Healthcare). The 25 pl reaction volume contained 19 ul of dH20, 0.4 uM of each primer and 4 ul of extracted DNA. PCR was run under the following conditions: initial denaturation for 5 min at 95 °C followed by five cycles of 30 s at 95 °C (de- naturation), 30 s at 58 °C (annealing) and 1 min at 72 °C (extension); in the re- maining 35 cycles, the annealing temperature was decreased to 56 °C; the PCR schedule ended with a final extension for 7 min at 72 °C. PCR products were pu- rifled and sequenced by Macrogen Inc. (Amsterdam, the Netherlands; https:// www.macrogen.com) or alternatively cleaned with ExoSAP (Affymetrix, Santa Clara, California, USA) and sequenced by FIMM Genomics (https://www2.hel- sinki.fi/en/infrastructures/genome-analysis/infrastructures/fimm-genomics). The primers ITS1F (Gardes and Bruns 1993) and ITS4 (White et al. 1990) were used both for PCR amplification and sequencing of the ITS regions. Phylogenetic analyses All the ITS sequences of Protoblastenia available in GenBank (n = 55) were downloaded (1.3.2025). Psora decipiens (Hedw.) Hoffm. and P. rubiformis (Ach.) Hook. were used as outgroups, based on the phylogenies of Miadlikow- ska et al. (2014) and Evankow et al. (2025). A total of 141 ITS sequences were aligned (Table 1) with MUSCLE v.3.8.31 (Edgar 2004) using EMBL-EBI’s freely available web service (http://www.ebi. ac.uk/Tools/msa/muscle/). The aligned dataset was subjected to Maximum Likelihood analysis (ML). The analysis was performed with RAXML v.8.1.15 (Stamatakis 2014) on the CSC — IT Center for Science server ((http://www. csc.fi). The ITS region was divided into three partitions: ITS1, 5.8S and ITS2. These partitions were analysed under the universal GTR-GAMMA model. Node support was estimated with 1000 bootstrap replications using the rapid bootstrap algorithm. Branches with bootstrap values = 70% were considered strongly supported. Results and discussion We generated 83 new ITS sequences obtained from the Finnish Protoblastenia specimens. In the ITS phylogeny (Fig. 1), the sequenced Finnish specimens were divided into 20 lineages of which 16 do not match any described species. These lineages, when represented by multiple samples, received high support values (96-100%) and are described here as new species. Typically, infraspe- cific differences in ITS in the Finnish Protoblastenia species are small. As expected, the phylogeny, based on only ITS sequences, remains unsup- ported at deeper nodes. However, six well-supported (ML bootstrap support > 70%) lineages represented by more than one species were identified: 1) P. fen- noarctica sp. nov. + P. rikkinenii sp. nov. + P saanaensis sp. nov. + P. timdalii sp. nov. + P pseudoincrustans ined. 2) P dolomitica sp. nov. + P westbergii sp. nov., MycoKkeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 195 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Table 1. Specimens used in the phylogenetic analyses. New sequences are in bold. Taxon Protoblastenia arupii P. aurata P. aurata P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. borealis P. calva P. calva P. calva P. calva P calva P. calva “P. calva” P calvella P calvella P calvella P calvella P calvella P calvella P calvella P compressa P compressa P. compressa P. compressa P compressa P compressa MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 Country Finland Austria Austria Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Ireland Austria Germany France Germany Norway Croatia Germany Germany Austria Germany Finland Finland Sweden Finland Finland Finland Finland Finland Finland Voucher specimen Pykald 44130 (H) Kainz 950 (M) Wittmann (M) Pykala 16567 (H) Pykald 23710 (H) Pykala 25292 (H) Pykala 26097 (H) Pykala 29704 (H) Pykala 32757 (H) Pykala 33037 (H) Pykald 33101 (H) Pykald 38784 (H) Pykala 39151 (H) Pykald 39942 (H) Pykald 40695 (H) Pykala 41355 (H) Pykala 42802 (H) Pykald 44641 (H) Pykald 44906 (H) Pykala 45759 (H) Pykala 47217 (H) Pykala 57501 (H) Pykald 58196 (H) Pykala 58392 (H) Pykald 60511 & Kuusisto (H) Sipman 30678 (B) Kainz 981 (M) Kainz 501 (M) Rambold 6242 (M) Kainz 852 (M) Edvardsen & Ekman NO6 (BG) Gueidan CG659 (DUKE) Kainz 764 (M) Kainz 765 (M) Kainz 967 (M) Kainz 805 (M) Pykald 44807 (H) Pykald 43503 (H) Ekman & Eide Ekman 5821 (UPS) Pykala 35585 (H) Pykald 36521 (H) Pykald 38349 (H) Pykala 38646 (H) Pykald 38965 (H) Pykald 41376 (H) PV766636 AY425658 AY425659 PV766637 PV766638 PV766639 PV766640 PV766641 PV766642 PV766643 PV766644 PV766645 PV766646 PV766647 PV766648 PV766649 PV766650 PV766651 PV766652 PV766653 PV766654 PV766655 PV766656 PV766657 PV766658 AY425628 AY425642 AY425643 AY425644 AY425645 EF524319 HQ650618 AY425646 AY425647 AY425648 AY425649 0Q604661 0Q604662 OR773065 PV766659 PV766660 PV766661 PV766662 PV766663 PV766664 GenBank Accession number (ITS) 196 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Taxon P compressa P compressa P. cyclospora P cyclospora P. dolomitica P. dolomitica P. dolomitica P. dolomitica P. dolomitica P. dolomitica P. dolomitica P. ekmanii P fennoarctica P. fennoarctica P fennoarctica P fennoarctica P incrustans P incrustans P incrustans P incrustans P incrustans P incrustans P incrustans P laeta P laeta P laeta P laeta “P. lilacina” P lilacina P lilacina P lilacina P lilacina P lilacina P lilacina P lilacina P lilacina P minuta P. oulankaensis P oulankaensis P. oulankaensis P. oulankaensis P. oulankaensis P oulankaensis P. oulankaensis P. pseudocompressa P. pseudocompressa P pseudocompressa MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 Country Finland Finland Italy Greece Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Svalbard Svalbard Germany Germany Germany Germany Germany Germany Germany Germany France Italy France Germany Greece Germany Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Voucher specimen Pykala 41511b (H) Pykald 42124 (H) Tretiach (M) Kainz 913 (M) Pykala 36072 (H) Pykalad 39217 (H) Pykald 39851 (H) Pykalad 43340 (H) Pykald 43466 (H) Pykala 43855 (H) Pykala 44115 (H) Pykala 37508 (H) Pykalad 43105 (H) Pykalad 43194 (H) Pykala 43656 (H) Pykala 44116 (H) Zhang Zhang ZT2013200 Kainz 498 (M) Kainz 855 (M) Kainz 842 (M) Kainz 228 (M) Kainz 220 (M) Kainz 856 (M) Kainz 737 (M) Kainz 741 (M) Rambold 6253 (M) Kainz 161 (M) Sipman 22890 (B) Kainz 809 (M) Kainz 900 (M) Kainz 788 (M) Pykald 27009 (H) Pykald 27629 (H) Pykala 37012 (H) Pykala 57976 (H) Pykala 44275 (H) Pykala 39629 (H) Pykald 39810 (H) Pykala 40201 (H) Pykala 44928 (H) Pykald 44949 (H) Pykala 44953 (H) Pykala 55995 (H) Pykala 35901 (H) Pykala 36325 (H) Pykala 39367 (H) GenBank Accession number (ITS) PV766665 PV766666 AY425667 AY425668 PV766667 PV766668 PV766669 PV766670 PV766671 PV766672 PV766673 PV766674 PV766675 PV766676 PV766677 PV766678 KP314370 KP314440 AY425630 AY425631 AY425632 AY425669 AY425670 AY425650 AY425651 AY425652 AY425653 AY425629 AY425660 AY425661 AY425662 AY425663 PV766679 PV766680 PV766681 PV766682 PV766683 PV766684 PV766685 PV766686 PV766687 PV766688 PV766689 PV766690 PV766691 PV766692 PV766693 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Taxon P. pseudocompressa P. pseudocompressa P. pseudocompressa P pseudoincrustans ined. P pseudoincrustans ined. P. pseudoincrustans ined. P. pseudoterricola P remota P remota P rikkinenii P rikkinenii P. rikkinenii P rupestris P rupestris P rupestris P rupestris P rupestris P rupestris “P. rupestris” P rupestris P rupestris “P. rupestris” P rupestris P rupestris P rupestris P rupestris P rupestris P rupestris “P. aff. rupestris” “P. aff. rupestris” P. saanaensis P. saanaensis P. siebenhaariana P terricola P terricola P terricola P terricola P. timdalii P timdalii P violacea P violacea P violacea P westbergii P westbergii P westbergii P westbergii Psora rubiformis P. decipiens Country Finland Finland Finland Austria Germany Germany Finland Finland Finland Finland Finland Finland Germany Germany Germany France Norway France Norway Canada Norway UK Spain USA Finland Finland Finland Finland USA USA Finland Finland Germany Austria Austria Switzerland Finland Finland Finland Finland Finland Finland Finland Finland Finland Finland Norway Greenland Voucher specimen Pykala 39693 (H) Pykald 39987 (H) Pykala 58457 (H) Kainz 949 (M) Kainz 488 (M) Kainz 489 (M) Pykald 44157 (H) Pykala 8547 (H) Pykald 39455 (H) Pykala 43252 (H) Pykald 43910 (H) Pykala 44387 (H) Kainz 714 (M) Kainz 791 (M) Kainz 773 (M) Rambold 6252 (M) Edvardsen & Ekman NO8 (BG) Gaya 415 & Guiedan (BCN) Millanes (O) McMullin (OAC) Johnsen (BG) Cannon (k) Garrido-Benavent IGB1018 (VAL) Lendemer 49650 (NY) Pykala 5963 (H) Pykald 27569 (H) Pykala 34043 (H) Pykald 46413 (H) Leavitt SL19096 Leavitt SL19099 Pykald 44431 (H) Pykald 44438 (H) Kainz 188 (M) Hafellner 42151 (GZU) Kainz 977 (M) Sippman 30291 (B) Pykald 44876 (H) Pykala 43624 (H) Pykald 44177 (H) Pykald 43015 (H) Pykala 43524 (H) Pykald 43605 (H) Pykald 43196 (H) Pykala 43279 (H) Pykald 44314 (H) Pykald 44316 (H) Timdal (0) Timdal 10078 (0) MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 PV766694 PV766695 PV766696 AY425664 AY425665 AY425666 PV766697 PV766698 PV766699 PV766700 PV766701 PV766702 AY425654 AY425655 AY425656 AY425657 EF524318 EU639582 KY266981 KT695366 MG926010 MZ159571 0Q550133 MK092337 PV766703 PV766704 PV766705 PV766706 MZ922194 MZ922195 PV766707 PV766708 AY425627 AY425639 AY425640 AY425641 PV766709 PV766710 PV766711 PV766712 PV766713 PV766714 PV766715 PV766716 PV766717 PV766718 KY266957 EF524326 GenBank Accession number (ITS) 198 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland 3) P compressa sp. nov. + P pseudocompressa sp. nov., 4) P. lilacina + P. ou- lankaensis sp. nov., 5) P. borealis sp. nov. + P ekmanii sp. nov. + P. rupestris, 6) P calva + P. calvella + P pseudoterricola sp. nov. + P terricola + P violacea sp. nov. All the new Finnish species with more than one sequence show a barcode gap, i.e. their genetic distance to the nearest neigbour exceeds their maximum intraspe- cific distance (Table 2). Note that the gap is slightly smaller if GenBank sequences here identified as P oulankaensis and P. remota are included. The maximum ge- netic distance within the species including the Italian sequence of P oulankaensis (AY425629) is 1.7%. If the two North American sequences (MZ922194, MZ922195) are included in P remota, the maximum genetic distance of P remota is 1.6%. The specimens previously treated as P siebenhaariana by Stenroos et al. (2016) belong to six different species, all characterised by a violet hypothe- cium: Protoblastenia arupii sp. nov., P dolomitica sp. nov., P pseudoterricola sp. nov., P. terricola, P violacea sp. nov. and P. westbergii sp. nov. However, all these species have more densely occurring apothecia than P. siebenhaariana. Furthermore, the specimens previously identified as P incrustans belong to P compressa sp. nov. and P pseudocompressa sp. nov. Specimens treated as Protoblastenia rupestris belong in nine species: P rupes- tris, P calvella, P borealis sp. nov., P fennoarctica sp. nov., PR minuta sp. nov., P. re- mota sp. nov., P rikkinenii sp. nov., P saanaensis sp. nov. and P timdalii sp. nov. The sequenced specimens previously identified as P. calva belong to P lilacina, P oulan- kaensis sp. nov. and P pseudocompressa. Previously only Finnish specimens on soil have been identified as P terricola, but the species is also epilithic in Finland. Unfortunately, sequences of P coniasis, P. geitleri and P szaferi were not available in GenBank. However, all three species differ morphologically from our new species: Protoblastenia coniasis resembles P incrustans, but has a yellow thallus that gradually turns brownish-rose in the herbarium (Poelt 1957). Table 2. Genetic distances within and between the new species of Protoblastenia, based on the Finnish and GenBank sequences. P arupii (n = 1) P borealis (n = 22) P. compressa (n = 8) P. dolomitica (n = 7) P. ekmanii (n = 1) P. fennoarctica (n = 4) P minuta (n = 1) P. oulankaensis (n = 7) P. pseudocompressa (n = 6) P. pseudoterricola (n = 1) P. remota (n = 2) P. rikkinenii (n = 3) P. saanaensis (n = 2) P. timdalii (n = 2) P. violacea (n = 3) P westbergii (n = 4) Maximum intraspecific | Minimum distance to distance (%) nearest neighbour (%) Barcodegap (6) Meare Sineigneour 9.9 P. dolomitica 0.6 250 1.9 P. ekmanii 0.8 2.0 1.2 P. pseudocompressa 1.1 3.9 2.8 P. westbergii 25 P. borealis 0.8 4.2 3.4 P. rikkinenii 9.2 P timdalii ded 9.6 8.4 P lilacina 0.6 2.0 1.4 P. compressa 4.9 P terricola 0.4 14.0 13.6 P incrustans 0.6 4.4 3.8 P. fennoarctica 0.2 5.2 5.0 P. timdalii 0.7 2.8 2.1 P. pseudoincrustans ined. 0.2 6.9 6.7 P terricola 2:2 4.6 2.4 P. dolomitica MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 199 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Psora decipiens EF 524326 Psora rubiformis KY266957 Protoblastenia siebenbaariana AY425627 100 P. aurata AY425659 P. aurata AY 425658 92, P. rikkinenii 43252 P. rikkinenii 43910 79 P. rikkinenii 44387 *P. rupestris” KY266981 100 P. fennoarctica 44116 P. fennoarctica 43105 70 P. fennoarctica 43656 P. fennoarctica 43194 . 87 P. pseudoincrustans ined. AY425564 29 P. timdalii 43624 94 P. timdalii 44177 99 —P. pseudoincrustans ined. AY425565 P. pseudoincrustans ined. AY425566 P. saanaensis 44431 P. saanaensis 44438 P, minuta 44275 a 004P. westbergii 43196 96 P. westbergii 43279 P. westbergii 44316 99 P. westbergli 44314 P. dolomitica 44115 96 P. dolomitica 36072 P. dolomitica 39851 P. dolomitica 43855 P. dolomitica 43340 P. dolomitica 39217 P. dolomitica 43466 P. arupii 44130 b 100 P. incrustans AY425630 100 _4P. incrustans AY425670 71 P. incrustans AY425669 100 P. incrustans KP314440 P. incrustans KP314370 gg ]P.incrustans AY425632 P. incrustans AY425631 P. pseudocompressa 39367 100 JP. psc 36325 . pseudocompressa 39987 P. pseudocompressa eee 100 P. pseudocompressa BS P. pseudocompressa 35901 P. compressa 36521 P. compressa 35585 P. compressa 42124 98 JP. compressa 38349 P. compressa 38965 97|°. compressa 38646 P. compressa 41376 E 1511 P. compresses P. oulankaensis 44949 P. oulankaensis 39810 P. oulankaensis 40201 97]P. oulankaensis 39629 P oulankaensis 55995. 100 P. oulankaensis 44953 “P. lilacina’ AY425629 P. oulankaensis 44928 P. lilacina 57976 97 P. lilacina 27009 P. lilacina 27629 951 P. lilacina AY425660 P. lilacina 37012 1007 VP. lilacina AY425661 P. lilacina AY425663 P. lilacina AY425662 P. rupestris 46413 P. rupestris 27569 100 P. rupestris AY425656 P. rupestris AY425657 P. rupestris MG926010 P. rupestris 5963 P. rupestris AY425655 P. rupestris EF524318 7 P. rupestris MK092337 P. rupestris OQ550133 74 ; P. rupestris KT695366 P. rupestris AY425654 P. rupestris 34043 P. rupestris EU639582 P. ekmanii 37508 P. borealis 44906 P. borealis 57501 P. borealis 29704 99 P. borealis 16567 P. borealis 40695 P. borealis 47217 P. borealis 33101 P. borealis 44641 P. borealis 60511 gg JP. borealis 41355 P. borealis 33037 P. borealis 23710 . borealis 42802 borealis 45759 borealis 26097 borealis 38784 borealis 39151 borealis 32757 borealis 39942 borealis 58196 borealis 25292 . borealis 58392 91-P. remota 39455 100 P. remota 8547 P. aff. rupestris MZ922194 P. aff. rupestris MZ922195 100 P. cyclospora AY425668 P. cyeisnore AY425667 . laeta AY425652 90 Up. laeta AY425650 P. laeta AY425651 P. laeta AY425653 VVUUV UU UU 100 ’P. rupestris” MZ159571 87 0.07 MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 100 P. calva AY425642 58 P. calva AY425644 67 P. calva AY425645 100 P. calva AY425643 P. calva EF 524319 P. calva AY425628 95 fF. terricola AY425639 P. terricola AY425641 100 P. terricola AY425640 P. terricola 44876 : 100 P. violacea 43015 75 P. violacea 43605 P. violacea 43524 P. pseudoterricola 44157 99 »P. calvella OR773065 P. calvella AY425649 1 99 83 P. calvella AY425648 P. calvella OQ604662 P. calvella OQ604661 95 91 4P. calvella AY425646 P. calvella AY425647 »P. calva” HQ650618 200 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Table 3. The most important apothecial characters between the new species of Protoblastenia. P. aruplii P. borealis P compressa P. dolomitica P. ekmanii P. fennoarctica P minuta P. oulankaensis P. pseudocompressa P. pseudoterricola P remota P_rikkinenii P. saanaensis P. timdalii P. violacea P. westbergii Mean spore length and width Size of hypothecium (min and Width of apothecia (min and max) (um) max) (um) (mm) 12.8 x 6.8 50-60 032260:5 11.2 x 6.2 40-160 O92=1:0 11.4x 5.9 40-100 0.2-0.8 12.8 x 5.4 40-100 O.15=6:7 10.2 x 5.9 120-200 0.4-0.8 10.0 x 5.3 60-280 0:3=0.9 7.9X 4.5 70-100 0.2-0.4 13.9 x 7.4 60-220 0.3-0.9 10.6 x 5.5 50-120 0.2-0.7 9.9x5.5 60-100 0.3-0.8 11.4 x 5.6 100-200 0.2-0.6 9.6x 5.1 80-300 0.3-0.8 8.4x4.8 40-100 0.3>0.5 9.2x48 60-100 0.3-0.8 9.2x4.6 100-280 0.3=1.0 12.0 x 6.1 60-150 OST Protoblastenia geitleri has globose spores (Zahlbruckner 1936), a feature not present in any of the Finnish species. Protoblastenia szaferi is characterised by an endolithic thallus, mainly flat apothecia immersed in rock and relatively distinct apothecial margins (Hafellner 2006). Such a combination of morpho- logical characteristics was not observed in the Finnish material. The number of new species is surprisingly high, particularly compared to the global number of accepted species. Even though the new species are geneti- cally distinct, their morphological identification may be difficult. On average, all species may differ morphologically, but for several species, more material is needed to confirm this. In addition, there is considerable overlap in morphologi- cal characteristics between the species. Most of the species may be semi-cryp- tic, a feature that may be common in lichens (e.g. Orange (2012); Pykala et al. (2020)). Some specimens can be identified by morphology, while identification of other specimens remains uncertain. Table 3 presents the most important apothecial characters between the new species. Numerous studies have shown that DNA barcoding is efficient in species de- limitation and in finding morphologically cryptic or semi-cryptic species (e.g. Diva- kar et al. (2016); Licking et al. (2020a); Zhang et al. (2022); Printzen et al. (2023)). However, in several lichen groups, lack of resolution in recently evolving species complexes may occur (Lticking et al. 2020a, b). Thus, lineages receiving high sup- port values in the ITS phylogeny and showing a barcode gap can be described as new species even if they are morphologically cryptic or semi-cryptic. Our results show that ITS is well suitable for species delimitations of Protoblastenia. Figure 1. Phylogenetic relationships of the studied Protoblastenia species, based on a Maximum Likelihood (ML) analysis of the ITS dataset. Maximum likelihood bootstrap values > 50% are shown at nodes. Thickened lines indicate bootstrap values > 70%. New species described in this study are indicated with shadowed boxes. GenBank accession numbers for sequences downloaded from GenBank and collection numbers for the specimens sequenced for this study are shown after the taxon names. Holotype specimens of the new species are shown in bold. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 201 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland The calcareous fells in Enontekio in NW Finland were identified as hot spots of Protoblastenia diversity. Nine newly-described species are restricted to this area in Finland. The importance of the area for rare lichens has been previously shown in several studies (see, for example, Pykala (2010); Pykala et al. (2017); Pykala and Lommi (2021)). Taxonomy The species descriptions are based on the sequenced specimens collected in Finland. All specimens are deposited in H. The ITS sequences of only four spe- cies present in GenBank match the Finnish specimens: Protoblastenia calvel- la, P. lilacina, P rupestris and P terricola. However, P. calvella may be hetero- geneous. More material is needed before any taxonomic conclusions can be made. No specimens whose barcode matched GenBank specimens of P calva, P incrustans or P. siebenhaariana were found. The Finnish specimens also dif- fer from the morphological descriptions of these species (Kainz and Rambold 2004; Cannon et al. 2022). Thus, these species have been deleted from the Finnish lichen biota. Based on ITS sequences, P calva in GenBank may be di- vided into two species: one sequenced from Central Europe, the other from Ireland and Norway. Similarly, in our phylogeny, the cited GenBank sequences of P incrustans seem to represent three species. Protoblastenia arupii Pykala & Myllys, sp. nov. MycoBank No: 859623 Fig. 2A Diagnosis. Differs from P dolomitica and P. westbergii in having only a slightly rimose thallus. Type. FINLAND * Enonteki6n Lappi, Enonteki6, Kilpisjarvi, Saana, fell, steep NE- slope, dolomite rock outcrop, on NE-facing wall, 820 m a.s.I., 69°02'N, 20°51'E, 11 August 2011, J. Pykala 44130 (H9250904 — holotype, GenBank accession number: PV766636). Description. Thallus grey, slightly rimose, ca. 0.02-0.05 mm thick, K-, UV-. Apothecia orange, 0.2-0.5 mm, slightly convex to convex, 1/2-immersed in thallus; ca. 60 apothecia / cm?. Epihymenium brown-yellow to orange-brown, 12-15 um thick, K+ violet. Hymenium 50-60 um thick. Hypothecium red-vi- olet, ca. 50-60 um thick. Paraphyses ca. 1.5-2 um thick, apex not thick- ened. Ascospores O-septate, (10.3-)11.8-12.8-13.8(-15.1) x (5.3-)5.7- 6.8-7.8(-9.4) um (n = 27). Habitat and distribution. Only one specimen has been found on the calcare- ous Saana fell in NW Finland. The species was associated with Polyblastia sp. and Rhizocarpon petraeum (Wulfen) A. Massal. Etymology. The specific epithet is in honour of Dr Ulf Arup (Lund) for his many important contributions on the taxonomy (particularly Teloschistaceae and Lecanoraceae) and biogeography of especially Fennoscandian lichens. Notes. Protoblastenia arupii is related to P dolomitica and P. westbergii in the ITS tree, although the relationship remains unsupported. The only known spec- imen is characterised by a thin red-violet hypothecium and large spores, some MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 202 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland of them particularly broad. Protoblastenia dolomitica and P. westbergii differ in their rimose to areolate thallus and slightly narrower spores. An ITS marker as a DNA barcode may be needed for unambiguous identification. Protoblastenia borealis Pykala & Myllys, sp. nov. MycoBank No: 859624 Fig. 2B Diagnosis. Differs from P rupestris in having a mostly rimose thallus. Type. FINLAND * Varsinais-Suomi, Lohja, Lylyinen, Saaksniemi, N of summer cottages, small calcareous rock outcrop, on SW-slope, 50 m az.s.I., 60°15'N, 23°57'E, 16 September 2003, J. Pykala 23710 (H9250905 — holotype, UPS - isotype, GenBank accession number: PV766638). Description. Thallus white to pale grey, endolithic (rarely), rimose (usually) to are- olate, areoles 0.1-0.8 mm, ca. 0.02-0.2 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells 5-12(-17) um. Apothecia yellow-orange to dirty brown-orange, 0.2- 1.0 mm, slightly convex to strongly convex, ¥-immersed in thallus to superficial, occasionally leaving shallow pits, K+violet; ca. 30-140 apothecia / cm?. Epihyme- nium orange-brown, 10-18 um thick, K+ violet. Hymenium 50—90(-100) um thick. Hypothecium colourless to pale yellow, rarely lower part reddish-brown, occasion- ally with oil droplets, ca. 40-160 um thick. Paraphyses ca. (1.5-)2-2.5 pm thick, apex not or slightly thickened, sparingly branched and anastomosing. Ascospores 0-septate, (7.6-)9.6-11.2-12.7(-16.1) x (4.8-)5.4-6.2—7.0(—-7.7) pm (n = 146). Habitat and distribution. This species is the most common species of Protoblastenia in Finland. It occurs from the hemi-boreal zone to middle Lapland. It grows on calcareous and calciferous rocks on both sun-exposed and shady habitats, but may prefer the latter. It also often grows in lime quar- ries. Specimens from concrete probably also belong in this species, but no se- quences of them are available. Typical companion species include, for example, Acarospora glaucocarpa (Ach.) K6rb., Clauzadea monticola (Schaer.) Hafellner & Bellem., Farnoldia jurana (Schaer.) Hertel, Lathagrium fuscovirens (With.) Otdalo- ra, P. M. Jorg. & Wedin, Phaeophyscia sciastra (Ach.) Moberg, Scytinium parvum (Degel.) Otdlora, PR M. Jg@rg. & Wedin, Verrucaria deversa Vain., V. epilithea Vain., V. muralis Ach. and V. nigrescens Pers. Etymology. The name refers to the geographic distribution of the species in the boreal vegetation zone. Notes. Protoblastenia borealis is morphologically very variable and may dif- fer in the size and density of apothecia and in the thickness of the hypothecium. The sister species Protoblastenia ekmanii is morphologically similar, but ap- pears to differ in having a thicker hypothecium. The new species is also difficult to distinguish from the closely-related P rupestris. However, P rupestris does not have any strongly convex apothecia and has, on average, a thicker thallus, which is mostly areolate. Protoblastenia pseudocompressa often/usually has plane apothecia, but some specimens with convex apothecia are somewhat similar to P. borealis. The two Finnish specimens of Protoblastenia calvella re- semble P. borealis in morphology; they have strongly convex apothecia and a thick hypothecium which were also observed in a few P. borealis specimens. Due to these overlaps, ITS is needed for unambiguous identification of P borealis. Mycokeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 203 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Other sequenced specimens. FINLAND * Varsinais-Suomi, Karkkila, Haavis- to, Herneojankallio, pyroxene gneiss rock outcrop, on high SW-facing over- hanging wall, scarce, 80 m a.s.l., 60°29'N, 24°20'E, 18.5.1996, J. Pykala 16567 (H); * Lohja, Torhola, 300 m N-NE of Kallioranta, young pine plantation, S-slope, almost 1 m high rock wall of calcareous rock outcrop, rather scarce, 60 m a.s.l., 60°15'N, 23°52’E, 22.7.2004, J. Pykala 25292 (H); * Karjalohja, Karkali, 200 m SW Karkali farm, on a shore of Lohjanjarvi Lake, small cal- careous rock outcrop, on stone, 33 m a.s.|., 60°14'N, 23°49'E, 19.10.2004, J. Pykala 26097 (H); * Pohja, Kuovila, 300 m SW of Vihreamaki, abandoned lime quarry, NE-facing wall, 37 m a.s.l., 60°08'N, 23°24'E, 12.10.2006, J. Pykala 29704 (H); * Suomusjarvi, Sallittu, Huuttavanmaki, calciferous rock outcrop, on W-facing wall, 110 m a.s.I., 60°18'N, 23°37'E, 28.6.2008, J. Pykala 32757 (H); * Sarkisalo, Férby, E of Vahamaankaula, abandoned lime quarry, on N-facing wall, 7 m a.s.l., 23.7.2008, J. Pykala 33037 (H); » Sarkisalo, Férby, E of Vahamaankaula, abandoned lime quarry, on NW-facing wall, 7 m a.s.l, 23.7.2008, J. Pykala 33101 (H); * Kemidnsaari (Vastanfjard), Finsj6, Verk- viksudden, calcareous rock outcrop, on 1.2 m high E-facing wall, 5 m a.s.l., 60°00'N, 22°41'E, 13.7.2010, J. Pykala 38784 (H); * Salo (Suomusjarvi), Hint- tala, W of Kalattomansuo, Picea abies-dominated herb-rich forest, aban- doned lime quarry, on S-facing wall, 110 m a.s.|, 60°23'N, 23°39'E, 28.9.2010, J. Pykala 41355 (H); * Etela-Savo, Kerimaki, Ruokojarvi, Pitkaniemi, close by abandoned lime quarry, young Pinus sylvestris-forest, on calcareous boulder, 90 ma.s.l., 61°56'N, 29°00'E, 15.9.2011, J. Pykala 45759 (H); * Pohjois-Karja- la, Juankoski, Siikajarvi, Huosiaisniemi, Nature Reserve, mixed herb-rich for- est, dolomite rock outcrop, on W-facing wall, scarce, 100 m a.s.I., 63°12'N, 28°21'E, 25.7.2011, J. Pykala 42802 (H); * Juankoski, Siikajarvi, Lauantaijoki, Kalliola 250 m N, calciferous serpentine rock outcrop, on NW-facing wall, a y s 4 EP , - a eo i “ 3 —_— 7. en Figure 2. A. Protoblastenia arupii (ho (holotype). Scale bars: 0.5 mm. lotype); B. P. borealis (holotype); C. P compressa (holotype); D. P dolomitica MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 204 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland 132 ma.s.l., 63°09'N, 28°36'E, 9.9.2013, J. Pykala 47217 (H); * Keski-Pohjan- maa, Vimpeli, Saaksjarvi, Huosianmaankallio, lime quarry, quarry spoil, on calcareous boulder, rather scarce, 135 m a.s.|., 63°09'N, 24°04'E, 1.9.2010, J. Pykala 40695 (H); * Kainuu, Hyrynsalmi, Moisiovaara, Hyyrylainen SW, Pinus sylvestris-dominated heath forest, VMT site type, calciferous serpen- tine rock outcrop, on 1 m high W-facing wall, 190 m a.s.I., 64°32'N, 29°05 E, 30.5.2021, J. Pykala 57501 (H); * Koillismaa, Kuusamo, Oulanka National Park, Kiutako6ngas 400 mN, Pinus sylvestris - herb-rich forest, small dolomite rock outcrop, on SE-slope, rather scarce, 175 m a.s.l., 66°22'N, 29°19'E, 3.8.2010, J. Pykala 39151; * Kuusamo, Juuma, Oulanka National Park, Hautaniityn- vuoma, gorge, calciferous (dolomite) schistose rock outcrop, on SW-facing wall, 195 ma.s.l, 66°15'N, 29°27'E, 14.8.2010, J. Pykala 39942 (H); » Kuusa- mo, Juuma, Oulanka National Park, Hautaniitynvuoma, gorge, NE-slope, cal- ciferous (dolomite) schistose rock outcrop, on NE-facing wall, 179 m a.s.l., 66°15'N, 29°26'E, 21.8.2011, J. Pykala 44641 (H); « Salla, Oulanka National Park, Savilampi 1.4 km NE, bank of Savinajoki River, cliff, dolomite rock out- crop, on SE-facing wall, 185 m a.s.|., 66°26'N, 29°11'E, 23.8.2011, J. Pykala 44906 (H); * Kuusamo, Kallunki, Juhonlampi, young Pinus sylvestris-dominat- ed heath forest, calcareous rock outcrop, SW-slope, on 80 cm high SW-facing wall, 236 m a.s.l., 66°22'N, 29°03'E, 5.8.2021, J. Pykala 58196 (H); » Kuusa- mo, Vasarapera, Havukkalampi N, mixed herb-rich heath forest, calcareous rock outcrop, on 100 cm high SW-facing wall, 246 m a.s.|., 66°06'N, 28°44'E, 8.8.2021, J. Pykala 58392 (H); + Kittilan Lappi, Kittila, Tepsa, Herravaara SE, clear-cut heath forest, small calcareous rock outcrop, on 1.5 m high SE-fac- ing wall, scattered, 15.9.2022 J. Pykala & |. Kuusisto 60511 (H). Protoblastenia calvella Kainz & Rambold, Bibl. Lichenol. 88: 290 (2004) Description. Thallus white to whitish-grey, rimose to areolate, areoles 0.2- 0.5 mm, ca. 0.02-0.2 mm thick, K-, C-, UV-, algal cells 5-8 um. Apothecia or- ange-yellow to orange, 0.3-0.6 mm, strongly convex, superficial, not leaving pits to leaving shallow pits, K+violet; ca. 40-100 apothecia /cm?. Epihymenium dark orange-brown, 12-15 um thick, K+ violet. Hymenium 80-100 um thick. Hypothecium colourless to pale yellow, ca. 100-180 um thick. Paraphyses ca. 2-2.5 um thick, apex slightly thickened, often branching. Ascospores 0-sep- tate, (7.6-)9.4-10.7-12.0(-12.7) x (4.4—-)5.2-5.9-6.7(-7.8) um (n = 29). Habitat and distribution. Only two specimens are known from northern Fin- land (Pykala 2023). The habitats are sun-exposed calcareous and calciferous rock outcrops where the species grows on rock or on pebbles. The species is associated with Enchylium polycarpon (Hoffm.) Otdlora, P. M. J@rg. & Wedin and Rhizocarpon petraeum. Notes. In the ITS phylogeny, the P calvella specimens are divided into three strongly-supported lineages: the first one includes three specimens collect- ed in Austria, Germany and Sweden; the second group includes two Finnish specimens and the third group consists of specimens collected in Croatia and Germany (Fig. 1). More material is needed to determine whether P calvella rep- resents several species. Protoblastenia calvella is a cryptic species which can- not be identified without molecular data (Wirth et al. 2013). Older apothecia MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 205 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland are often brown-orange to dark red-brown (Svensson et al. 2024), but such co- louration was not observed in the Finnish specimens. It may be confused with several other Protoblastenia species. A few specimens of P borealis have a thick hypothecium and/or hymenium and thus resemble the Finnish specimens of P. calvella. The Finnish specimens of P rupestris have a thinner hymenium and hypothecium compared to P calvella. Furthermore, the apothecia tend to be predominantly orange, while in P. calvella, they are orange yellow. Protoblas- tenia ekmanii may differ in having darker orange apothecia. Protoblastenia fen- noarctica can be rather similar to P. calvella, but its apothecia vary from plane to strongly convex and its hymenium is thinner (ca. 50-80 um thick). Specimens examined. « Enonteki6n Lappi, Enontekio, Porojarvet, Toskalhar- ji, Toskalpahta, fell, dolomite rock outcrop, on steep SW-slope, 820 m a.s.l., 69°11'N, 21°30'E, 4.8.2011, J. Pykala 43503 (H); * Koillismmaa, Kuusamo, Juuma, Niskakoski, river bank, calciferous (dolomite) schistose rock outcrop, on peb- bles, 224 m a.s.l., 66°16'N, 29°24'E, 22.8.2011, J. Pykala 44807 (H). Protoblastenia compressa Pykala & Myllys, sp. nov. MycoBank No: 859625 Fig. 2C Diagnosis. Differs from P pseudocompressa in having plane to slightly convex apothecia. Type. FINLAND * Varsinais-Suomi, Parainen (Korppoo), Elfsj6, Hummelskar Island, close by shore of the Baltic Sea, calcareous rock outcrop, on shady 1 m high NE-facing wall, 5 m a.s.I., 60°08'N, 21°24'E, 31 July 2009, J. Pykala 35585 (H9250940 — holotype, GenBank accession number: PV766659). Description. Thallus white to pale grey, continuous, rimose to rarely areolate, areoles 0.2—0.8 mm, ca. 0.01-0.15 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells 4-9 um. Apothecia yellow, yellow-orange to orange, 0.2-0.8 mm, plane to slightly convex, 1/4—1-immersed in thallus, occasionally leaving shal- low pits, K+ violet; ca. 20-190 apothecia / cm?. Epihymenium orange-brown, 12-18 um thick, K+ violet. Hymenium 50-60 um thick. Hypothecium colourless to yellow, ca. 40-100 um thick. Paraphyses ca. 2-2.5 um thick, apex not thick- ened to slightly thickened, sparingly branched and anastomosing. Ascospores 0-septate, (8.6-)9.8-11.4-12.6(-13.5) x (4.8-)5.1-5.9-6.7(—7.4) um (n = 45). Habitat and distribution. This species has ca. 40 known localities in the hemi-boreal vegetation zone in SW Finland (seven localities confirmed by ITS). It grows on calcareous rocks and in lime quarries, often on N-facing walls. It seems to prefer shady habitats. In the SW archipelago of Finland, P compressa may be the most common Protoblastenia species. Typical companion species are Bagliettoa baldensis (A. Massal.) Vézda s.lat, Clauzadea monticola, Lepraria finkii (B. de Lesd.) R. C. Harris, Polyblastia abscondita (Nyl.) Arnold, Sagiolechia protuberans (Ach.) A. Massal., Thelidium decipiens (Nyl.) Kremp., T: incavatum Mudd, Verrucaria foveolata (Flérke) A. Massal., V. muralis, V. nigrescens s.lat. and V. viridula (Schrad.) Ach. Etymology. The name refers to the flattened apothecia of the species. Notes. The species is characterised by plane to slightly convex apothe- cia. Protoblastenia pseudocompressa is a sister species with a rather similar MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 206 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland morphology. However, P pseudocompressa often has convex apothecia which have not been seen in P compressa. The occurrence of Protoblastenia pseudo- compressa has been confirmed only in the Kuusamo-Salla area ca. 800 kilome- tres north from the distribution area of P compressa. Protoblastenia incrustans differs in having apothecia that leave pits (P compressa rarely leaves shallow pits). Protoblastenia coniasis differs in having a yellow thallus that gradually turns brownish-rose in the herbarium (Poelt 1957). Protoblastenia compressa may also be difficult to separate from P rupestris, even though they are not closely related. However, P compressa usually has thinner apothecia and a thinner thallus which is rarely areolate (mostly areolate in P rupestris). ITS may sometimes be needed for unambiguous identification of P compressa. Other sequenced specimens. FINLAND * Varsinais-Suomi, Kemidnsaari (Vastanfjard), S6dersundvik, Nasudden, calcareous rock outcrop on shore of the Baltic Sea, cliff, on N-facing wall, rather scarce, 12 m a.s.|., 60°03'N, 22°46'E, 25.8.2009, J. Pykala 36521 (H); * Parainen (Korppoo), Elfsj6, Stora Limskar Island, abandoned lime quarry, beneath N-facing wall, bottom, on pebbles, 4 m a.s.l., 60°09'N, 21°26'E, 22.6.2010, J. Pykala 38349 (H); * Parainen (Korppoo), Afvensar, Ronudden, abandoned lime quarry, on NW-facing wall, 2m a.s.|.,60°18'N, 21°32'E, 2.7.2010, J. Pykala 38646 (H); * Parainen (Korppoo), Afvensar, Ronudden, aban- doned lime quarry on shore of the Baltic Sea, on overhanging NW-facing wall, 2maz.s.l.,60°17'N, 21°32’E, 15.7.2010, J. Pykala 38965 (H); * Salo (Suomusjarvi), Rautsuo, 200 m NE of Kukutin, herb-rich forest, abandoned lime quarry, on NW-facing wall, 85 m a.s.I., 60°21'N, 23°37'E, 28.9.2010, J. Pykala 41376 (H); * Parainen, Skrabbole, Skrabbdle lime quarry, by the lime quarry, N-slope, on stone, 24 m a.s.l., 60°17'N, 22°17'E, 5.10.2010, J. Pykala 41511b (H); * Parainen, Attu, Skdtudden SW, abandoned lime quarry on shore of the Baltic Sea, on NE-facing wall, 2 maz.s.|., 60°11'N, 22°18'E, 16.6.2011, J. Pykala 42124 (H). Protoblastenia dolomitica Pykala & Myllys, sp. nov. MycoBank No: 859628 Fig. 2D Diagnosis. Differs from P siebenhaariana in having more densely occurring apothecia and from P arupii in having a rimose to areolate thallus and often olive-yellow apothecia. Type. Finland » Enontekion Lappi, Enonteki6, Porojarvet, Toskalharji, Toska- ljarvi N, fell, brook, W-shore, dolomite rock outcrop, on N-slope, 710 m a.s.l., 69°11'N, 21°26'E, 3 August 2011, J. Pykala 43466 (H9235018 — holotype, Gen- Bank accession number: PV766671). Description. Thallus white to whitish-grey, rarely pale brown, rimose to areolate, areoles 0.2—0.9 mm, ca. 0.03-0.25 mm thick, K-, C-, UV-, one specimen UV+ blu- ish-white, algal cells 4—8(-—12) um. Apothecia yellow to dirty orange-yellow, some- times olive-yellow, 0.15-0.7 mm, plane, slightly convex to convex, 1/2-immersed in thallus; ca. 40-160 apothecia / cm. Epihymenium dirty olive-yellow to or- ange-brown, 12-20 um thick, K+ violet. Hymenium 50-70 um thick. Hypothecium dark brown, reddish-brown or usually violet, ca. 40-100 um thick. Paraphyses ca. 1.5-2 um thick, apex not thickened to slightly thickened, branching. Ascospores 0-septate, (7.6-)11.0-12.8-14.7(-19.4) x (4.4-)4.8-5.4—-5.9(-7.4) um (n = 117). MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 207 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Habitat and distribution. This species has been found only in the parishes of Salla (in the biogeographical Province Ks) and Enontekid (biogeographical Province EnL) in N Finland. It grows on dolomite rock outcrops, once collected from dolomite stones and pebbles. The species may prefer sun-exposed and half-shady sites on fells and in gorges. Companion species include, for exam- ple, Acarospora glaucocarpa, Hymenelia rhodopis (Sommerf.) Lutzoni, Opegra- pha dolomitica (Arnold) Clauzade & Cl. Roux ex Torrente & Egea, Polyblastia spp., Protoblastenia fennoarctica, Synalissa ramulosa (Bernh.) K6rb., Thelidium auruntii (A. Massal.) Kremp. and T. declivum Pykala & Myllys. Etymology. The species grows on dolomite rocks. Notes. Protoblastenia dolomitica is closely related to P westbergii, which has slightly larger apothecia, broader spores and sometimes a thicker hypoth- ecium. The specimens have been previously identified as P. siebenhaariana in Finland. However, this species has only 1-10 apothecia / cm? (Kainz and Ram- bold 2004) and the Finnish specimens are not closely related to a GenBank specimen of P. siebenhaariana. Protoblastenia dolomitica is characterised by rather small, often dirty olive-yellow apothecia and a thin hypothecium. Proto- blastenia arupii, based on the inspection of only one known specimen, has or- ange apothecia, on average, broader spores and an only slightly thinner rimose thallus. More material from the morphologically related species is needed to determine whether P dolomitica can be identified by morphology only. Other specimens examined. FINLAND * Koillismaa, Salla, Oulanka National Park, Pikkuk6ngas, bank of River Oulankajoki, high cliff, calciferous (dolomite) schistose rock, on overhanging SW-facing wall, 180 m a.s.l., 66°25'N, 29°09'E, 10.8.2009, J. Pykala 36072 (H); * Salla, Oulanka National Park, Pikkukongas, bank of River Ou- lankajoki, high cliff, calciferous (dolomite) schistose rock outcrop, on SE-facing wall, 182 ma.s.l., 66°25'N, 29°08'E, 4.8.2010, J. Pykala 39217 (H); * Salla, Oulanka National Park, Pikkuk6ngas, bank of River Oulankajoki, high cliff, calciferous (do- lomite) schistose rock outcrop, beneath SW-facing wall, on pebble, 180 m a.s.l., 66°25N, 29°08'E, 13.8.2010, J. Pykala 39851 (H); * Enontekidn Lappi, Enontekio, Porojarvet, Toskalharji, Toskaljarvi N, fell, dolomite rock outcrop, gentle SE-slope, on dolomite pebbles, 730 m a.s.I., 69°12'N, 21°26'E, 2.8.2011, J. Pykala 43340 (H); * Enonteki6, Porojarvet, Toskalharji, Toskaljarvi N, lake shore, dolomite scree, on dolomite stones, 705 m a.s.|., 69°11'N, 21°26, 7.8.2011, J. Pykala 43855 (H); * Enonteki6, Kilpisjarvi, Saana, fell, steep NE-slope, dolomite rock outcrop, on NE-facing wall, 820 m a.s.I., 69°02'N, 20°51'E, 11.8.2011, J. Pykala 44115 (H). Protoblastenia ekmanii Pykala & Myllys, sp. nov. MycoBank No: 859630 Fig. 3A Diagnosis. Differs from P borealis in having a thicker hypothecium. Type. FINLAND « Varsinais-Suomi, Salo (Kisko), Kirkonkyla, 450 m NW of Kolko, shore of Maarjarvi Lake, calcareous rock outcrop, on W-slope, 43 ma.s.l., 60°12'N, 23°31'E, 20 May 2010, J. Pykala 37508 (H9224071 — holotype, Gen- Bank accession number: PV766674). Description. Thallus white to greyish-white, rimose to areolate, areoles 0.3-0.6 mm, ca. 0.05-0.2 mm thick, K-, UV+ bluish-white, algal cells 5-8 um. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 208 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Apothecia dirty brown-orange, 0.4-0.8 mm, convex to strongly convex, su- perficial; ca. 30-50 apothecia / cm?. Epihymenium dark orange-brown, 10- 20 um thick, K+ violet. Hymenium 50-70 um thick. Hypothecium yellow- ish-white, ca. 120-200 um thick, in some apothecia base patchily red-violet. Paraphyses ca. 2-2.5 um thick, apex not or slightly thickened. Ascospores 0-septate, (8.1—)8.7-10.2-11.6(-12.5) x (5.0—)5.1—-5.9-6.6(—7.2) (n = 12). Habitat and distribution. Only one specimen is known from a calcareous rock outcrop on a lake shore in SW Finland. The sun-exposed site suggests that the species may have a southern distribution. The species is apparently very rare in Finland. Companion species include Acarospora glaucocarpa, Clauzadea monticola, Placynthium nigrum (Huds.) Gray and Thelidium incavatum. Etymology. The name is in honour of Dr Stefan Ekman (Uppsala) for his many important contributions on the phylogeny and taxonomy of lichens in Fennoscandia and elsewhere. Notes. The species is closely related to P. borealis (only 2.5% difference in ITS), but seems to differ in having a thicker hypothecium. The hypothecium of P. borealis is very variable, but in no specimens does it reach the upper limit of the thickness in the P ekmanii specimen. The Finnish specimens of P calvella have orange-yellow to orange apothecia and thus resemble P ekmanii. More material is needed to determine whether the species can be unambiguously morphologically distinguished from P calvella. Protoblastenia fennoarctica Pykala & Myllys, sp. nov. MycoBank No: 859631 Fig. 3B Diagnosis. Differs from P calvella in having thinner hymenium, from P minuta in having rimose to areolate thallus and from P rikkinenii in usually having more densely occurring apothecia. Type. FINLAND * EnontekiOn Lappi, Enonteki6, Porojarvet, Toskalharji, 1.2 km NE of Toskaljarvi, fell, SW-slope, scree, on dolomite boulder, 875 ma.s.l., 69°12'N, 21°28'E, 5 August 2011, J. Pykala 43656 (H9220129 — holotype, GenBank accession number: PV766677). Description. Thallus white to pale greyish-brown, rimose to areolate, are- oles 0.15-1.0 mm, ca. 0.03-0.25 mm thick, K-, C-, UV-, algal cells 5-10 um. Apothecia yellow, orange-yellow to orange, 0.3-0.9 mm, plane to strongly convex, immersed in thallus to superficial; ca. 20-110 apothecia / cm. Epi- hymenium orange-yellow to brown, 12-20 um thick, K+ violet. Hymenium 50-80 um thick. Hypothecium colourless to yellow, base sometimes patch- ily reddish-brown, ca. 60-280 um thick. Paraphyses ca. 2-3 um thick, cells 5-8 um long, apex not thickened to slightly thickened, sparingly branched and anastomosing. Ascospores 0-septate, (6.6-)8.7-10.0-11.3(-12.5) x (4.2- )4.8-5.3-5.9(-7.2) um (n = 93). Habitat and distribution. This species has been found only in the Parish Enonteki6 (biogeographical Province EnL in NW Finland). It grows on fells on dolomite rock outcrops, boulders and pebbles. Companion species include Enchylium polycarpon, Farnoldia jurana, Physcia caesia (Hoffm.) Firnr., Poly- blastia spp., Protoblastenia dolomitica and Verrucaria vacillans Pykala & Myllys. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 209 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland (holotype). Scale bars: 0.5 mm. Etymology. The species occurs in the most northern, almost arctic (oroarc- tic) sites in Finland (Fennia). Notes. Protoblastenia fennoarctica morphologically resembles P. calvella, P minuta, P remota and P rikkinenii in having a pale hypothecium and small spores. In the ITS phylogeny, the species is most closely related to P. rikkinenii (Fig. 1). Protoblastenia rikkinenii differs in having more sparsely occurring apothecia, but more material is needed to confirm whether the two species can be identified by morphology. Protoblastenia calvella has a thicker hymenium (80-100 um thick). Protoblastenia minuta and P. remota have an endolithic to thin thallus and P minuta has smaller spores compared to P. fennoarctica, while P remota may have slightly larger spores. Other specimens examined. FINLAND * EnontekiOn Lappi, Enontekio, Porojarvet, Toskalharji, Toskalpahta fell, SW-slope, scree, on dolomite pebbles, 795 m a.s.l., 69°11'N, 21°29'E, 1.8.2011, J. Pykala 43105 (H); * Enonteki6, Porojarvet, Toskalharji, Toskaljarvi N, fell, dolomite scree, gentle SE-slope, on dolomite boulder, 715 m a.s.|., 69°11'N, 21°26'E, 2.8.2011, J. Pykala 43194 (H); * Enonteki6, Kilpisjarvi, Saana, fell, steep NE-slope, dolomite rock outcrop, on NE-facing wall, 820 ma.s.|., 69°02'N, 20°51'E, 11.8.2011, J. Pykala 44116 (H). Protoblastenia lilacina Poelt & Vézda, Cas. slezsk. Mus. Opavé, Ser. A 19: 26 (1970) Description. Thallus white to grey, endolithic, farinose to small areoles, K-, C-, UV- to UV+ bluish-white, algal cells 3-8 um. Apothecia orange to dark orange, 0.2-0.6 mm, slightly convex to strongly convex, 1/4-immersed to superficial, sometimes leaving shallow pits, K- or K+ fairly weakly red; ca. 40-100 apothecia MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 210 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland / cm?. Epihymenium (yellowish) orange-brown, 12-20 um thick, K- to K+ orange or carmine red, not into solution. Hymenium 50-120 um thick, with oil drop- lets. Hypothecium pale yellow, sometimes patchily violet, ca. 60-120 ym thick. Paraphyses ca. 2-3 um thick, apex not thickened to slightly thickened, spar- ingly branched and anastomosing. Ascospores 0-septate, (8.4—)10.7-12.4- 14.2(516:6) x45.2=)6:0-6 7-7 3(=7.8).pimin = 30), Habitat and distribution. The species is rare and occurs only on calcareous rock outcrops and boulders in the hemi-boreal zone in SW Finland. This seems to be the northern limit of this predominantly temperate species. Protoblaste- nia lilacina grows both on fairly sun-exposed and shady habitats. It may prefer rather dry calcareous rocks. Companion species include, for example, Bagliet- toa baldensis s.lat, B. calciseda (DC.) Gueidan & Cl. Roux, Catillaria lenticularis (Ach.) Th. Fr., Hymenelia spp., Lecidella stigmatea (Ach.) Hertel & Leuckerrt, Staurothele rupifraga (A. Massal.) Arnold, Verrucaria deversa Vain., V. foveolata and V. viridula. Notes. The species differs from most species of Protoblastenia in having K- or weakly K+ red apothecia. The closely-related Protoblastenia oulankaensis is the only other species with this characteristic. However, the two species have different distribution areas as Protoblastenia oulankaensis is only known from the Oulanka area in NE Finland. Morphologically, P oulankaensis differs in having a thicker hypothecium and often more sparsely occurring apothecia (in most specimens 10-30 apothecia / cm’). Specimens examined. FINLAND « Varsinais-Suomi, Kemi6nsaari (Vastanfjard), Vesterillo, Strommen 200-400 m SE, Pinus sylvestris-dominated forest, calcar- eous rock outcrop, on 70 cm high E-facing wall, 17 m a.s.l., 60°00'N, 22°44'E, 9.9.2009, J. Pykala 37012 (H); * Lohja, Moisio, between Lohjanharjuntie and Nummentie roads, NW-slope of Lohjanharju esker, Pinus sylvestris-heath forest, on calcareous boulder, 82 ma.s.I.,60°15'N, 24°06 E, 18.6.2005, J. Pykala 27009 H); * Lohja, Lohja, Pitkaniemi industrial area, 5m from shore of Lohjanjarvi Lake, deciduous forest, calcareous rock outcrop, on NW-facing 2 m high rock wall, 35 m a.s.l., 60°15'N, 24°02’E, 19.8.2005, J. Pykala 27629 (H); * Lohja, Lohja, Pitkaniemi industrial area, deciduous forest ca. 5 m from shore of Lohjanjar- vi Lake, calcareous rock outcrop, on 1.7 m high overhanging NW-facing wall, 35 ma.s.l.,60°15'N, 24°02'E, 12.7.2021, J. Pykala 57976 (H). Protoblastenia minuta Pykala & Myllys, sp. nov. MycoBank No: 859654 Fig. 3C Diagnosis. Differs from the other Protoblastenia species with a pale hypoth- ecium in having small apothecia and spores and apothecia which leave pits. Type. FINLAND * Enontekion Lappi, Enonteki6, Kilpisjarvi, Saana Nature Reserve, E-part, fell, steep SW-slope, dolomite rock outcrop, on S-facing wall, scarce, 715 m a.s.|, 69°02'N, 20°51'E, 12 August 2011, J. Pykala 44275 (H9250941 — holotype, GenBank accession number: PV766683). Description. Thallus ochraceous, endolithic, K-, UV-. Apothecia orange, 0.2-0.4 mm, slightly convex to strongly convex, immersed in rock to superfi- cial, leaving shallow to fairly deep pits; ca. 80 apothecia / cm?. Epihymenium MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 211 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland brown-yellow, 10-12 um thick, K+ violet. Hymenium 45-55 um thick. Hypothe- cium colourless to yellow, ca. 70-100 um thick. Paraphyses ca. 2-3 um thick, apex not thickened to slightly thickened. Ascospores 0-septate, (7.3—)7.6-7.9- 8.3(-8.8) x (4.1-)4.3-4.5-4.7(-4.8) um (n = 9). Habitat and distribution. Only one specimen has been found on the calcar- eous Saana fell in NE Finland, on a dolomite rock outcrop. Companion species include Acarospora macrospora (Hepp) A. Massal. ex Bagl., Enchylium polycar- pon, Placynthium asperellum (Ach.) Trevis. s.lat, Romjularia lurida (Ach.) Timdal and Thalloidima alutaceum Ani. Etymology. The name refers to the small size of the apothecia and spores. Notes. The species is rather isolated amongst the sequenced species of Protoblastenia (minimum difference from other species over 9% in ITS). The species has smaller apothecia and spores than most other Protoblastenia species with a pale hypothecium. Furthermore, other Finnish Protoblastenia species do not leave pits or only rarely leave shallow pits. However, only one specimen is known and the species may be expected to be morphologically more variable. More material is needed to evaluate whether the species can be identified by morphology only. Protoblastenia saanaensis has a rimose thallus and the apothecia do not leave pits. Protoblastenia szaferi differs in having larger apothecia and spores (see Hafellner (2006)). Protoblastenia oulankaensis Pykala & Myllys, sp. nov. MycoBank No: 859655 Fig. 3D Diagnosis. Differs from P lilacina in often having slightly larger and more sparse apothecia and an often thicker hypothecium. Type. FINLAND °* Koillismaa, Salla, Oulanka National Park, W of Savikoski, cliff, dolomite rock outcrop, on NE-facing wall, 190 m a.s.l.,66°25'N, 29°10'E, 17 August 2010, J. Pykala 40201 (H9224130 — holotype, UPS — isotype, GenBank accession number: PV766686). Description. Thallus white to whitish-grey, endolithic, farinose or rimose, ca. 0-0.1 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells 5-9 um. Apothecia orange to dark orange, 0.3-0.9 mm, convex to strongly convex, 1/4-immersed to superficial, often leaving shallow pits, K+ weakly red; ca. 10-50 apothe- cia / cm? (in one specimen, ca. 50-80 apothecia / cm’). Epihymenium dark orange-brown, 12-25 um thick, K- to K+ orange or carmine red. Hymenium 70-120 um thick, with oil droplets, in one specimen, partly violet in few apoth- ecia. Hypothecium colourless to yellow, more rarely patchily red-brown, ca. 60-220 um thick. Paraphyses ca. 2-3 um thick, apex not thickened to slightly thickened, sparingly branched and anastomosing. Ascospores 0-septate, (8.8- )11.8-13.9-16.0(-18.8) x (5.2-)6.6-7.4-8.2(—9.6) um (n = 92). Habitat and distribution. The species occurs only in the Oulanka area in the parishes of Kuusamo and Salla in NE Finland. It grows on N- and S-fac- ing walls of dolomite rock outcrops. Companion species include, for ex- ample, Diploschistes gypsaceus (Ach.) Zahlbr., Gyalecta jenensis (Batsch) Zahlbr., Lempholemma isidioides (Nyl. ex Arnold) H. Magn., Protoblastenia borealis, Sagiolechia protuberans, Scytinium parvum, Verrucaria foveolata and MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 212 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Xanthocarpia crenulatella (Nyl.) Frodén, Arup & S@chting s.lat. Based on the ITS phylogeny, one GenBank specimen of P lilacina from northern Italy (AY425629) belongs to P. oulankaensis (Fig. 1). Etymology. In Finland, the species occurs only in the Oulanka area in the northeast. Notes. Protoblastenia oulankaensis is closely related to P lilacina, but differs clearly from that species in its ITS profile (10% difference). It is also morphologi- cally rather similar to P lilacina, having a weak K+ reaction of apothecia and large spores. Protoblastenia oulankaensis tends to have slightly larger apothecia, which usually occur more sparsely (in most specimens ca. 10-50 apothecia /cm?) and, on average, a thicker hypothecium. Protoblastenia lilacina is only known from the hemi-boreal zone of SW Finland, ca. 800 kilometres south to south-west. Other specimens examined. FINLAND ° Koillismaa, Salla, Oulanka National Park, Savikoski 300 m N, Pinus sylvestris-forest, steep N-slope, dolomite rock outcrop, on NW-facing wall, st pc — sp, 180 ma.s.l.,66°25'N, 29°10'E, 10.8.2010, J. Pykala 39629 (H); + Salla, Oulanka National Park, Savilamminniemi, shore of Savilampi Lake, cliff, dolomite rock outcrop, on E-facing wall, sp, 180 m a.s.l., 66°25'N, 29°10'E, 12.8.2010, J. Pykala 39810 (H); * Salla, Oulanka National Park, Savilampi 1.4 km NE, bank of Savinajoki river, cliff, dolomite rock outcrop, upper slope, on NE-facing wall, 192 m a.s.l., 66°26'N, 29°11'E, 23.8.2011, J. Pykala 44928 (H); Salla, Hautajarvi, Kurtinniittykuru, dolomite rock outcrop, on SW-facing wall, 195 m a.s.l., 66°26'N, 29°09'E, 24.8.2011, J. Pykala 44949 (H)); * Salla, Hautajarvi, Kurtinniittykuru, dolomite rock outcrop, on SE-facing wall, 195 ma.s.l., 66°26'N, 29°09'E, 24.8.2011, J. Pykala 44953 (H); * Kuusamo, Juu- ma, Myllyniemi, dolomite rock outcrop, on NE-facing wall, 235 m a.s.|., 66°16'N, 29°22'E, 11.8.2020, J. Pykala 55995 (H). Protoblastenia pseudocompressa Pykala & Myllys, sp. nov. MycoBank No: 859656 Fig. 4A Diagnosis. Differs from P compressa in often having convex apothecia. Type. FINLAND * Koillismaa, Salla, Onkamo, Latvarova SW, clear-cut heath forest, on calcareous boulder, 258 m a.s.I., 66°42'N, 28°055'E, 9 August 2021, J. Pykala 58457 (H9250942 - holotype, UPS - isotype, GenBank accession number: PV766696). Description. Thallus white to grey, continuous, rimose (usually) to areolate, rarely endolithic, ca. 0-0.15 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells 6-12 um. Apothecia orange-yellow to orange, 0.2—0.7 mm, plane to strongly con- vex, 1/4-1-immersed in thallus or rock to superficial, sometimes leaving shallow pits; ca. 20-120 apothecia / cm2. Epihymenium yellow-brown to orange-brown, 12-18 um thick, K+ violet. Hymenium 50-60 um thick. Hypothecium colourless to pale yellow, rarely pink (in one specimen), ca. 50-120 um thick. Paraphyses ca. 1.5-2.5 um thick, sparsely branched and anastomosing. Ascospores 0-sep- tate, (6.8—-)9.1-10.6-12.1(-13.0) x (4.2—)4.7—5.5-6.2(—8.0) um (n = 51). Habitat and distribution. Protoblastenia pseudocompressa has been verified by ITS only from the Oulanka area (parishes Kuusamo and Salla) in NE Finland. However, a morphologically similar specimen was found in the neighbouring MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 213 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland & = a. Su a . . 4 Figure 4. A. Protoblastenia pseudocompressa (holotype); B. P pseudoterricola (holotype); C. RP remota (holotype); D. P. rikkinenii (holotype). Scale bars: 0.5 mm. biogeographical Province of Kainuu. The species cannot be identified with certainty by morphology only and it is likely that more populations occur in the Oulanka area. It mainly grows on dolomite rock outcrops (one specimen was collected from a dolomite boulder) and may prefer half-shady habitats. Typical companion species are Acarospora glaucocarpa, Clauzadea monticola, Farnoldia jurana, Hymenelia prevostii (Duby) Kremp., H. rhodopis, Lathagrium fuscovirens, Placynthium nigrum, Thelidium declivum, Verrucaria deversa, V. mu- ralis, V. kuusamoensis Pykala, Kantelinen & Myllys and V. subtilis. Etymology. The name refers to the close relationship in ITS and morpholog- ical similarity to P compressa. Notes. Protoblastenia pseudocompressa is closely related to P compressa and the two species are also morphologically similar. Protoblastenia pseudo- compressa has, on average, slightly smaller spores than P compressa, but there is a wide overlap. Furthermore, P pseudocompressa specimens with plane to slightly convex apothecia may be confused with P compressa. However, the apothecia of P pseudocompressa are often convex, while the apothecia of P compressa are always plane to slightly convex. Based on the distribution of sequenced specimens, the two species are geographically well separated. Protoblastenia pseudocompressa specimens with convex apothecia may also be confused with P borealis, which, however, often has a thicker hymenium. Other specimens examined. FINLAND * Koillismaa, Kuusamo, Oulanka Na- tional Park, Pikkukonkaankuru, dolomite rock outcrop, small W-facing wall, 175 ma.s.l., 66°21'N, 29°19'E, 8.8.2009, J. Pykala 35901 (H); * Salla, Oulan- ka National Park, 400 m N of Savilampi, bank of River Savinajoki, dolomite rock outcrop, on NW-facing wall, 178 m a.s.|., 66°25'N, 29°10'E, 13.8.2009, J. Pykala 36325 (H); * Kuusamo, Oulanka National Park, Kiutak6ngas 400 m N, SW-slope, Pinus sylvestris-dominated forest, small dolomite rock outcrop, MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 214 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland on SW-facing wall, scarce, 175 m a.s.I., 66°22'N, 29°19'E, 6.8.2010, J. Pykala 39367 (H); * Kuusamo, Oulanka National Park, Kiutak6ngas, bank of Oulan- kajoki River, by the rapids, calciferous (dolomite) schistose rock outcrop, on rather flat rock, 150 m a.s.l., 66°22'N, 29°20'E, 10.8.2010, J. Pykala 39693 (H); * Kuusamo, Oulanka, Putaanoja, 500 m W-NW of Hautala, dolomite rock outcrop, on E-facing wall, scarce, 232 m a.s.|., 66°22'N, 29°25'E, 15.8.2010, J. Pykala 39987 (H). Protoblastenia pseudoterricola Pykala & Myllys, sp. nov. MycoBank No: 859657 Fig. 4B Diagnosis. Differs from P terricola in having a thinner thallus and hypothecium. Type. FINLAND * Enonteki6n Lappi, Enonteki6, Kilpisjarvi, Saana, fell, steep NE-slope, dolomite rock outcrop, beneath NE-facing wall, on dolomite pebbles, 820 m a.s.l., 69°02'N, 20°51'E, 11 August 2011, J. Pykala 44157 (H9235068 — holotype, GenBank accession number: PV766697). Description. Thallus white, rimose, ca. 0.05-0.1 mm thick, K-, UV+ blu- ish-white. Apothecia orange (young apothecia) to dirty orange-brown, 0.3-0.8 mm, convex to strongly convex, superficial; ca. 30 apothecia / cm’. Epihymenium orange-brown, 10-15 um thick, K+ violet. Hymenium 70 um thick. Hypothecium dark reddish-brown to violet, ca. 60-100 um thick. Ascospores 0-septate, (7.9-)8.6-9.9-11.2(-12.4) x (4.5-)4.8-5.5-6.2(—7.0) um (n = 20). Habitat and distribution. Only one specimen of this species is known. It was collected from dolomite pebbles beneath an NE-facing wall of a dolomite rock outcrop on the Saana fell (NW Finland). Companion species include Farnoldia jurana, Polyblastia inconspicua Savic & Tibell and Thelidium huuskonenii Pykala & Myllys. Etymology. The name refers to the similarities in morphology and ITS to P terricola. Notes. The species is closely related to P terricola and P violacea. Protoblas- tenia terricola has rimose to areolate thallus (when growing on rock) mainly ex- ceeding 0.1 mm in thickness and an often thicker hypothecium. Protoblastenia violacea has orange apothecia and smaller spores. More material is needed to confirm whether these three species can be distinguished by morphology only. Protoblastenia westbergii resembles the new species, but has more densely occurring apothecia (ca. 40-160 apothecia / cm?) and larger spores. Protoblastenia remota Pykala & Myllys, sp. nov. MycoBank No: 859658 Fig. 4C Diagnosis. Differs from P borealis and RP pseudocompressa by often smaller more convex apothecia and a smaller thallus. Type. FINLAND * Varsinais-Suomi, Pernio, Lupaja, Alhonmaki, abandoned lime quarry, on W-slope, 9 August 1991, J. Pykala 8547 (H9250943 — holotype, GenBank accession number: PV766698). MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 215 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Description. Thallus white to grey, endolithic to sparse small areoles, 0.2- 0.5 mm wide, ca. 0-0.15 mm thick, K-, C-, UV- to UV+ bluish-white. Apothecia orange, 0.2-0.6 mm, convex to strongly convex, 1/2-immersed to mainly su- perficial; ca. 15-80 apothecia / cm?. Epihymenium yellow-brown to dark or- ange, 12-20 um thick, K+ violet. Hymenium 50-80 um thick. Hypothecium co- lourless to pale yellow, ca. 100-200 um thick. Paraphyses ca. 2-2.5 um thick, apex not thickened to thickened to ca. 3-5 um thick. Ascospores 0-septate, (8.4-)9.8-11.4-12.9(-14.4) x (4.8-)5.0-5.6-6.1(-7.1) um (n = 36). Habitat and distribution. Two specimens of this species are known: one from a lime quarry in SW Fnland and one from a calciferous schistose cliff in NE Finland. Companion species include Lathagrium fuscovirens, Mycobilimbia tetramera (De Not.) Vitik., Ahti, Kuusinen, Lommi & T. Ulvinen ex Hafellner & Turk and Verrucaria deversa. Furthermore, two closely-related specimens from North America and previously identified as Protoblastenia aff. rupestris (MZ922194, MZ922195) (Munger et al. 2022) may belong to P. remota (Fig. 1). Etymology. The name refers to the large difference in ITS of the species com- pared to other Protoblastenia species (differs by ca. 15-18% from other species). Notes. Based on ITS sequences, this species is not closely related to any sequenced Protoblastenia species. The species is morphologically a typical Protoblastenia and very difficult to distinguish from several species, particularly from P. borealis and P pseudocompressa. However, these two species have, on average, less convex apothecia, a thinner hypothecium and often a better de- veloped thallus, but there is a wide overlap. In addition, Protoblastenia borealis often has larger apothecia (up to 1 mm). Other specimens examined. FINLAND * Koillismaa, Kuusamo, Oulanka Na- tional Park, Paahkanankallio, high cliff, calciferous (dolomite) schistose rock outcrop, on SE-facing wall, st pc, 195 m a.s.l., 66°16'N, 29°31'E, 7.8.2010, J. Pykala 39455 (H). Protoblastenia rikkinenii Pykala & Myllys, sp. nov. MycoBank No: 859659 Fig. 4D Diagnosis. Differs from P. fennoarctica in always having convex to strongly con- vex apothecia and from P. saanaensis and P timdalii in having sparsely occur- ring apothecia and a thick hypothecium. Type. FINLAND * Enontekion Lappi, Enontekid, Porojarvet, Toskalharji, Toska- ljarvi N, fell, dolomite rock outcrop, on dolomite pebbles, 735 m a.s.I., 69°12'N, 21°26'E, 2 August 2011, J. Pykala 43252 (H9250944 — holotype, GenBank ac- cession number: PV766700). Description. Thallus white, grey to pale brown, rimose to areolate, ca. 0.05-0.2 mm thick, K-, C-, UV-, algal cells 5-9 um. Apothecia yellow-orange, 0.3-0.8 mm, convex to strongly convex, 1/2-immersed to superficial; ca. 20-40 apothecia /cm?. Epihymenium dirty yellow to orange-yellow, 12- 18 um thick, K+ violet. Hymenium 50-70 um thick. Hypothecium pale yellow, ca. 80-300 um thick. Paraphyses ca. 2—2.5 um thick, apex slightly thickened to 3 um thick. Ascospores 0-septate, (7.4-)8.5-9.6-10.7(-12.6) x (4.4-)4.7- 5.1-5.5(-6.3) um (n = 52). MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 216 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Habitat and distribution. The species occurs on dolomite rock outcrops, do- lomite stone and dolomite pebbles on the fells. It may prefer sun-exposed sites. Companion species include Halecania alpivaga (Th. Fr.) M. Mayrhofer, Hymene- lia heteromorpha (Kremp.) Lutzoni, H. prevostii, H. rhodopis, Lecidea polycocca Sommerf., Polyblastia sp., Rhizocarpon petraeum, Thelidium huuskonenii and Verrucaria lapponica Pykala. One GenBank sequence (KY266981) originates from northern Norway. Etymology. The name is in honour of Prof Jouko Rikkinen (Helsinki) for his innovative work on various fields of lichenology. Notes. The species is related to P. fennoarctica, which often has more dense- ly occurring, often plane and/or yellow apothecia. However, an ITS sequence may be needed for unambiguous identification of specimens. Protoblastenia saanaensis and P timdalii have a thinner hypothecium and more densely occur- ring apothecia. Protoblastenia calvella has more densely occurring apothecia and slightly larger spores. Other specimens examined. FINLAND * Enonteki6n Lappi, Enonteki6, Kilpis- jarvi, Saana Nature Reserve, W-part, fell, dolomite rock outcrop, on W-facing wall, 700 m a.s.I., 69°03'N, 20°48'E, 9.8.2011, J. Pykala 43910 (H); « Enonteki6, Kilpisjarvi, Saana Nature Reserve, fell, steep SW-slope, dolomite rock outcrop, beneath SW-facing wall, on dolomite stone, 840 m a.s.l., 69°02'N, 20°50'E, 14.8.2011, J. Pykala 44387 (H). Protoblastenia rupestris (Scop.) J. Steiner, Verh. Kaiserl.-K6nigl. zool.-bot. Ges. Wien 61: 47 (1911) Lichen rupestris Scop., Fl. carniol., Edn 2 (Wien) 2: 363 (1772). Description. Thallus whitish-grey to rarely pale brown, rimose to predom- inantly areolate, areoles 0.1-0.8 mm, ca. 0.1-0.2 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells 4-12 um. Apothecia yellow-orange to or- ange, 0.2-0.9 mm, slightly convex to convex, 3/4-immersed in thallus to superficial; ca. 40-100 apothecia / cm?. Epihymenium orange-brown, 12- 20 um thick, K+ violet. Hymenium 50-80 um thick. Hypothecium pale yellow, ca. 60-80 um thick. Paraphyses ca. 1.5-2.5 um thick, apex not thickened to slightly thickened to 2-3 um thick, sparingly branched and anastomos- ing. Ascospores 0-septate, (8.2—)9.1-10.3-11.6(-13.0) x (5.2—-)5.6-6.3- 7.1(-7.6) um (n = 43). Habitat and distribution. Sequenced specimens were collected from rather sun-exposed calcareous rock outcrops in the hemi-boreal zone in Finland. The species may prefer calcareous rock outcrops close to shores. Notes. Protoblastenia rupestris sequences available in GenBank are hetero- geneous and distributed in three different groups in our ITS phylogeny (Fig. 1). Specimens similar to those in Kainz and Rambold (2004) are likely to belong to P rupestris s. stricto as the species was described from Central Europe. The se- quence MZ159571 (UK) is related to P. incrustans and the sequence KY266981 (Norway) belongs in P. rikkinenii. Morphologically, the Finnish specimens of P rupestris may be confused with the closely-related P. borealis. However, the P rupestris specimens have a predominantly areolate thallus, while the thalli MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 217 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland of P borealis are more often rimose and usually do not exceed 0.1 mm in thickness. Furthermore, P. borealis often has a thicker hypothecium. Despite these minor differences, the two species often cannot be distinguished with certainty by morphology only. Specimens examined. FINLAND * Varsinais-Suomi, Lohja, Skraatila, Kirkko- vuori E, cliff, calcareous rock outcrop, on S-slope, 8.8.1990, J. Pykala 5963 (H); * Lohja, Lohja, Pitkaniemi industrial area, by the sauna, on shore of Lohjanjarvi Lake, flat calcareous rock outcrop, 33 m a.s.|., 60°15'N, 24°02'E, 19.8.2005, J. Pykala 27569 (H); * Karjalohja, Saarenpaa, Saarenpdanniemi, calcareous rock outcrop on shore of Lohjanjarvi Lake, on small SW-facing wall, 35 m a.s.l., 60°13'N, 23°47'E, 7.10.2008, J. Pykala 34043 (H); * Salo (Sarkisalo), Kaukosalo, Klintinmaki, cliff, calciferous schistose rock outcrop, on SW-facing wall, st pc, 35 ma.s.l., 60°06'N, 22°58'E, 4.10.2011, J. Pykala 46413 (H). Protoblastenia saanaensis Pykala & Myllys, sp. nov. MycoBank No: 859660 Fig. 5A Diagnosis. Differs from P minuta in apothecia not leaving pits, from P rikkinenii in having more densely occurring apothecia, from P. fennoarctica in having a thinner hypothecium and from P. timdalii in often having smaller apothecia. Type. FINLAND * Enontekion Lappi, Enonteki6, Kilpisjarvi, Saana Nature Reserve, fell, steep SW-slope, dolomite rock outcrop, SW-slope, on dolomite pebbles, 835 m a.s.|., 69°02'N, 20°50'E, 14 August 2011, J. Pykala 44431 (H9250945 — holotype, GenBank accession number: PV766707). Description. Thallus white, grey or pale brown, rimose to areolate, areoles 0.3-0.8 mm, ca. 0.03-0.1 mm thick, K-, C-, UV-, algal cells 5-12 tum. Apoth- ecia yellow to orange-yellow, 0.3-0.5 mm, slightly convex to strongly con- vex, 3/4-immersed to superficial; ca. 20-80 apothecia / cm*. Epihymenium orange-yellow, 12-17 ym thick, K+ violet. Hymenium 50-70 um thick. Hy- pothecium pale yellow (some orange-yellow in squash), ca. 40-100 um thick. Paraphyses ca. 2-2.5 um thick, apex not thickened to slightly thickened, branching. Ascospores 0-septate, (5.4—)7.1-8.4-9.7(-11.4) x (3.8-)4.3- 4.8-5.4(-6.1) um (n = 47). Habitat and distribution. The species has only been found on the calcare- ous Saana fell in NW Finland. The specimens are from a dolomite stone and dolomite pebbles in sun-exposed to rather shady habitats. Companion species include Acarospora glaucocarpa. Etymology. The name refers to the Saana fell where both known specimens were collected. Notes. The species is most closely related to P. timdalii (ca. 5-6% difference in ITS) from which it is difficult to distinguish by morphology. It may have slight- ly smaller apothecia and spores. It also resembles P minuta and the closely-re- lated P rikkinenii in having similar small spores. The only known specimen of P minuta has apothecia which leave shallow to deep pits. Protoblastenia rikki- nenii has sparsely occurring apothecia. Protoblastenia fennoarctica resembles P. saanaensis, but usually has a thicker hypothecium, slightly larger spores and sometimes plane apothecia. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 218 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland 4 * an - <= Figure 5. A. Protoblastenia saanaensis (holotype); B. P. timdalii (holotype); C. PR. violacea (holotype (holotype). Scale bars: 0.5 mm. ); D. PR. westbergii Other specimens examined. FINLAND * Enonteki6n Lappi, Enonteki6, Kilpis- jarvi, Saana Nature Reserve, fell, steep SW-slope, dolomite rock outcrop, under overhanging SW-facing wall, on dolomite stone, 835 m a.s.l., 69°02'N, 20°50'E, 14.8.2011, J. Pykala 44438 (H). Protoblastenia terricola (Anzi) Lynge, Rep. Sci. Res. Norweg. Exped. Novaya Zemlya, 1921 43: 216 (1928) Biatora rupestris var. terricola Anzi, Cat. Lich. Sondr.: 78 (1860). Description. Thallus white to whitish-grey, rimose to areolate, areoles 0.2- 0.8 mm, ca. 0.05-0.2 mm thick, K-, C-, UV+ bluish-white. Apothecia dirty or- ange to brown-orange (some with an olive tinge), 0.4—0.9 mm, strongly con- vex, superficial, K+ violet; ca. 70-120 apothecia /cm?. Epihymenium dark orange-brown, 8-15 um thick, K+ violet. Hymenium 60-100 um thick. Hypoth- ecium violet to violet-brown, 70-180 um thick. Paraphyses ca. 2—2.5 um thick, apex not thickened to slightly thickened, sparingly branched and anastomos- ing. Ascospores O-septate, (9.4-)9.7-10.8-11.9(-12.4) x (4.7-)4.9-5.5- 6.2(—7.3) um (n = 16). Habitat and distribution. The species occurs only in northern Finland (bio- geographical Provinces Ks and EnL) with one separate population in the bio- geographical Province PK (not sequenced) in eastern Finland. It grows on cal- careous rock outcrops and on calcareous soil. Notes. The specimens on calcareous soil (not sequenced) have a squa- mulose thallus, thus differing from the other species of Protoblastenia (Ka- inz and Rambold 2004). However, on dolomite rock outcrops, the thallus MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 219 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland is more weakly developed, rimose to areolate. The apothecia of P terrico- la tend to be dirty orange to brown. Protoblastenia terricola may be most difficult to distinguish from P. pseudoterricola. The only known specimen of P pseudoterricola has a 60-100 um thick hypothecium and a 0.05- 0.1 mm thick thallus. Specimen sequenced. FINLAND * Koillismaa, Salla, Oulanka National Park, Savilampi 1.2 km NE, bank of Savinajoki River, dolomite rock outcrop, on SE-fac- ing wall, 66°26'N, 29°11'E, 23.8.2011, J. Pykala 44876 (H). Protoblastenia timdalii Pykala & Myllys, sp. nov. MycoBank No: 859661 Fig. 5B Diagnosis. Differs from P saanaensis in having slightly larger apothecia and spores. Type. FINLAND * Enonteki6n Lappi, Enonteki6, Kilpisjarvi, Saana, fell, steep NE-slope, dolomite rock outcrop, on NE-facing wall, 820 m a.s.l., 69°02'N, 20°51'E, 11 August 2011, J. Pykala 44177 (H9250946 — holotype, O — isotype, GenBank accession number: PV766711). Description. Thallus white, grey or pale brown, endolithic, rimose to areolate, areoles 0.1—0.4 mm, ca. 0-0.15 mm thick, K-, C-, UV-. Apothecia yellow-orange, 0.3-0.8 mm, convex to strongly convex, semi-immersed to superficial; ca. 40-60 apothecia / cm?. Epihymenium brown-yellow to dirty yellow, K+ violet. Hymenium 60-70 um thick, pale, partly yellow. Hypothecium colourless to yellow, ca. 60-100 um thick. Paraphyses ca. 2-2.5 um thick, apex not thick- ened to slightly thickened. Ascospores 0-septate, (6.8—)7.9-9.2—10.5(—12.6) x (3.8-)4.2—4.8-5.5(-—7.0) um (n = 46). Habitat and distribution. Two specimens are known from the calcareous fells of Enonteki6, one from a NE-facing wall of a dolomite rock outcrop and one from dolomite stones on Dryas octopetala heath. This suggests that the species may have considerable microhabitat variation on the fells. Companion species include Halecania alpivaga, Polyblastia spp. and Thelidium auruntii. Etymology. The name is in honour of Prof Einar Timdal (Oslo) for his numer- ous important contributions to the taxonomy, biogeography and conservation of Fennoscandian lichens, including studies on the related genus Psora and an important technical edition of the Nordic lichen journal Graphis Scripta. Notes. The species is most closely related to P pseudoincrustans ined. This undescribed species differs in having plane to convex apothecia, a thick hymenium (80-115 um thick) and larger spores (mean 12 x 7 um) (Kainz and Rambold 2004). However, in our phylogeny, P pseudoincrustans is divided into two species. The closely-related P saanaensis (differing ca. 5% in ITS) is also morphologically very similar to P. timdalii. Protoblastenia saanaensis may have slightly smaller apothecia and spores, but more material is needed to confirm this. Other specimens examined. FINLAND * Enonteki6n Lappi, Enontekio, Porojar- vet, Toskalharji, 1.2 km NE of Toskaljarvi, fell, SW-slope, gentle E-slope, Dryas heath, on dolomite stones, 875 m a.s.|., 69°12'N, 21°28'E, 5.8.2011 J. Pykala 43624 (H). MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 220 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Protoblastenia violacea Pykala & Myllys, sp. nov. MycoBank No: 859662 Fig. 5C Diagnosis. Differs from most Protoblastenia species with violet hypothecium in having a thick hypothecium and from P. westbergii in having less densely occurring apothecia and smaller spores. Type. FINLAND * Enontekion Lappi, Enontekid, Porojarvet, Toskalharji, Toska- lpahta, fell, dolomite rock outcrop, beneath SW-facing wall, on dolomite peb- bles, 800 ma.s.l.,69°11'N, 21°30'E, 4 August 2011, J. Pykala 43524 (H9250947 — holotype, GenBank accession number: PV766713). Description. Thallus white, grey or pale brown, rimose to areolate, areoles 0.15-0.4 mm, ca. 0.01-0.15 mm thick, K-, C-, UV-, algal cells 6-10 um. Apothe- cia orange, 0.3-1.0 mm, convex to mainly strongly convex, superficial, high; ca. 20-40 apothecia / cm’, sometimes forming clusters of smaller apothecia on surface of old apothecia. Epihymenium orange-brown, 12-14 um thick, K+ vio- let. Hymenium 60-80 um thick. Hypothecium purple, orange-red to red-violet, ca. 100-280 um thick. Paraphyses ca. 2—2.5(—3) um thick, apex not thickened to slightly thickened, often branching. Ascospores O0-septate, (6.3—)7.9-9.2- 10,.5(—12.7)} x (3.7-)4,2-4.6=5.1(-5,6) pmid(n = 52), Habitat and distribution. The three specimens are known from the calcare- ous Toskalharji fell (NW Finland), all from dolomite pebbles. Companion spe- cies include Farnoldia jurana and Lecidella stigmatea. Etymology. The name refers to the violet hypothecium of the species. Notes. The species is related to P pseudoterricola and P terricola, which have mainly brown apothecia and broader spores. Protoblastenia violacea differs from P. siebenhaariana in having more densely occurring apothecia (only ca. 1-10 apothecia / cm? in P. siebenhaariana; Kainz and Rambold (2004)), slightly smaller apothecia and a thinner hymenium. Protoblastenia westbergii has more densely occurring apothecia and larger spores. Other specimens examined. FINLAND * Enonteki6n Lappi, Enontekio, Porojar- vet, Toskalharji, Toskalpahta, fell, SW-slope, on dolomite pebbles, 765 m a.s.l., 69°11'N, 21°29'E, 1.8.2011, J. Pykala 43015 (H); * Enontekid, Porojarvet, To- skalharji, 1.2 km NE of Toskaljarvi, fell, SW-slope, gentle E-slope, Dryas heath, on dolomite pebbles, 875 m a.s.I.,69°11'N, 21°30'E, 5.8.2011, J. Pykala 43605 (H). Protoblastenia westbergii Pykala & Myllys, sp. nov. MycoBank No: 859663 Fig. 5D Diagnosis. Differs from P dolomitica in having slightly broader spores, darker orange apothecia and a thicker hypothecium. Type. FINLAND * Enontekion Lappi, Enontekio, Kilpisjarvi, Saana Nature Re- serve, E-part, fell, steep SW-slope, on dolomite boulder, 910 m a.s.I., 69°02'N, 20°51'E, 13 August 2011, J. Pykala 44316 (H9229821 — holotype, GenBank ac- cession number: PV766718). Description. Thallus white to whitish-grey, rimose to areolate, areoles 0.15- 0.8 mm, ca. 0.05-0.3 mm thick, K-, C-, UV- to UV+ bluish-white, algal cells MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 221 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland 4-8 um, thallus often strongly invaded by cyanobacteria. Apothecia orange to dark brownish-orange, 0.2—1.1 mm, slightly convex to convex, 3/4-immersed to superficial, in one specimen with pale margin ca. 30-50 um thick; ca. 40-200 apothecia / cm?. Epihymenium dirty yellow to orange-brown, 10-15 um thick, K+ violet. Hymenium 40-70 um thick, often with oil droplets. Hypothecium red- brown to red-violet, ca. 60-150 um thick. Paraphyses ca. 2—2.5 um thick, apex not thickened to slightly thickened to 3 um thick, some branched and anas- tomosing. Ascospores 0-septate, (8.4—)10.5-12.0-13.6(-15.0) x (5.0-)5.4- 6.1-6.9(-7.8) um (n = 39). Habitat and distribution. Four specimens are known from the calcareous Saa- na and Toskalharji fells in NW Finland. On Saana, both specimens occurred close to each other on a steep SW-slope on a dolomite boulder. On Toskalharji, the species was found on a SE-facing slope on dolomite pebbles. The species may prefer dry, sun-exposed habitats. Companion species include Farnoldia jurana, Parmeliella thriptophylla (Ach.) Mull. Arg., Thelidium auruntii and Verrucaria sp. Etymology. The name is in honour of Dr Martin Westberg (Uppsala) for his im- portant work on the taxonomy and biogeography of northern European lichens. Notes. The species is closely related to P dolomitica (4% difference in ITS). Protoblastenia dolomitica has, on average, smaller and paler, sometimes ol- ive-yellow apothecia, narrower spores and a thinner hypothecium. However, P westbergii often also has a thin hypothecium. Protoblastenia arupii has slight- ly broader spores, a thin hypothecium and possibly thinner thallus. Other specimens examined. FINLAND * Enontekion Lappi, Enonteki6, Poro- jarvet, Toskalharji, Toskaljarvi N, fell, dolomite scree, gentle SE-slope, on do- lomite pebbles, 715 m a.s.l., 69°11'N, 21°26'E, 2.8.2011, J. Pykala 43196 (H); * Enonteki6, Porojarvet, Toskalharji, Toskaljarvi N, fell, dolomite rock outcrop, SE-slope, on dolomite pebbles, 720 m a.s.l., 69°12'N, 21°26E, 2.8.2011, J. Pykala 43279 (H); * Enontekio, Kilpisjarvi, Saana Nature Reserve, E-part, fell, steep SW-slope, on dolomite boulder, 910 m a.s.I., 69°02'N, 20°51'E, 13.8.2011, J. Pykala 44314 (H). A preliminary key of Protoblastenia in Finland 1 Apothecia K-06 KAsweakhy red sii iba dara coevondha es inode suvnsbattonnddauesdes estes 2 = ADOUNeCld K-AWIOIER Mh uot 5) decet er bs men eunr ts ried oan t ha SONS Tm Sonne ld 3 2 (1) Apothecia 0.2—-0.6 mm, 40-100 apothecia / cm?, hemiboreal....P. lilacina = Apothecia 0.3-0.9 mm, 10-50(-80) apothecia / cm?, northern boreal ee RT ree eC ro ok PR eet Kir ey ore FEN P. oulankaensis Bi(2)* KAY DOTe Cla WIOle tO DFOW MPa .vecatensee. Sos terrerauee Manish eden avantctdeanelenrsaveeee: 4 = Hy POtMe Cla Ww iRILG Toy GLOW .25. horas cant rate eas ore cass ar cement doreasaent ee tes hear 9 AW O\y Oa. LOSAO a pOlNeClay CM .csicte sa clcaa seh he cage Ne eit SALE RAD Tole esr sbha Savas 5 a Cara 160 rapotneClay Chika ke. tag ec treet rt ea eet 6 5(4) — Apothecia mainly brown, hypothecia 60-100 um thick... A Ae Ro Ripe POR ARTS 2 ee eT ah DO MOTT AES SORE TR RT P. pseudoterricola = Apothecia orange, hypothecia 100-280 um thick................ P. violacea 6(4) — Thallus slightly rimOSe........... ee cecceecceeeceeeceecseeeeceeeseeseeseeeeseeseens P. arupii = TMMAULS FIMO SEMOsarSOlALS «oc dhcwenscisesSeeleadence tat dierppeievennnee sites pesanenenens 7 7 (6) — Apothecia yellow, dirty orange yellow or olive yellow........ P. dolomitica e Apothecia orange to Drown OFainge ..............:cccceesseceeesseceesseeeeeeseeeeessaeees 8 MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 222 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland 8 (7) Apothecia convex to strongly convex, mean spore size 10.0 x LINE Ub femaeenrir ont MCS er ef elh at acid ead le cei eter te OU caer P. terricola Apothecia slightly convex to convex, mean spore size 12.6 x BESANT A iantcenaM tectenesnuine Mertens ere cat RRO deetnreus tebe ae ded P. westbergii 9(3) Apothecia plane to slightly CONVEX ...........ccececcesseesceteetsceseeseessenseees 10 = Apothecia slightly convex to strongly CONVEX............::cccceesceeessseeeeeees 11 GVO) WH SMMED ORS xc .1 i ccc cosiceMenccsanen tvvesiaattonehastavestnettapesreesveoks P. compressa = NOGIMER MN DORCAS ta Sicko. 4 ousde ms haere he veloc en acters P. pseudocompressa HEIRS; MOAN OW INOS cists Secs aR sel once dled esate oh Maw tM oka ML red ast iee ae? 12 = OMeGal eae Ouse Saket pe OM oy ec apn Sl aeava rs er gir snia te etoasererboinedioes 17 12 (11) Apothecia convex to strongly convex, hypothecia 100-200 um thick.. _exnanr Hae acs te. ean oh AEM ah Ins 5: be nt sa ae Peas Anika 7 Peale Are Wp 13 = Apothecia slightly convex to convex, hypothecia 40-160 um thick.....15 13 (12) Thallus endolithic to sparse small areoles...............ccceceeeee P. remota = TANS FIMO SETOzareO lates a, 2.8.8. a vhssaves Aca naedtatasenpecleadeoere idee eicaeness 14 14 (13) Apothecia orange-yellow to orange, 40-100 apothecia / cm’............... Pern OeAal Ars dustMetescaeas Read taalte A Atecalt wes re Miah anol Mec agthses does iad P. calvella Apothecia dirty brown-orange, 30-50 apothecia / cm”........ P. ekmanii 15 (12) Thallus mainly areolate, 0.1-0.2 mm thick................cce P rupestris ? Thallus mainly rimose, usually 0.02-0.1 mm thick............... eee 16 16 (15) Apothecia plane to strongly convex, hymenia 50-60 um thick............. TO oh TORR Jet» ARTE ROO Is MN UPS yh der OORT P. pseudocompressa 5 Apothecia slightly convex to strongly convex, hymenia 50-90 um PEDIC KOREN, 95525! directs bcc AD, sae 6 faces AGRO doar SERN Bo EN, SRA Yi P. borealis 17(11) Apothecia leave shallow to fairly deep pits, thallus endolithic.....P minuta = Apothecia not leave pits, rarely leave shallow pits, thallus usually (s101 [Ni] Lo aMMee ae. Lavi Sasa ee ea ae OMT sce ee OE ee ee oe 18 TBCl7)« HY pothecia 60=30Gshini thicken 2c nce tome ee los ar 19 = hypothecia 40=TO0sim, (nicks, nese sews. oc ee eee Aa 20 19 (18) Apothecia convex to strongly convex, 20-40 apothecia / cm?............. icenaaeaenneaae aaNet ta dil a) eaten sezagnl eer aecan senao7 OO aaren: erzaeaeata aeRaRT naa P. rikkinenii - Apothecia plane to strongly convex, 20-110 apothecia / cm7’.............. Seaiettaiseeeatidacrdaaiet seus dt i aevtett ee neeatibared daaubrrsscaal Adee Jeane etaaa Aare P. fennoarctica 20 (18) Apothecia 0.3-0.5 mm, mean spore size 8.4 x 4.8 um......P. saanaensis 7 Apothecia 0.3-0.8 mm, mean spore size 9.2 x 4.8 um.......... P. timdalii Acknowledgements We would like to thank Marijke Iso-Kokkila, Emelie Winquist and Diana Weck- man for their help with the laboratory work and Teuvo Ahti, Starri Heidmarsson and Einar Timdal for their constructive comments on the manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 293 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Use of Al No use of Al was reported. Funding The fieldwork and manuscript preparation were mainly conducted as parts of the research projects “Threatened lichens of calcareous rocks” (Grant number YTB059), “Key habitats for red-listed lichens” (Grant number 700T-MTX003) and “Inventory of calcareous and ser- pentine rocks” financed by the Finnish Ministry of the Environment. The first two projects were part of the Research Programme of Deficiently Known and Threatened Forest Spe- cies (PUTTE). DNA barcoding was supported by the Kone Foundation, the Finnish Cultural Foundation and the Academy of Finland through their grants to the Finnish Barcode of Life (FinBOL) and the Finnish Biodiversity Information Facility (FinBIF). In the project “Key habitats for red-listed lichens”, 32 specimens of Protoblastenia were barcoded. Author contributions All authors have contributed equally. Author ORCIDs Juha Pykala © https://orcid.org/0000-0002-7566-9310 Leena Myllys © https://orcid.org/0000-0002-9566-9473 Data availability All of the data that support the findings of this study are available in the main text. References Cannon P Aptroot A, Coppins B, Orange A, Sanderson N, Simkin J (2022) Lecanorales: Psoraceae, including the genera Brianaria, Protoblastenia, Protomicarea and Psora. Revisions of British and Irish Lichens 28: 1-11. Castello M, Nimis PL (1995) A critical revision of Antarctic lichens described by C. W. Dodge. Bibliotheca Lichenologica 57: 71-92. Divakar PK, Leavitt SD, Molina MC, Del-Prado R, Lumbsch HT, Crespo A (2016) A DNA barcoding approach for identification of hidden diversity in Parmeliaceae (Ascomycota): Parmelia sensu stricto as a case study. Botanical Journal of the Linnean Society 180: 21-29. https://doi.org/10.1111/boj.12358 Edgar RC (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792-1797. https://doi.org/10.1093/nar/ gkh340 Ekman S, Blaalid R (2011) The devil in the details: Interactions between the branch- length prior and likelihood model affect node support and branch lengths in the phy- logeny of the Psoraceae. Systematic Botany 60: 541-561. https://doi.org/10.1093/ sysbio/syr022 Evankow AM, Yin A, Zulfiqar R, Anmad UF, Nordenhaug P, Khalid AN, Wang L, Timdal E (2025) Psora mediterranea (Lecanorales, Psoraceae), a new lichen species from Europe, including a new concept for P. himalayana and a revised key to the European species. Mycological Progress 24: 26. https://doi.org/10.1007/s11557-025-02045-8 Gardes M, Bruns TD (1993) ITS primers with enhanced specificity for basidiomycetes —- application to the identification of mycorrhizae and rusts. Molecular Ecology 2: 113-118. https://doi.org/10.1111/j.1365-294X.1993.tb00005.x MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 904 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Hafellner J (2006) Protoblastenia szaferi (lichenized Ascomycotina) — new to the Alps. Herzogia 19: 23-33. Kainz C (2004) Protoblastenia. In: Nash Ill TH, Ryan BD, Diederich P, Gries C, Bungartz F (Eds) Lichen Flora of the Greater Sonoran Desert Region. Volume Il. Lichens Unlimited, Arizona State University, Tempe, 424-425. Kainz C, Rambold G (2004) A phylogenetic study of the lichen genus Protoblastenia (Lecanorales, Psoraceae) in Central Europe. Bibliotheca Lichenologica 88: 267-299. Lucking R, Aime MC, Robbertse B, Miller AN, Ariyawansa HA, Aoki T, Cardinali G, Crous PW, Druzhinina IS, Geiser DM, Hawksworth DL, Hyde KD, Irinyi L, Jeewon R, Johnston PR, Kirk PM, Malosso E, May TW, Meyer W, Opik M, Robert V, Stadler M, Thines M, Vu D, Yurkov ANM, Zhang N, Schoch CL (2020a) Unambiguous identification of fungi: Where do we stand and how accurate and precise is fungal DNA barcoding? IMA Fungus 11: 1-32. https://doi.org/10.1186/s43008-020-00033-z Licking R, Nadel MRA, Araujo E, Gerlach A (2020b) Two decades of DNA barcoding in the genus Usnea (Parmeliaceae): How useful and reliable is the ITS? Plant and Fungal Systematics 65: 303-357. https://doi.org/10.35535/pfsyst-2020-0025 Miadlikowska J, Kauff H, H6gnabba F, Oliver JC, Molnar K, Fraker E, Gaya E, Hafellner J, Hofstetter V, Gueidan C, Otalora MAG, Hodkinson B, Kukwa M, Lucking R, Bjork C, Sipman HJM, Burgaz AR, Thell A, Passo A, Myllys L, Goward T, Fernandez-Brime S, Hestmark G, Lendemer J, Lumbsch HT, Schmull M, Schoch CL, Sérusiaux E, Maddi- son DR, Arnold AE, Lutzoni F, Stenroos S (2014) A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infra- generic taxa, 317 genera and 66 families. Molecular Phylogenetics and Evolution 79: 132-168. https://doi.org/10.1016/j.ympev.2014.04.003 Munger |, Baugh M, Henrie JR, Hollinger J, Crepeau R, Leavitt SD (2022) Integra- tive biodiversity inventories: Characterizing lichen-forming fungal diversity in Glen Canyon National Recreation Area using DNA barcoding and vouchered specimens. Western North American Naturalist 82(2): 213-233. https://doi. org/10.3398/064.082.0201 Myllys L, Velmala S, Holien H, Halonen P Wang LS, Goward T (2011) Phylogeny of the ge- nus Bryoria. Lichenologist (London, England) 45: 617-638. https://doi.org/10.1017/ $0024282911000132 Nowak J (1974) Protoblastenia szaferi sp. nov., a new lichen species in the calcareous part of the Polish Tatra Mts. Fragmenta Floristica et Geobotanica 20: 529-533. Orange A (2012) Semi-cryptic marine species of Hydropunctaria (Verrucariaceae, liche- nized Ascomycota) from north-west Europe. Lichenologist (London, England) 44: 299-320. https://doi.org/10.1017/S002428291 1000867 Poelt J (1957) Mitteleuropadische Flechten. V. Mitteilungen der Botanischen Staatssam- mlung Munchen 17-18: 386-399. Printzen C, Holien H, Kantelinen A, Myllys L, Ratschow F, Stepanchikova |, Weber L, Timdal E (2023) DNA barcoding indicates the presence of unrecognized species and phylogenetic diversity within the Biatora vernalis-and B. meiocarpa-groups. Plant and Fungal Systematics 68: 262-279. https://doi.org/10.35535/pfsyst-2023-0011 Pykala J (2010) Notes on the lichen flora of Saana and Malla fells in northern Finland. Memoranda Societatis pro Fauna et Flora Fennica 86: 34-42. Pykala J (2023) Additions to the lichen flora of Finland. X. Graphis Scripta 35(3): 14-29. Pykala J, Lommi S (2021) Lichen flora of Finland — short history of Finnish lichenology and updated species statistics. Memoranda Societatis pro Fauna et Flora Fennica 97: 73-88. MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 295 Juha Pykala & Leena Myllys: Lichen genus Protoblastenia in Finland Pykala J, Launis A, Myllys L (2017) Four new species of Verrucaria from calcareous rocks in Finland Lichenologist 49: 27-37. https://doi.org/10.1017/S002428291 6000542 Pykala J, Kantelinen A, Myllys L (2020) Taxonomy of Verrucaria species characterised by large spores, perithecia leaving pits in the rock and a pale thin thallus in Finland. MycoKeys 72: 43-92. https://doi.org/10.3897/mycokeys.72.56223 Stamatakis A (2014) RAxML version 8: A tool for phylogenetic analysis and post-analy- sis of large phylogenies. Bioinformatics (Oxford, England) 30(9): 1312-1313. https:// doi.org/10.1093/bioinformatics/btu033 Stenroos S, Velmala S, Pykala J, Ahti T [Eds] (2016) Lichens of Finland. Norrlinia 30: 1-896. Svensson M, Ekman §S, Arup U, Eide Ekman L, Hammarstrom O, Isaksson R, Jonsson F, Palice Z, Vicente R, Westberg M (2024) Further additions to the Swedish flora of lichenised fungi. Graphis Scripta 36(2): 15-49. Timdal E (1987) Problems of generic delimitation among squamiform members of the Lecideaceae. In: Peveling E (Ed.) Progress and problems in lichenology in the eight- ies. Bibliotheca Lichenologica 25: 243-247. White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal DNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR Protocols: a guide to methods and applications. Academic Press, San Diego, 315-322. https://doi.org/10.1016/B978-0-12-3721 80-8.50042-1 Wirth V, Hauck M, Schultz M (2013) Die Flechten Deutschlands. Band 2. Ulmer, Stuttgart, 1244 pp. Zahlbruckner A (1936) Neue Flechten. XII. Annales Mycologici 34: 159-179. Zhang Y, Clancy J, Jensen J, McMullin RT, Wang L, Leavitt SD (2022) Providing scale to a known taxonomic unknown—At least a 70-fold increase in species diversity in a cosmopolitan nominal taxon of lichen-forming fungi. Journal of Fungi (Basel, Swit- zerland) 8: 490. https://doi.org/10.3390/jof8050490 MycoKeys 124: 193-226 (2025), DOI: 10.3897/mycokeys.124.162802 226