Two new species of Cora (lichenized Basidiomycota, Lichenomphaliaceae) and additional records from Bolivia

1 Two new species of Cora (lichenized Basidiomycota, Lichenomphaliaceae) and additional records from Bolivia Magdalena Oset1, Adam Flakus2, Beata Guzow-Krzemińska1 1 DepartmentofPlantTaxonomyandNatureConservation,UniversityofGdańsk,WitaStwosza59,PL-80-308Gdańsk,Poland 2 W.SzaferInstituteofBotany,PolishAcademyofSciences,Lubicz46,PL-31-512Krakow,Poland Correspondingauthor:MagdalenaOset([email protected]) Copyright: © Magdalena Oset et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Two new species of lichens, Cora neoparabovei Oset, Flakus & Guzow-Krzem. and C. neopseudobovei Oset, Flakus & Guzow-Krzem., are described based on material collected in Bolivia. The study is based on morphological and anatomical examinations, molecular phylogenetic analysis and haplotype network analysis of the nuITS rDNA sequences. Phenotypically, Cora neoparabovei is characterised by the long, concentrically arranged setae on the surface of the thallus and densely pilose margins, whereas in C. neopseudobovei the diagnostic features are the brown upper surface thallus when fresh, pale yellow to orange-yellow when dry, with slightly visible concentric colour zonation when dry and also with involute, creamy white margins. In addition, three species, i.e., Cora arcabucana B. Moncada, C. Rodr. & Lücking, C. cuzcoensis Holgado, Rivas Plata & Perlmutter, and C. undulata L.Y. Vargas, B. Moncada & Lücking, are reported as new to Bolivia. New records of C. aspera Wilk, Lücking & E. Morales from Bolivia are reported. Key words: Basidiolichens, cyanolichens, molecular barcoding, neotropics, new species Introduction According to Lücking et al. (2014c) lichenized Basidiomycota make up less than one percent of all lichenized fungi in terms of species number. However, in ecosystems and habitats such as tropical alpine regions and conserved forests, lichenized Basidiomycota play an important role as a component of tropical forests (see also Petersen 1967; Oberwinkler 1970, 1984, 2012; Gómez 1972; Parmasto 1978; Lutzoni and Vilgalys 1995; Redhead et al. 2002; Lawrey et al. 2007, 2009; Ertz et al. 2008; Yánez et al. 2012). Within Basidiomycota, the family Lichenomphaliaceae (Lücking & Redhead) Vizzini, Consiglio & P. Alvarado is represented by eight genera: Acantholichen P.M. Jørg., Arrhenia Fr., Cora Fr., Corella Vain., Cyphellostereum D.A. Reid, Dictyonema C. Agardh ex Kunth, Eonema Redhead, Lücking & Lawrey, and Lichenomphalia Redhead, Lutzoni, Moncalvo & Vilgalys (Vizzini et al. 2024). The most diverse genus within Lichenomphaliaceae is Cora, with over 100 species formally described using a combination of morphology, anatomy, habitat ecology, and nuITS rDNA barcoding, but many still await formal recognition as 450 species are predicted to exist (Lücking et al. 2013, 2014b, 2014c, 2014d, 2015, 2017, 2020; Dal Forno et al. 2019). Academic editor: Gerhard Rambold Received: 16 July 2025 Accepted: 4 November 2025 Published: 2 December 2025 Citation: Oset M, Flakus A, GuzowKrzemińska B (2025) Two new species of Cora (lichenized Basidiomycota, Lichenomphaliaceae) and additional records from Bolivia. MycoKeys 126: 1–18. https://doi.org/10.3897/ mycokeys.126.165395 MycoKeys 126: 1–18 (2025) DOI: 10.3897/mycokeys.126.165395 2 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia Cora is characterised by foliose to macrosquamulose thalli forming distinct layers (cortex, photobiont layer, medulla) and producing corticoid basidiocarps on the lobe underside (see Dal Forno et al. 2013). In many lichen groups taxonomy can be based on morphological, anatomical or chemical characters. However, in the case of the genus Cora, traditional revisions based largely on herbarium collections failed to recognise important field characters (e.g. lobe arrangement, colour, and substrate or hymenophore anatomy) to establish species which led to numerous misclassifications of taxa. These traits are not widely used for diagnostic purposes due to the excessive plasticity of the thallus and changes in morphology resulting, from among other things, habitat conditions or variations in these traits due to changes in thallus hydration. This makes species within this genus much more difficult to distinguish. According to Dal Forno et al. (2022), it was the main reason why historically only a single foliose species Cora glabrata (Spreng.) Fr. (previously named Dictyonema pavonium (Sw.) Parmasto and subsequently D. glabratum (Spreng.) D. Hawksw. was recognised. According to Dal Forno et al. (2022) the best method for species identification in the genus Cora is DNA barcoding using nuITS rDNA. In the genus Cora it has been generally found that the topology of phylogeny based on ITS is highly consistent with the topology of other markers, such as nuLSU and RPB2, suggesting that ITS accurately distinguishes these lineages (Dal Forno et al. 2022). Therefore, for Cora taxonomy, DNA barcoding is the best and most efficient research method, which has already significantly increased the number of taxa described. This conclusion is also supported by this study. Despite the intensification of research on the genus Cora in Neotropical areas (Dal Forno et al. 2019, 2022; Lücking et al.2013, 2014b, 2014c, 2014d, 2015, 2017, 2020), there are still areas where this genus has not been thoroughly investigated. One such country is Bolivia, which is one of the two landlocked South American countries located in the central part of the continent. Bolivia is the continent’s highest country in terms of average altitude, with more than half of the country’s area made up of highlands and mountains. Also, the country is characterized by tropical and, in many places, dry climates. According to Ibisch and Mérida (2004) and Josse et al. (2003), twelve ecoregions and twenty-three sub-regions are present in Bolivia. All together these make it very diverse in terms of biodiversity and the country is ranked among the ones with the highest biodiversity in the world (e.g. Ibisch and Mérida 2004; Josse et al. 2003; Rodriguez-Flakus et al. 2016). Biodiversity studies in Bolivia remain rather poorly explored, and lichens are no exception. The late development of lichenological research in Bolivia meant that by 1998, only 150 lichen species had been recorded. As a result of surveys conducted in subsequent years, this number has increased to about 2,000 species. This suggests that only about half of Bolivia’s lichen biota has been documented so far, and many additional new taxa are undoubtedly yet to be discovered there (Rodriguez-Flakus et al. 2016 and literature cited therein). To date, 18 species of the genus Cora have been recorded from Bolivia (e.g. Flakus and Lücking 2008; Lücking et al. 2013, 2014c, 2017; Ertz et al. 2015; Flakus et al. 2016; Rodriguez-Flakus et al. 2016 and lit. cited therein; Guzow-Krzemińska et al. 2019; Oset et al. 2024). Recently, modern approaches in Cora taxonomy, including phylogenetic analyses, have been applied to Bolivian collections, resulting in the recording of additional species, including taxa new to science. 3 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia This paper is a further contribution and presents the descriptions and records of five additional Cora species, including two new to science (C. neoparabovei and C. neopseudobovei), three new to Bolivia (C. arcabucana B. Moncada, C. Rodr. & Lücking, C. cuzcoensis Holgado, Rivas Plata & Perlmutter, and C. undulata L.Y. Vargas, B. Moncada & Lücking), and new records of C. aspera Wilk, Lücking & E. Morales. Materials and methods Taxon sampling Cora specimens were collected in the Yungas and Tucumano-Boliviano regions during fieldwork carried out between 2010 and 2014. The specimens are deposited at KRAM, LPB, and UGDA herbaria. Morphology and anatomy were examined using Nikon SMZ800N dissecting microscope and ZEISS Axioskop compound microscope. Character assessment was based on the morphological and anatomical traits from Cora described by Lücking et al. (2013, 2014a, 2017) and Vargas et al. (2014). Secondary compounds were analysed using thin-layer chromatography (TLC) in solvents A and C (Orange et al. 2001). DNA extraction, PCR amplification and sequencing A total of 18 new specimens of Cora from Bolivia were used for the molecular study. The new nuITS rDNA sequences were compared with Cora sequences available in the GenBank database using BLAST search (Altschul et al. 1997). The nuITS rDNA sequences of representative taxa belonging to the genus Cora, based on the analyses of Dal-Forno et al. (2022) and Oset et al. (2024), were aligned with new sequences. Corella tomentosa Vain. (KJ780617) and C. zahlbruckneri Schiffn. (KJ780592) were used as an outgroup (Suppl. material 1). For new DNA extractions from Bolivian material, two separate thallus fragments were taken from each specimen to allow cross-checking of the results and account for potential sample contamination. Total genomic DNA was extracted using the Plant & Fungi DNA Purification Kit (Eurx, Poland), following the manufacturer’s protocol and modified CTAB method (Guzow-Krzemińska and Węgrzyn 2000). Fungal nuITS rDNA was amplified using the primers ITS5 and ITS4 (White et al. 1990). The same primers were used for sequencing. For PCR amplification, Start-Warm HS-PCR Mix (A&A Biotechnology, Poland) was used with the following parameters: an initial denaturation at 94 °C for 3 min and 33 cycles of denaturation at 94 °C for 30 sec; annealing at 52° for 45 sec; and extension at 72 °C for 1 min followed by a final extension at 72 °C for 10 min. The PCR products were purified with the Clean-Up Kit (A&A Biotechnology, Poland) according to the manufacturer’s instructions. Sequencing was performed in Macrogen (the Netherlands) (http://www.macrogen.com). Sequence editing and alignments Forward and reverse sequences of the nuITS rDNA region were assembled using BIOEDIT v. 7.25. (Hall 1999). The assembled sequences were analysed using BLAST search (Altschul et al. 1997) at NCBI (https://www.ncbi.nlm.nih.gov). Sequences 4 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia used for the phylogenetic analyses were selected based on Dal-Forno et al. (2022) and Oset et al. (2024) and are presented in Suppl. material 1 together with GenBank accession numbers, voucher numbers, country of origin, and source publication. A single sequence was used for most species, with multiple sequences used for species that are newly recorded. The final alignment of representatives of Cora spp. was generated in MAFFT using auto option and default parameters (Kuraku et al. 2013; Katoh et al. 2019). The alignment was trimmed using MEGA-11 (Tamura et al. 2021). The final alignment consisted of 109 sequences and 840 sites. A second alignment of newly sequenced samples together with sequences of reference taxa obtained from GenBank was made in Seaview software (Galtier et al. 1996; Gouy et al. 2010) employing the muscle option. The alignment included only the closely related species: C. bovei, C. palaeotropica, C. parabovei, C. pseudobovei, C. neoparabovei, C. neopseudobovei, Cora sp., and consisted of 20 sequences and 752 sites. Network and phylogenetic analyses IQ-TREE analysis was performed to find the best-fitting nucleotide substitution model (Nguyen et al. 2015) with the model selection restricted to models implemented in MrBayes. TIMe+I+G4 was chosen based on BIC. The search for maximum likelihood tree with 100 bootstrap replicates was performed using IQ-TREE (Nguyen et al. 2015) on the CIPRES Web Portal (Miller et al. 2010). Bayesian analysis was carried out using a Markov Chain Monte Carlo (MCMC) method, in MrBayes v. 3.2.6 (Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003) on the CIPRES Web Portal (Miller et al. 2010) using a previously selected model. Two parallel MCMC runs were performed, each using four independent chains and ten million generations, sampling every 1000th tree. The resulting log files were analysed using Tracer 1.7.2 (Rambaut et al. 2018). Posterior probabilities (PP) were determined by calculating a majority-rule consensus tree after discarding the initial 25% trees of each chain as the burn-in. The convergence of the chains was confirmed by the convergent diagnostic of the Potential Scale Reduction Factor (PSRF), which approached 1, and the ‘average standard deviation of split frequencies’ was < 0.01 (Ronquist et al. 2005). Phylogenetic trees were visualised using FigTree v. 1.4.3 (Rambaut 2009) and modified in Inkscape (https://inkscape.org/). Bootstrap support (BS values ≥ 70) and PP values (values ≥ 0.95) are given near the branches on the phylogenetic tree. PopART software (https://popart.maths.otago.ac.nz) employing the TCS network option was used for haplotype network analyses (Clement et al. 2002). This analysis is supplemented with a table showing variable positions in the alignment of nuITS rDNA marker of Cora bovei, C. neoparabovei, C. neopseudobovei, C. palaeotropica, C. parabovei, C. pseudobovei and Cora sp. (Suppl. material 2) as PopArt masks sites with gaps and missing data. Results and discussion Eighteen new nuITS rDNA sequences were generated for this study and used in phylogenetic analysis together with reference sequences representing different species of Cora (Fig. 1). Nine of these were found to represent previously 5 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia 0.04 Corella zahlbruckneri KJ780592 Cora fimbriata KJ780537 Cora neopseudobovei PV647978 Cora setosa KJ780535 Cora minutula KJ780381 Cora schizophylloides KX772719 Cora asperaPV647986 Cora viliewoa EU825956 Cora dalfornoae PP356557 Cora comaltepeca KX772556 Cora barbifera KX772682 Cora leslactuca KJ780507 Cora applanata KJ780473 Cora arcabucana KX772585 Cora crispoleslia KJ780443 Cora aspera KF443231 Cora neopseudobovei PV647977 Cora guajalitensis KF443239 Cora arborescens KJ780599 Cora hawksworthiana KJ780600 Cora strigosa KF443241 Cora urceolata KJ780677 Cora maxima KJ802420 Cora davicrinita KJ780440 Cora caraana KX772512 Cora caliginosa KJ802414 Cora barbulata EU825961 Cora casasolana KX772540 Cora paraciferrii KJ780428 Cora parabovei KJ780571 Cora aspera PV647987 Cora aspera PV647989 Cora smaragdina EU825960 Cora inversa KF443237 Cora garagoa KX772578 Cora dalehana KJ780434 Cora corani KJ780369 Cora hymenocarpa EU825959 Cora fuscodavidiana KJ780411 Cora sp. KJ780522 Cora arachnoidea KF443233 Cora suturifera KJ780399 Cora paraminor KJ780610 Cora quillacinga KX772701 Cora neoparabovei PV647979 Cora bovei KJ780579 Cora elephas KJ780448 Cora pikynasa KX772511 Cora udebeceana KJ780520 Cora soredavidia KJ780649 Cora dulcis KX772551 Cora casanarensis KJ780530 Cora davidia KX772658 Cora neopseudobovei PV647976 Cora gomeziana KJ780598 Cora santacruzensis KJ780565 Cora subdavicrinita KJ780441 Cora haledana KJ780607 Cora undulata KJ780536 Cora palaeotropica KX772742 Cora pseudobovei KJ780372 Cora pastorum KX772660 Cora caucensis KX772716 Cora reticulifera KJ780655 Cora hafecesweorthensis KJ780519 Cora canari KJ780394 Cora ciferrii KF443238 Cora verjonensis KX772457 Cora pichinchensis KJ780391 Cora campestris KJ780578 Cora dewisanti KJ780405 Cora accipiter KJ780433 Cora minor EU825968 Cora gigantea KJ780515 Cora cyphellifera KF443242 Cora celestinoa KX772596 Cora hirsuta KF443235 Cora undulata PV647990 Cora arcabucana PV647985 Cora rothesiorum KJ780445 Cora aturucoa KX772607 Cora corelleslia KJ780498 Cora imi EU825958 Cora neoparabovei PV647980 Cora pseudocorani KJ780368 Cora hochesuordensis KJ780367 Cora arcabucana PV647984 Cora terrestris KJ780596 Cora aspera PV647988 Cora auriculeslia KJ780378 Cora neopseudobovei PV647974 Cora byssoidea KF443234 Cora neopseudobovei PV647975 Cora cuzcoensis KJ802411 Cora squamiformis KF443240 Cora arachnodavidea KJ780477 Cora itabaiana KX772629 Cora boleslia KJ802416 Cora palustris KJ780597 Cora neoparabovei PV647981 Cora aspera PV647991 Cora davibogotana KJ780504 Cora rubrosanguinea KJ780400 Cora putumayensis KX772611 Cora cuzcoensis PV647983 Cora neoparabovei PV647982 Cora terricoleslia KJ802419 Corella tomentosa KJ780617 Cora yukiboa KX772744 85/1 92/1 87/1 92/1 100/1 100/1 73/1 100/1 100/1 85/- 93/0.99 96/1 71/- 77/0.99 99/1 95/1 96/1 91/1 100/1 77/1 92/1 78/1 77/1 98/1 94/1 78/1 73/0.98 100/1 75/0.96 71/0.98 85/1 78/1 -/0.95 73/- 94/1 95/1 100/1 81/0.98 100/1 100/1 77/0.99 100/1 88/1 -/0.95 -/1 -/0.99 -/0.99 -/0.99 -/0.96 -/0.98 Figure 1. IQ-tree based on a nuITS rDNA dataset for Cora spp. The names of species are followed with their GenBank accession numbers (see Suppl. material 1). Newly sequenced specimens are marked in bold. Bootstrap support values (%) from IQ-tree analysis and PP support from Bayesian analysis are indicated near the branches. Sequences of Corella tomentosa KJ780578 and Corella zahlbruckneri KJ780592 were used as outgroup taxa. 6 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia described Cora species (Lücking et al. 2013, 2017; Vargas et al. 2014). These are C. arcabucana (2 specimens), C. cuzcoensis (1 specimen), and C. undulata L (1 specimen), which are new to Bolivia, and five records belong to C. aspera, which was recorded only on three localities in Bolivia (Fig. 1). Nine sequences form two, closely related, distinct lineages in the Cora tree (Fig. 1), suggesting the presence of two new species, named here as C. neoparabovei and C. neopseudobovei. These species are related to C. parabovei Dal-Forno, Kukwa & Lücking. Other taxa closely related to these new species are C. bovei Speg., C. campestris Dal-Forno, Eliasaro & A.A. Spielm., C. palaeotropica Weerakoon, Aptroot & Lücking, C. pseudobovei Wilk, Dal-Forno & Lücking, and C. celestinoa B. Moncada, Cabr.-Amaya & Lücking. All these species are characterized by terricolous thallus with more or less brown or olive thallus when fresh. Based on their morphological similarity and the tree topology, we propose that the group of all of these species should be named as the Cora bovei group. The first novel species, C. neoparabovei is represented by four specimens from Bolivia. All specimens are characterised by the long, concentrically arranged setae on the surface of the thallus and densely pilose in the margins. This species is phylogenetically, morphologically and ecologically similar to C. parabovei and other members of the clade (Fig. 1). In the haplotype network (Fig. 2), the sequences of C. neoparabovei are represented by two haplotypes (but see also Suppl. material 1 and Suppl. material 2). The three specimens (PV647979, PV647980, PV647981), share the same haplotype, which differs from the second (PV647982) haplotype in a single site. These differ from C. parabovei (KJ780571) by thierteen or fourteen nucleotide substitutions (Fig. 2, Suppl. material 2). The second of the new species, C. neopseudobovei is represented by five specimens from Bolivia. To compare the genetic diversity between the new species and other related taxa, we carried out a haplotype network analysis. In the haplotype network (Fig. 2), the sequences of C. neopseudobovei are represented by two haplotypes (see also Suppl. material 1 and Suppl. material 2). The sequences of two specimens (PV647974, PV647977), share the same haplotype (Fig. 2), and differ from the second haplotype (PV647975, PV647976, PV647978) in a single position. Five sequences of C. pseudobovei (KJ780540, KJ780539, KJ780470, KJ780426, KJ780417) have the same haplotype, which differs by thirteen or fourteen nucleotide substitutions from C. neopseudobovei. It seems interesting that the new Cora species described here have only been found in single localities, which may presumably suggest their endemism. The occurrence of endemic species of Cora has been noted, for example, in the United States of America, where Cora timucua is endemic to Florida (Lücking et al. 2020; Dal Forno et al. 2021) and from Galapagos, including C. galapagoensis Dal-Forno, Bungartz & Lücking, and C. santacruzensis Dal-Forno, Bungartz & Yánez-Ayabaca (Dal Forno et al. 2017; Lücking et al. 2017). However, as the genus is still not well studied in many regions, some species may appear more widespread. In this paper, we report the first record of C. arcabucana from Bolivia, which has been known only from Colombia so far (Lücking et al. 2017), C. cuzcoensis previously known only from the type locality near Machu Picchu in Peru (Lücking et al. 2017), and C. undulata, previously noted only in Colombia (Vargas et al. 2014). 7 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia Taxonomy New species from Bolivia Cora neoparabovei Oset, Flakus & Guzow-Krzem., sp. nov. MycoBank No: 860019 Fig. 3 Diagnosis. Species very similar to C. parabovei, but differing in the distinct phylogenetic position within the genus, and in substitution of several nucleotide positions in nuITS rDNA (see also Suppl. material 2). Taxon is characterized by long, concentrically arranged setae present on the thallus surface and densely pilose margins. Type. Bolivia • Dept. La Paz, Prov. Nor Yungas, PNANMI Cotapata, N of Unduavi, by Sillu Tincara pre-Columbian route, 16°16'33"S, 67°52'60"W, elev. 3429 m, Yungas cloud forest, 27 June 2010, A. Flakus 16965 & P. Rodriguez (holotype KRAM; isotype LPB). Description. Thallus lichenized, terricolous over and between bryophytes, foliose, up to 8.5 cm across, composed of 3 semicircular, adjacent lobes; individual lobes up to 8 cm wide and 2.5–6.5 cm long, turned upwards, circular patches sparsely branched, without radial branching sutures, surface olive green to pale yellow in the centre with a slight concentric zonation of colour when fresh but when dried yellowish-grey. Margins thin, involute, green becoming yellow-green in the herbarium. Upper surface evenly to shallowly undulating when fresh, rugose when dry, with yellowish setae throughout, with greater density towards the centre of the thallus; setae consisting of hyphae up to 300 μm long and 5 μm wide, hyphae irregular in shape, sparingly branched and anastomosing, hyaline to very pale brown; lower surface ecorticate, felty-arachnoid (representing the exposed medulla), green-yellow when fresh, becoming yellow to brown Figure 2. Haplotype network showing relationships between nuITS rDNA sequences from Cora bovei, C. neoparabovei, C. neopseudobovei, C. palaeotropica, C. parabovei and C. pseudobovei and Cora sp.. The names of species are followed with their GenBank accession numbers. The newly sequenced samples are marked in bold (see Suppl. material 1). Mutational changes are presented as hatch marks. Each species is marked in colour with ellipse. 8 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia when dried. Thallus in the section up to 550 μm thick, with upper cortex, photobiont layer, and medulla; upper cortex collapsed compacted; photobiont layer up to 250 μm thick, orange-brown (upper portion) to aerugious (lower portion); medulla 100 μm thick; clamp connections absent, papilliform hyphae absent. Hymenophore not present. No substances detected by TLC. Habitat and distribution. Cora neoparabovei grows on bryophytes and is known only from three localities in Nor Yungas province in La Paz department, occurring at elevations between 3210 m and 3429 m in Yungas cloud forest. Etymology. The name is derived from the Greek prefix néos, meaning “new”, combined with “parabovei”, referring to its similarity to C. parabovei. Additional material examined. Bolivia • Dept. La Paz; Prov. Nor Yungas, below Unduavi village, on the road La Paz – Chulumani, 16°18'27"S, 67°53'48"W, elev. 3211 m, Yungas cloud forest, on bryophytes, 31 May 2011, A. Flakus 22219 & O. Plata (KRAM L-71638; LPB). elev. 3210 m, Yungas cloud forest, on bryophytes, 31. May 2011, A. Flakus, O. Plata (UGDA–20029; LPB). • PNANMI Cotapata, 5 hours walking from Unduavi by Sillu Tincara pre-Columbian route, 16°16'33"S, 67°52'60"W, elev. 3429 m, Yungas cloud forest, 27 June 2010, A. Flakus 16855/2 & P. Rodriguez (KRAM; LPB). Notes. Cora neoparabovei is phylogenetically most closely related to C. bovei, C. campestris, C. celestinoa, C. parabovei, C. palaeotropica, C. pseudobovei, but morphologically and ecologically it is the most similar only to C. parabovei (Lücking et al. 2013, 2017). Both species are characterized by a rather unique Figure 3. Morphology of Cora neoparabovei (holotype). A. Upper surface (on the left side thallus is dry, on the right side thallus is wet); B. Felty-arachnoid lower surface (thallus is dry). Scale bars: 10 mm (A, B). 9 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia pattern of dense hairs formed in concentric zones. Cora neoparabovei, is one of four described species with concentrically arranged surface hairs, aside from C. dewisanti B.Moncada, Suár.-Corr. & Lücking from Venezuela, Ecuador and Colombia, and C. maxima Wilk, Dal-Forno & Lücking and C. parabovei also from Bolivia. Nevertheless, C. dewisanti differs in mostly glabrous surface except for concentric bands of whitish setae, and with distinct, whitish, glabrous margins and presents a corticioid-cyphelloid hymenophore, C. maxima has glabrous lobe margins and is much larger, whereas C. parabovei has different colour of thallus when is dried (grey), which has concentric zonation both when fresh and dried (Lücking et al. 2017). Cora neopseudobovei Oset, Flakus & Guzow-Krzem., sp. nov. MycoBank No: 860020 Fig. 4 Diagnosis. Species very similar to C. pseudobovei, but differing in the distinct phylogenetic position within the genus, and in substitution of several nucleotide posititions in nuITS (see also Suppl. material 2). The diagnostic features of the species are the small, macrosquamulose, up to 2 cm across brown upper surface thallus when fresh, pale yellow to orange-yellow when dry, with slightly visible concentric colour zonation when dry and also with involute, creamy white margins. Type. Bolivia • Dept. La Paz; Prov. Franz Tamayo, ANMIN Apolobamba, near Puyo Puyo village, 14 °56'55"S, 69°07'58"W, elev. 4888 m, high Andean open vegetation, 5 July 2010, A. Flakus 17603 & P. Rodriguez (holotype KRAM; isotype LPB). Description. Thallus lichenized, terricolous, foliose, small, macrosquamulose, up to 2 cm across, composed of up to 8 semicircular, adjacent to subimbricate lobes; individual lobes up to 1.2 cm wide and 6 mm long, moderately branched, without radial branching sutures, surface brown when fresh, with slightly visible concentric colour zonation when dry, with involute, creamy white margins. Upper surface shallowly undulate or scabrous when fresh and dry, involute margins undulate; lower surface excorticate, felty-arachnoid (representing the exposed medulla), brown-olive. Thallus in section 200–300 μm thick, with upper cortex, photobiont layer, and medulla; upper cortex collapsed compacted, formed by up 40 μm thick layer of loosely packed, irregularly arranged; photobiont layer up to 100 μm thick, orange above, aeruginous below, separated from the medulla by a thick, compacted layer of brownish hyphae; medulla 100 μm thick, often indistinct; clamp connections absent, papilliform hyphae absent. Hymenophore in section 50 μm thick, spores 5 × 3 μm, basidioles 20– 25 × 3–6 μm. Sterigmata 4. Chemistry: No substances detected by TLC. Habitat and distribution. Cora neopseudobovei is known from four localities in La Paz department (Bautista and Franz Tamayo provinces), occurring at elevations between 3780 m and 4888 m in open area with shrubs and high Andean vegetation. The species was found on soil. Etymology. The name refers to the similarity in morphology to Cora pseudobovei. Additional material examined. Bolivia • Dept. La Paz; Prov. Bautista Saavedra, ANMIN Apolobamba, between la Curva and Charazani, 15°08'09"S, 69°02'03"W, 16 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia mushrooms (Basidiomycota: Agaricales: Hygrophoraceae). Mycological Research 113(10): 1154–1171. https://doi.org/10.1016/j.mycres.2009.07.016 Lücking R, Dal-Forno M, Lawrey JD, Bungartz F, Rojas MEH, Hernández MJE, Marcelli MP, Moncada B, Morales EA, Nelsen MP, Paz E, Salcedo L, Spielmann AA, Wilk K, WillWolf S, Yánez-Ayabaca A (2013) Ten new species of lichenized Basidiomycota in the genera Dictyonema and Cora (Agaricales: Hygrophoraceae),with a key to all accepted genera and species in the Dictyonema clade. Phytotaxa 139(1): 1–38. https://doi. org/10.11646/phytotaxa.139.1.1 Lücking R, Grall A, Thüs H (2014a) Typification of Thelephora pavonia Sw. and reinstatement of Cora ciferrii (Tomas.) comb. nov. Lichenologist (London, England) 46(6): 825–828. https://doi.org/10.1017/S0024282914000401 Lücking R, Dal-Forno M, Wilk K, Lawrey JD (2014b) Three new species of Dictyonema (lichenized Basidiomycota: Hygrophoraceae) from Bolivia. Acta Nova 6: 4–16. Lücking R, Dal-Forno M, Sikaroodi M, Gillevet PM, Bungartz F, Moncada B, Yánez A, Chaves JL, Coca LF, Lawrey JD (2014c) A single macrolichen constitutes hundreds of unrecognized species. Proceedings of the National Academy of Sciences of the United States of America 111(30): 11091–11096. https://doi.org/10.1073/ pnas.1403517111 Lücking R, Barrie FR, Genney D (2014d) Dictyonema coppinsii, a new name for the european species known as Dictyonema interruptum (Basidiomycota: Agaricales: Hygrophoraceae), with a validation of its photobiont Rhizonema (Cyanoprokaryota: Nostocales: Rhizonemataceae). Lichenologist (London, England) 46(3): 261–267. https://doi.org/10.1017/S0024282913000352 Lücking R, Cáceres MES, Silva NG, Alves RJV (2015) The genus Cora in the South Atlantic and the Mascarenes: Two novel taxa and inferred biogeographic relationships. The Bryologist 118(3): 293–303. https://doi.org/10.1639/0007-2745-118.3.293 Lücking R, Dal Forno M, Moncada B, Coca LF, Vargas-Mendoza Y, Aptroot A, Arias LJ, Besal B, Bungartz F, Cabrera-Amaya DM, Cáceres MES, Chaves JL, Eliasaro S, Gutiérrez MC, Hernández-M JE, Herrera-Campos MA, Holgado-Rojas ME, Jonitz H, Kukwa M, Lucheta F, Madriñán S, Marcelli MP, Martins SMA, Mercado-Díaz JA, Molina JA, Morales EA, Nelson PR, Nugra F, Ortega F, Paredes T, Patiño AL, Peláez-Pulido RN, Pérez-Pérez RE, Perlmutter GB, Rivas-Plata ME, Robayo J, Rodríguez C, Simijaca DF, Soto-Medina E, Spielmann AA, Suárez-Corredor A, Torres JM, Vargas CA, Yánez-Ayabaca A, Weerakoon G, Wilk K, Celis-Pacheco M, Diazgranados M, Brokamp G, Borsch T, Gillevet PM, Sikaroodi M, Lawrey JD (2017) Turbo-taxonomy to assemble a megadiverse lichen genus: seventy new species of Cora (Basidiomycota: Agaricales: Hygrophoraceae), honouring David Leslie Hawksworth’s seventieth birthday. Fungal Diversity 84(1): 139–207. https://doi.org/10.1007/s13225-016-0374-9 Lücking R, Kaminsky L, Perlmutter GB, Lawrey JD, Dal Forno M (2020) Cora timucua (Hygrophoraceae), a new and potentially extinct, previously misidentified basidiolichen of Florida inland scrub documented from historical collections. The Bryologist 123(4): 657–673. https://doi.org/10.1639/0007-2745-123.4.657 Lutzoni F, Vilgalys R (1995) Omphalina (Basidiomycota, Agaricales) as a model system for the study of coevolution in lichens. Cryptogamic Botany 5: 71–81. Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees in Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA, 1–8. https://doi. org/10.1109/GCE.2010.5676129 17 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia Nguyen LT, Schmidt HA, Von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300 Oberwinkler F (1970) Die Gattungen der Basidiolichenen. Vortr Gesamtgeb Bot 4: 139– 169. Oberwinkler F (1984) Fungus-alga interactions in basidiolichens. Beiheft zur Nova Hedwigia 79: 739–774. Oberwinkler F (2012) Basidiolichens. In The Mycota. 2nd Edn, Vol. IX. Fungal Associations. In: B. Hock (Ed.) Berlin, Heidelberg, New York. 341–362. Springer. https://doi. org/10.1007/978-3-642-30826-0_16 Orange A, James PW, White FJ (2001) Microchemical methods for the identification of lichens. British Lichen Society, London. Oset M, Guzow-Krzemińska B, Flakus A, Rodriguez-Flakus P, Kukwa M (2024) New records of Cora species (Hygrophoraceae, Agaricales, lichenized Basidiomycota) from Bolivia confirmed by molecular data. Phytotaxa 670(2): 083–096. https://doi. org/10.11646/phytotaxa.670.2.1 Parmasto E (1978) The genus Dictyonema (‘Thelephorolichenes’). Nova Hedwigia 29: 99–144. Petersen RH (1967) Notes on clavarioid fungi. VII. Redefinition of the Clavaria vernalis –C. mucida complex. American Midland Naturalist 77(1): 205–221. https://doi. org/10.2307/2423440 Rambaut A (2009) FigTree v1.4., A Graphical Viewer of Phylogenetic Trees. http://tree. bio.ed.ac.uk/software/figtree/ Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior Summarization in Bayesian Phylogenetics Using Tracer 1.7. Systematic Biology 67(5): 901–904. https://doi.org/10.1093/sysbio/syy032 Redhead SA, Lutzoni F, Moncalvo JM, Vilgalys R (2002) Phylogeny of agarics: Partial systematics solutions for core omphalinoid genera in the Agaricales (euagarics). Mycotaxon 83: 19–57. Rodriguez-Flakus P, Kukwa M, Etayo J, Lücking R, Meneses RI, Rivas Plata E, Stanton D, Truong C, Vargas R, Flakus A (2016) Preliminary catalogue of lichens and lichenicolous fungi from Bolivia. http://bio.botany.pl/lichens-bolivia/en,strona,catalogue,5. html [Version 1.5, 31 December 2016] Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics (Oxford, England) 19(12): 1572–1574. https://doi. org/10.1093/bioinformatics/btg180 Ronquist F, Huelsenbeck J, van der Mark P (2005) MrBayes 3.1 manual. Distributed with the Program by the Authors. Tamura K, Stecher G, Kumar S (2021) MEGA11: Molecular Evolutionary Genetics Analysis version 11. Molecular Biology and Evolution 38(7): 3022–3027. https://doi. org/10.1093/molbev/msab120 Vargas LY, Moncada B, Lücking R (2014) Five new species of Cora and Dictyonema (Basidiomycota: Hygrophoraceae) from Colombia: chipping away at cataloging hundreds of unrecognized taxa. The Bryologist 117(4): 368–378. https://doi. org/10.1639/0007-2745-117.4.368 Vizzini A, Alvarado P, Consiglio G, Marchetti M, Xu J (2024) Family matters inside the order Agaricales: Systematic reorganization and classification of incertae sedis clitocyboid, pleurotoid and tricholomatoid taxa based on an updated 6-gene phylogeny. Studies in Mycology 107(1): 67–148. https://doi.org/10.3114/sim.2024.107.02 18 MycoKeys 126: 1–18 (2025), DOI: 10.3897/mycokeys.126.165395 Magdalena Oset et al.: New species of Cora (lichenized Basydiomycota, Hygrophoraceae) from Bolivia White TJ, Bruns T, Lee S, Taylor J (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innnis MA, Gelfand DH, Sninsky JJ & White TJ (Eds) PCR protocols: a guide to methods and applications. Academic Press, New York, 315–22. https://doi.org/10.1016/B978-0-12-372180-8.50042-1 Yánez A, Dal-Forno M, Bungartz F, Lücking R, Lawrey JD (2012) A first assessment of Galapagos basidiolichens. Fungal Diversity 52(1): 225–244. https://doi.org/10.1007/ s13225-011-0133-x Supplementary material 1 Specimens of Cora used in molecular analysis Authors: Magdalena Oset, Adam Flakus, Beata Guzow-Krzemińska Data type: xlsx Explanation note: Specimens of Cora used in molecular analysis with GenBank accession numbers, voucher numbers, country of origin, and source publication. Sequences generated for this study are in bold. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.126.165395.suppl1 Supplementary material 2 Variable positions in the alignment of nuITS rDNA marker Authors: Magdalena Oset, Adam Flakus, Beata Guzow-Krzemińska Data type: xlsx Explanation note: Variable positions in the alignment of nuITS rDNA marker of Cora bovei, C. neoparabovei, C. neopseudobovei, C. palaeotropica, C. parabovei, C. pseudobovei and Cora sp. Variable characters are marked in bold, while diagnostic nucleotide position characters to distinguish these species from other taxa of this group are marked with a yellow background. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.126.165395.suppl2