683 MycoKeys MycoKeys 114: 259-276 (2025) DOI: 10.3897/mycokeys.114.139681 Research Article Sticta flakusiorum and S. kukwae—two additional new species from the Neotropics (Peltigerales, Peltigeraceae) Emilia Anna Ossowska™, Bibiana Moncada22“®, Robert Liicking?“®, Emmanuel Sérusiaux™®, Nicolas Magain>® on FF WoO NY — Department of Plant Taxonomy and Nature Conservation, Faculty of Biology, University of Gdarisk, Wita Stwosza 59, PL-80-308 Gdarisk, Poland Licenciatura en Biologia, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26D-54, Torre de Laboratorios, Herbario, Bogota D.C., Colombia Research Associate, Science & Education, The Field Museum, 1400 South Lake Shore, Chicago, IL 60605, USA Botanischer Garten und Botanisches Museum Berlin, Freie Universitat Berlin, Konigin-Luise-StraBe 6-8, 14195 Berlin, Germany Evolution and Conservation Biology, University of Liége, Sart Tilman B22, 4000 Liége, Belgium Corresponding author: Emilia Anna Ossowska (emilia.ossowska@ug.edu.p!) OPEN @ ACCESS Academic editor: Gerhard Rambold Received: 18 October 2024 Accepted: 24 January 2025 Published: 4 March 2025 Citation: Ossowska EA, Moncada B, Lucking R, Sérusiaux E, Magain N (2025) Sticta flakusiorum and S. kukwae—two additional new species from the Neotropics (Peltigerales, Peltigeraceae). MycoKeys 114: 299-276. https://doi.org/10.3897/ mycokeys.114.139681 Copyright: © Emilia Anna Ossowska 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). Abstract Two additional species of Sticta are described as new to science based on material from Bolivia and Peru and supported by phylogenetic analysis of the fungal ITS barcoding marker. The two new species represent lineages within clade | on the global Sticta phy- logeny. Sticta flakusiorum Ossowska, B. Moncada & Licking is a species in the S. hum- boldtii morphodeme and is characterized by lobes partly to entirely covered with white hairs, also covering the margins of submarginal and laminal apothecia, and the scabrid basal membrane of cyphellae, which is white to yellow, or partly brown, and when yellow K+ purple. The taxon was discovered at a single locality in Bolivia, but it is closely related to a potentially new Sticta species from Peru, which is here left undescribed. The other new species, S. kukwae Ossowska, Magain & Sérus., belongs to the S. weigelii mor- phodeme. It has lobes with sinuous margins and dark, palmate to corymbose phyllidia. It was collected at several locations in Peru and a single locality in Bolivia. Key words: Bolivia, diversity, integrative taxonomy, ITS rDNA, Lobarioideae, Peltigeraceae, Peru Introduction Lobarioid lichens, long treated in their own family, Lobariaceae (Moncada et al. 2013a), are now recognized as a subfamily, Lobarioideae, within Peltigera- ceae, along with subfamilies Nephromatoideae and Peltigeroideae (Kraichak et al. 2018; Licking 2019; Lumbsch and Leavitt 2019; Widhelm et al. 2019, 2021). Traditionally, the presence or absence of regular pores in the lower cor- tex, called cyphellae, was the main feature that differentiated the genus Sticta (Schreb.) Ach. from other genera in the family (Galloway and Elix 2013; Mon- cada et al. 2013a); however, apart from Sticta, a second lineage, nested within the Lobaria clade and separated in the genus Dendriscosticta B. Moncada & Licking, also features cyphellae (Moncada et al. 2013a; Simon et al. 2022). In addition to Dendriscosticta, another nine genera have recently been segregated 299 Emilia Anna Ossowska et al.: ShortTitle from the collective genera Lobaria (Schreb.) Hoffm., Pseudocyphellaria Vain., and Sticta, including, e.g., Yarrumia D.J. Galloway (Galloway 2015) and Emman- uelia Ant. Simon, Licking & Goffinet (Simon et al. 2020). Currently, close to 500 species have been accepted in the subfamily Lobarioideae (Kirk et al. 2008; Licking et al. 2017), almost half of them within the genus Sticta (Moncada et al. 2014a, 2021a; Licking et al. 2017; Ossowska et al. 2022a). The species of this genus are common in humid, cool to warm environments with high rain- fall or humidity and are most diverse in tropical areas (Moncada 2012; Mon- cada et al. 2014b, 2020). This is illustrated by the example of Colombia, where more than 150 Sticta species have been identified following extensive field and laboratory work (Moncada 2012; Moncada and Liicking 2012; Moncada et al. 2013b, 2014a, b, 2015, 2021b). In comparison, the knowledge on the genus Sticta in neighboring countries is limited. For Ecuador, 46 names have been listed (Consortium of Lichen Herbaria 2023; Yanez-Ayabaca et al. 2023), but at least eight of these are doubtful records, including some known New Zealand endemics. Twenty-two species are included in the revised checklist for Brazil (Aptroot 2002; Dal Forno et al. 2018; Torres et al. 2021); however, Torres et al. (2021) suggest that the Cerrado forest ecoregion may host a significant num- ber of novel Sticta species, indicating that the true diversity of species in Brazil may be higher. The checklist of lichens for Peru includes only ten Sticta species (Ramos 2014), but the majority of the records are historical and have not been critically checked. Meanwhile, twenty-eight Sticta taxa are known from Bolivia, mostly based on recent works (Moncada and Licking 2012; Ossowska 2021; Ossowska et al. 2022a, b, 2024a, b; Crous et al. 2023). This paper presents two additional new Sticta species, S. flakusiorum and S. kukwae, both supported by molecular data, from Peru and/or Bolivia. Sticta flakusiorum has been found so far at a single site in Bolivia, whereas S. kuk- wae has been collected from several localities in Peru and Bolivia. Detailed morphological and anatomical descriptions of both species are also given, together with a discussion on similar taxa. Materials and methods Taxon sampling Fresh material for this study was collected during fieldwork in Bolivia in 2010- 2017 and Peru in 2012. The collected material is deposited in the LPB, UGDA, and DUKE herbaria. All material was examined under a dissecting and a compound microscope (Nikon SMZ800N and ZEISS Axioskop). Character assessment was based on the morphological and anatomical traits for Sticta described by Monca- da (2012) and Moncada et al. (2014a). Spot reactions were done with K (potas- sium hydroxide solution), C (sodium hypochlorite solution), Pd (paraphenylenedi- amine), and KC (K followed by C) on close spots of exposed medulla of the same thallus fragments; secondary compounds were analyzed using the thin-layer chromatography method (TLC) in solvents A and C (Orange et al. 2001). Species that were informally distinguished by Moncada (2012) and Mon- cada et al. (2020) but have not yet been formally described are marked with quotes (e.g., ‘S. arachnosylvatica’). MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 260 Emilia Anna Ossowska et al.: ShortTitle DNA extraction, PCR amplification, and sequencing Genomic DNA from the Bolivian samples was isolated, and the nulTS rDNA marker was amplified following the protocol described in Ossowska et al. (2022a). Sequencing was performed in a Macrogen sequencing system (http:// www.macrogen.com). In the case of samples from Peru, DNA was extracted following the protocol of Cubero et al. (1999). PCR conditions and primers were the same as in Ossowska et al. (2022a). Alignment and sequence analyses The newly generated sequences were compared with available data from the genus Sticta (Suppl. material 1), using our previous alignment (Ossowska et al. 2022a) based on a recent master alignment (Moncada et al. 2020). The new sequences were added to the existing alignment using MAFFT 7.164 with the “—add” option (Katoh and Frith 2012; Katoh and Standley 2013), followed by manual checking in BIOEDIT 7.0.9 (Hall et al. 2011). Phylogenetic analysis was performed using maximum likelihood in RAXML 8.2.0 (Stamatakis 2014) onthe CIPRES Science Gateway (Miller et al. 2010), with non-parametric bootstrap- ping using 400 pseudoreplicates (based on an automated saturation criterion) under the universal GTR GAMMA model. Trees were visualized in FigTree 1.4.2 (Drummond and Rambaut 2007) and edited using Coral Draw 2019. Results We generated seven new nulTS rDNA sequences that form two distinct lineag- es in the Sticta tree (Fig. 1), suggesting the presence of three new species, two of which are closely related sister species. The new sequences align close to other Sticta species, such as S. sylvatica (Huds.) Ach. and S. aymara Ossowska et al., within clade | (fuliginosa clade) sensu Widhelm et al. (2018). The first of the new species, S. kukwae, is represented by one specimen from Bolivia and four from Peru. All specimens have a thallus with strongly sinuous margins and very dark phyllidia. One specimen (LG3227) from Peru had small and sparse apothecia, absent in the other specimens. However, other characteristics were consistent with the rest of the specimens in this clade (Fig. 1). Within clade I, the new species is closely related to S. umbili- cariiformis Hochst. ex Flot. (Fig. 1). The second new lineage is formed by two sister species. The lineage with a specimen from Bolivia is named in this paper S. flakusiorum and is closely related to the specimens from Peru, which potentially also represent a new species. At this point, we have named it Sticta sp. 36. It is a phyllidiate species (S. flakusiorum lacks vegetative diaspores), and morphologically it is similar to S. phyllidiokunthii B. Moncada & LUcking, with numerous, aggregated, palmate phyllidia that are marginal and laminal in S. sp. 36 and marginal in S. phyllidio- kunthii (Moncada et al. 2013b). However, the material of S. sp. 36 is too sparse for a formal description at this point. Both species, S. flakusiorum and S. sp. 36, are nested in the clade of S. viviana Alej. Suarez & Licking and are closely related to S. phyllidiofuliginosa B. Moncada, A. Suarez & Licking. MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 261 Emilia Anna Ossowska et al.: ShortTitle 100 Sticta macrothallina_KC732656_DNA5562b-MON0431b_Colombia_Coca_1376 Sticta macrothallina_KC732655_DNA5562a-MON0431a_Colombia_Coca_1376 Sticta macrothallina_KC732637_DNA5539-MON0408_Colombia_Coca_1115 Sticta andensis_KC732547_DNA5016-MON0199_ Colombia_Alfonso_2 Sticta pseudolimbata_KC732467_DNA4922-MON0031_Colombia_Moncada_4009 Sticta pseudolimbata_KC732564_DNA5340-MON0185_Colombia_Moncada_4022 Sticta pseudolimbata_KC732468_DNA4923-MON0032_Colombia_Sipman_33451 Sticta duplolimbata_OP999519_Tanzania_Kaasalainen_UK-170914b "Sticta duplolimbata_OP999460_Kenya_Rikkinen_JR-16374b Sticta duplolimbata_OP999582_Tanzania_Kaasalainen_UK-171513f Sticta cometiella_KC732606_DNA5470-MON0339_Colombia_Moncada_4669 *Sticta cometiella | KC732516_DNA4976-MON0127_Colombia_Moncada_4404 Sticta cometiella_KC011068_DNA5024-MON0227_Colombia_Moncada_4152 Sticta cometia_KC732642_DNA5544-MON0413_Colombia_Coca_1044 Sticta cometia_KC732627_DNA5527-MON0396_Colombia_Coca_1058 Sticta cometia_KC732641_DNA5543-MON0412_Colombia_Coca_1078 100 Sticta aymara_NR-189895_Bolivia_Flakus_17220 Sticta aymara_OP250126 Bolivia Flakus_17220 Sticta narinioana_OP 244961_MON2272_Colombia_Moncada_7525 sticta narinioana_OP244963_MON2280_Colombia_Moncada_7614 Sticta narinioana_OP244962_MON2696_Colombia_Simijaca_2044 100 97 Sticta gretae_MN700002_USA_Oregon_Di-Meglio_78 Sticta gretae_MN700003_USA_Califomia_Di-Meglio_84 Sticta gretae_MN700007_USA_Oregon_Di-Meglio_150 Sticta madidensis_Ecuador_ Benitez Sticta madidensis MON6164_Colombia_Moncada_11480 Sticta madidensis_PP273990_Bolivia_Kukwa_14879 Sticta madidensis_MON2317_Colombia_Moncada_7599 87) Sticta madidensis MON6155_Colombia_Moncada_11484 Sticta madidensis_MON2314_Colombia_Moncada_7613 Sticta madidensis_MON2318_Colombia_Moncada_7615 100 Sticta fasciculata_MN699993 Canada-British-Columbia_Di-Meglio_91 Sticta fasciculata_MN699995_USA-Oregon_Di-Meglio_161 Sticta fasciculata_MN699989_USA-Alaska_Dillman_DM-25-2018 100 = Sticta phyllidiofuliginosa_KC732764_DNA6342-MON0559_ Colombia_Moncada_4972 Sticta phyllidiofuliginosa_KC732508_DNA4969-MON0109_Colombia_Moncada_4052 a Sticta phyllidiofuliginosa_KC732495_DNA4958-MON0088_Colombia_Moncada_4051 16 Sticta flakusiorum_PQ461155_Bolivia_Kukwa_14677 Sticta sp 36_LG3235_Peru_Magain_sn Sticta sp 36_.G3229_Peru_Magain_sn 400 Sticta globulifuliginosa_KC 732601_DNA5464-MON0333_Colombia_Moncada_4747 96 Sticta globulifuliginosa_KC732678_DNA5592a-MON0461a_Colombia_Luecking_33318 93 Sticta globulifuliginosa_KC732582_DNA5445-MON0314_ Colombia_Moncada_4742 100 Sticta viviana_GB_DNA-MON4685_Colombia_Moncada_10507 Sticta viviana_KC732680_DNA5593-MON0462_Colombia_Luecking_33311 Sticta viviana_KC732692_DNA5611-MON0480_Colombia_Moncada_4808 2 Sticta viviana_KC732585_DNA5448-MON0317_Colombia_Moncada_4748 Sticta viviana_GB_DNA7244-MONO0704_CostaRica_Moncada_5793 100 Sticta atlantica_KT281734_LG3747_NA_Ireland_Serusiaux_sn Sticta atlantica_KT281737_LG3858_NA_Azores_Serusiaux_sn 82) Sticta atroandensis_KC732533_DNA4999-MON0167_Colombia_Fonseca_23 Sticta atroandensis_KC732528_DNA4989-MON0147_Colombia_Fonseca_188b Sticta atroandensis_KC732527_DNA4988-MON0146_Colombia_Fonseca_188a 96 Sticta pseudohumboldtii_KC732463_DNA4917-MON0025_Colombia_Moncada_4586 Sticta pseudohumboldtii_KC732737_DNA6300-MON0517_Colombia_Moncada_4921 i) Sticta pseudohumboldtii_KC732543_DNA5012-MON0190_Colombia_Moncada_4635a 95 | Sticta arachnofuliginosa_KC732512_DNA4973-MON0116_Colombia_Moncada_4049 Sticta arachnofuliginosa_KC732524_DNA4985-MON0143_Colombia_Moncada_4007a Sticta arachnofuliginosa_KC732458_DNA4911-MON0015_Colombia_Moncada_4595a 100 Sticta arachnosylvatica_KC732689_DNA5606-MON0475_Colombia_Moncada_4803 P9 Sticta arachnosylvatica_KC732580_DNA5443-MON0312_Colombia_Moncada_4733 Sticta arachnosylvatica_KC732718_DNA6236-MON0650_Colombia_Diaz_L8 81 Sticta macrofuliginosa_KC732747_DNA6316-MON0533_Colombia_Moncada_4042 174 Sticta parahumboldtii_| KC732550_DNA5019-MON0209_Colombia_Moncada_4016 4 Sticta parahumboldtii_| KC732573_DNA5425-MON0294_ Colombia_Betancourt_144f 88 Sticta hirsutofuliginosa_KC732612_DNA5479-MONO348_Colombia_Moncada_4734a 20; Sticta hirsutofuliginosa_KC732611_DNA5478-MON0347_Colombia_Moncada_4734c 78 Sticta hirsutofuliginosa_KC732501_DNA4964-MONO0099_Colombia_Moncada_4032 83 ag Sticta humbordtil_| MT507786_DNA14382-MON4382_Colombia_Luecking_39360 88 Sticta humboldtii_KC732705_DNA6202-MON0616_Colombia_Diaz_L7 Sticta humboldtii_KC732704_DNA6201-MON0615_Colombia_Diaz_L5 Sticta munda_OP999598_Tanzania_Kaasalainen_UK-171582i Sticta munda_OP999600_Tanzania_Kaasalainen_UK-171584u Sticta munda_OP999581_ Tanzania_Kaasalainen_UK-171510k Sticta afromontana_OP999554_Tanzania_Kaasalainen_UK-171439t 191 Sticta afromontana_OP999503_Tanzania_Kaasalainen_UK-170857a Sticta afromontana_OP999601_Tanzania_Kaasalainen_UK-171584v Sticta ambavillaria_}Q735978_LG0992_Reunion_Magain_sn Sticta limbata_MN700025_USA-Califomia_Di-Meglio_85 Sticta limbata_MG367428_DNA8131-MON1262_USA-Hawaii_Moncada_6995 Sticta limbata_KT281728_1G3544_France_Serusiaux_sn % Sticta limbata_| 7 KT281708_LG2749_Spain-Canary-Islands_Serusiaux_sn Sticta limbata_KT281707_LG2690_Scotland_Serusiaux_sn 79, Sticta fuliginosa_PP291713_Canada-Ontario_Brinker_7920 Sticta fuliginosa_MT526260_UK_unknown_S10 Sticta fuliginosa_OP999486_Tanzania_Kaasalainen_UK-170794g Sticta fuliginosa_AF350310_New-Zealand_Thomas_1061 Sticta fuliginosa_MG367426_DNA8112-MON1243_USA_Hawaii_Moncada_6978 Sticta aspratilis_OP999475_Kenya_Kaasalainen_UK-110551f Sticta aspratilis_OP999607_Tanzania_Kaasalainen_UK-171587j Sticta aspratilis_OP999530_Tanzania_Kaasalainen_UK-170975f Sticta pseudosylivatica_KC732498_DNA4961-MON0091_Colombia_Moncada_2074 99 Sticta pseudosylvatica_KC732725_DNA6277-MON0494_ Colombia_Suarez_305 Sticta pseudosylivatica_KC732724_DNA6276-MON0493_Colombia_Suarez_306 90 Sticta umbilicariiformis_OP999596_Tanzania_Kaasalainen_UK-171577h 87 Sticta umbilicariiformis_KT281697_LG0925 Rwanda_Magain_sn Sticta umbilicariiformis_OP999434_Kenya_Rikkinen_JR-16098 99 100 Sticta kukvae_PQ461157_LG3225 Peru_Magain_sn Sticta kukwae_PQ461156_Bolivia_Kukwa_14729a 934 Sticta kukwae_PQ461158_LG3221 Peru_Magain_sn 83Sticta kukwae_PQ461160_LG3227_Peru_Magain_sn Sticta kukwae_PQ461159_LG3223_Peru_Magain_sn 100 0.02 Figure 1. Best-scoring maximum likelihood tree of the Sticta target clade containing the new species S. flakusiorum from Bo- livia (blue), S. kukwae from Bolivia and Peru (red), and S. sp. 36 from Peru (green), based on the fungal ITS barcoding marker. Branches associated with high bootstrap support values ( = 70) are thickened and values are indicated near the branches. Detailed descriptions of morphological and anatomical characteristics of S. flakusiorum and S. kukwae, together with figures and comparisons with similar and related species, are given below. Discussion As agenus, Sticta is relatively easy to recognize in the field due to the foliose, large thallus with tomentum and cyphellae on the lower surface and the often charac- teristic fishy odor caused by the presence of methylamine products (Galloway MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 262 Emilia Anna Ossowska et al.: ShortTitle 1994, 1997; Moncada 2012). However, species within this genus are much more difficult to distinguish, due to the lack of a clear concept of within-species varia- tion, including the type of photobiont, the presence and type of vegetative prop- agules, their shape, distribution, and size, as well as the width and length of the lobes or their shape (Moncada 2012). Molecular data have helped to address this issue and to refine the set of potentially diagnostic characters, and Moncada et al. (2014a) identified a total of over 150 morphological and anatomical features that should be taken into account (see also Ossowska et al. 2024a). Even so, accurate species recognition may be hindered by intraspecific vari- ability, such as the formation of photosymbiodemes, apotheciate vs. non-apoth- eciate species pairs, and discrete morphodemes (Moncada et al. 2020, 2021b; Ossowska et al. 2022a, b, 2024a; Di Meglio and Goward 2023). For instance, the two vegetatively reproducing S. fuliginosa (With.) Ach. (with isidia) and S. limbata (Sm.) Ach. (with soredia) cannot be distinguished using the ITS barcoding mark- er (Moncada et al. 2014a; Magain and Sérusiaux 2015). Another case is found in the large foliose S. filix (Sw.) Nyl. vs. the delicate S. lacera (Hook. f. & Taylor) Mull. Arg., both New Zealand endemics (Liicking et al. 2022), or in S. antoniana B. Moncada & Liicking and S. tomentosa (Sw.) Ach. from Hawaii (Moncada et al. 2020, 2021a), as well as S. arenosella Di Meglio & Goward and S. torii Ant. Simon & Goward (Simon et al. 2018a; Di Meglio and Goward 2023; Ossowska et al. 2024a, b). The Sticta fuliginosa clade also contains apotheciate specimens de- void of isidia or soredia, suggesting the existence of individuals within the same species with different modes of reproduction. This was also observed in our new species, S. kukwae, with apothecia in one specimen from Peru, arranged in a clade together with non-apotheciate specimens. Similar cases were recently re- ported for S. scabrosa B. Moncada, Merc.-Diaz & Bungartz subsp. scabrosa and S. cellulosa Kaasalainen, originally described as sterile (Moncada et al. 2021b; Kaasalainen et al. 2023), but later found fertile in material from Bolivia (Ossows- ka et al. 2022b, 2024b). Some species, e.g., the widely distributed S. andina, show even greater variation in reproduction modes: this species was previously divided into three tentative taxa that differed in the type of propagation (Moncada et al. 2021a, b; Ossowska et al. 2022a; Kaasalainen et al. 2023), but later they were merged into one species due to genetic similarities (Moncada et al. 2021b). Traditional taxonomy in Sticta was largely based on morphodemes, i.e., partic- ular gross morphologies that were recognized as species, e.g., narrow-lobed, cya- nobacterial individuals with marginal isidia as S. weigelii (Ach.) Vain., broad-lobed, cyanobacterial specimens with laminal isidia as S. fuliginosa, or green-algal, apoth- eciate individuals as S. canariensis (Bory) Bory ex Delise, S. damicornis (Sw.) Ach., or S. dichotoma Bory ex Delise. Molecular data have shown that these morphode- mes consist of many, often only distantly related species (Moncada et al. 2013b, 2014a, 2015; Lticking et al. 2021; Ossowska et al. 2024a). The two additional new species of Sticta introduced in this paper are also part of morphodemes: S. kuk- wae is morphologically similar to S. weigelii, both having a brown, irregular to orbic- ular thallus with dark, marginal vegetative propagules, whereas S. flakusiorum, due to its hairy upper surface, represents the S. humboldtii morphodeme. In recent years, research on Sticta has intensified, resulting in the addition of many new species in various parts of the world (e.g., Lendemer and Goffinet 2015; Magain and Sérusiaux 2015; Simon et al. 2018a, b; Dal Forno et al. 2018; Torres et al. 2021; Ossowska et al. 2022a, 2024a; Kaasalainen et al. 2023; MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 263 Emilia Anna Ossowska et al.: ShortTitle Di Meglio and Goward 2023; Yanez-Ayabaca et al. 2023). Many regions in the Neotropics, e.g., Colombia, Bolivia (Ossowska et al. 2024a) and Puerto Rico (Mercado-Diaz et al. 2020), but also Africa (Simon et al. 2018b; Kaasalainen et al. 2023) and Oceania, e.g., Hawaii (Moncada et al. 2020, 2021a), are being explored, but many other regions remain poorly studied in terms of the genus Sticta. For instance, the checklist of lichens from Peru includes only ten Sticta species (Ramos 2014), and these have not yet been critically restudied. Among the names in the Peruvian checklist are S. fuliginosa, S. laciniata Ach., S. sylvati- ca, and S. weigelii, in which many new species have recently been distinguished (Moncada et al. 2021b; Di Meglio and Goward 2023; Ossowska et al. 2024a; this study). The new species presented here, S. kukwae, and the as-yet-undescribed S. sp. 36, are the first specimens from Peru supported by molecular data. Taxonomy Sticta flakusiorum Ossowska, B. Moncada & Licking, sp. nov. MycoBank No: 856228 Fig: 2 Diagnosis. Differing from S. humboldtii in the absence of true cilia, the presence of submarginal apothecia with entire to crenate margins, completely to partly covered by white hairs, spongy to fasciculate primary tomentum, and scabrid basal membrane of cyphellae, white to yellow (then K+ purple), or partly brown. Type. BoLiviA. * Dept. La Paz; Prov. Bautista Saavedra, Area Natural de Manejo Integrado Nacional AROLOBAMBA, between La Curva and Charazani, 15°08'09"S, 69°02'03"W, 3780 m, open area with shrubs, Ceja de Monte Supe- rior (Altimontano), on shrub, 15 Nov. 2014, M. Kukwa 14677 (holotype UGDA L-65223, isotype LPB). Description. Stipe absent. Thallus orbicular, up to 5 cm diam., moderately branched, with 3-5 branches per 5 cm radius, branching pleurotomous to poly- tomous; lobes suborbicular to flabellate, interspaced to adjacent, involute, with their apices rounded, revolute, and undulate and their margins sinuous, slightly thickened; lobe internodes 2-20 mm long, 3-15 mm broad; thallus coriaceous. Upper surface pitted to rugose, yellowish brown to chocolate brown, darker near the apices in the herbarium, shiny; lobes entirely hirsute or rarely with some parts lacking tomentum, covered by white hairs, without papillae and macu- lae; true cilia absent, but lower tomentum partly projecting beyond the margins and resembling cilia, fasciculated to agglutinated, white to pale brown, up to 0.5 mm. Apothecia submarginal and laminal, subaggregated, sessile to shortly stipitate, with pronounced invagination on the lower side, up to 2.0 mm diam.; disc brown to chestnut-brown; margin entire to crenate, completely to partly covered by white hairs, up to 1 mm long, simple to agglutinated, margin brown to dark brown. Vegetative propagules absent. Lower surface ribbed, brown; primary tomentum dense and usually thick to sparse to the margin, spongy to fasciculate, soft, white to brown; secondary tomentum present, arachnoid. Rhizines absent. Cyphellae 1-20 per cm? towards the thallus center and 41- 60 per cm? towards the margin, scattered, elongate to irregular, urceolate with wide pore to cupuliform, erumpent to sessile, remaining below the level of the primary tomentum, with the margin raised and involute to erect, cream to brown MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 264 Emilia Anna Ossowska et al.: ShortTitle colored, with tomentum up to the pore; pore up to 1.5 mm diam.; basal mem- brane scabrid, white to yellow, or partly brown, when yellow K+ purple and C+ red-orange, KC-, P-. Medulla compact, white to yellow, or partly brown, when yellow K+ purple and C+ red-orange, KC-, P—. No substances detected by TLC. Upper cortex paraplectenchymatous, up to 35 um thick, uniform, up of 5 lay- ers of cells, their walls up to 1.5 um thick and their lumina rounded to isodia- metric, up to 5-15 x 5-10 um diam. Photobiont layer up to 150 um thick, its cells up to 10 um diam. Medulla up to 120 um thick, its hyphae up to 5.0 um broad. Lower cortex paraplectenchymatous, up to 50 um thick, with up to 7 cell layers; cells up to 10 um diam. Upper primary tomentum up to 100 um long, simple or in fascicles formed of up to 7 hyphae, hyphae simple. Upper sec- ondary tomentum not seen on upper surface. Lower primary tomentum up to 200 um long, composed of fascicles formed of 10-15 hyphae, hyphae mostly simple, apically free, and flexuous. Lower secondary tomentum 30 um long, of single, simple to branched hairs, moniliform. Cyphellae cavity up to 220 um deep; cells of basal membrane without or rarely with up to 2 papillae. Apothecia biatorine, up to 500 um high, with indistinct stipe, about 20 um high; excipulum up to 400 um broad, with projecting hairs, up to 1 um long. Hymenium up to 300 um high; epihymenium up to 5 um high, orange-brown, pigment present in the gel and in the walls upper cells of paraphyses, with very gelatinous upper layer. Asci 4—-8-spored, ascospores fusiform, 1—3-septate, 25-35 x 6-8 um. Habitat and distribution. Sticta flakusiorum is an epiphytic species found in an open area with shrubs at an altitude of 3780 m in the Department La Paz, Bolivia. Etymology. The species is named in honor of two lichenologists, Adam Flakus and Pamela Rodriguez-Flakus, for their contributions to the taxonomy of lichens and lichenicolous fungi of Bolivia. Notes. The new species, S. flakusiorum, forms part of the S. humboldtii morphodeme, which also includes S. pseudohumboldtii B. Moncada & Lucking and S. parahumboldtii B. Moncada & Licking (Moncada et al. 2013b). How- ever, unlike in the other species, the upper surface of S. flakusiorum is rather hirsute, while in S. humboldtii and the other species, the hairs are very dense and resemble the primary tomentum present on the lower surface (Moncada 2012). In addition, S. parahumboldtii has marginal soredia and lacks apothe- cia (Moncada et al. 2013b). Furthermore, all species differ in the color of the lower surface and tomentum. In the new species, the lower surface is brown, and the primary tomentum is white to cream. Other species have a cream-col- ored lower surface, and the primary tomentum is cream in S. parahumboldtii, cream-white in S. pseudohumboldtii, and cream to grey-brown in S. humboldtii (Moncada 2012; Moncada et al. 2013b). All species belong to clade | on the Sticta phylogeny (see Fig. 1), but the new species is more closely related to S. viviana. Sticta humboldtii and S. parahumboldtii are related to ‘S. arachnosyl- vatica’, while S. pseudohumboldtii is close to S. arachnofuliginosa B. Monca- da & Liicking (Widhelm et al. 2018). Among the species of this morphodeme, S. humboldtii has been reported more frequently than other species (Moncada 2012; Moncada et al. 2013b), including records from Peru (Ramos 2014). How- ever, only the Colombian records are supported by molecular data (Moncada et al. 2013b, 2014a), and therefore its presence in Peru needs to be verified. Sticta pseudohumboldtii and S. parahumboldtii are known so far only from Colombia (Moncada 2012; Moncada et al. 2013b, 2014a, b). MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 265 Emilia Anna Ossowska et al.: ShortTitle % : ea hs - tt ea ‘ By Pate ta 8 ie af Pe ate Figure 2. Morphology of Sticta flakusiorum (holotype) A upper surface B lower surface C, D hirsute upper surface with apothecium and lower surface with tomentum and cyphellae E apothecia with entire to crenate margins, covered by white hairs F primary tomentum spongy to fasciculate and cyphellae with scabrid basal membrane. Scale bars: 1 mm (A-F). In the phylogenetic tree, S. flakusiorum forms a lineage sister to a clade of a potentially new species, referred to as Sticta sp. 36 (see above). This taxon is distinguished by its thallus with smooth upper surface, sparse and laminal apothecia, and abundant, marginal phyllidia. Furthermore, the primary tomentum is greyish gold, whereas in S. flakusiorum it is white to brown. The specimens of S. sp. 36 are fragmentary; thus, we have decided not to describe it at this mo- ment. Sticta sp. 36 was found in Peru in Puno (Lampa, Santa Lucia). The hirsute upper surface is also characteristic of ‘S. arachnosylvatica’, S. minutula B. Moncada, A. Suarez & Licking and S. hirta (Nyl.) Trevis (Moncada MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 266 Emilia Anna Ossowska et al.: ShortTitle 2012; Moncada et al. 2014a, 2020), but these taxa differ from S. flakusiorum in the structure of the lobes, the presence of vegetative propagules, as well as the color of the lower surface and the structure of primary tomentum. In particular, the lobe margins in all these species are entire to crenate, whereas in S. flakusiorum they are sinusoidal; in addition, ‘S. arachnosylvatica’ and S. minutula have isidia. The lower surface of ‘S. arachnosylvatica’ is cream-white with primary tomentum dense to the margin (Moncada 2012), and in S. minutula the lower surface is cream-white with primary tomentum scarce over the whole area. Additionally, the latter taxon is dis- tinguished by its sparse cyphellae (Moncada 2012). Sticta hirta has a creamy lower surface with irregular tomentum, sparse towards the margins, and it is fasciculate to spongy (Moncada 2012). All three species, ‘S. arachnosylvatica; S. minutula, and S. hirta, have been molecularly confirmed only from Colombia (Moncada 2012; Moncada et al. 2014a, 2020) but have not been reported from Bolivia and Peru. Sticta kukwae Ossowska, Magain & Sérusiaux, sp. nov. MycoBank No: 856229 Fig. 3 Diagnosis. Differing from S. weigellii in lobes with sinuous margins, in the presence of marginal phyllidia, and the scarce, submarginal apothecia, as well as the primary tomentum being light brown to brown, dense, and sparse towards the margins. Type. BOLiviA. * Dept. La Paz; Prov. Franz Tamayo, Area Natural de Manejo Integrado Nacional APOLOBAMBA, between la Cumbre and Pelechuco, close to Aguas Blancas, 14°49'12"S, 69°07'05'W, elev. 4070 m, open high Andean vegetation, Altoandino, saxicolous, 15 Nov. 2014, M. Kukwa 14729a (holotype UGDA L-65224, isotype LPB). Description. Stipe absent. Thallus suborbicular to irregular, up to 10 cm diam., moderately branched, with 3-5 branches per 5 cm radius, branching anisotomous to polytomous; lobes ligulate to flabellate, undulate, with their apices rounded, revolute, and their margins sinuous, not thickened; lobe inter- nodes 5-9 mm long, 4-9 mm broad; thallus coriaceous. Upper surface smooth to shallowly pitted, yellowish brown to brown when dry, shiny; surface glabrous, few lobes with papillae but without maculae; true cilia absent. Only two apoth- ecia found, submarginal, with slightly pronounced invagination on lower side, up to 1.5 mm diam.; disc brown; margin smooth, brown to dark brown. Phyl- lidia present, marginal and laminal, simple, branched, palmate to corymbose, vertical to obliquely arranged, globular at first, then spatulate to squamiform, usually darker than the thallus. Lower surface uneven, light brown; primary to- mentum dense and thick to the margin, sometimes absent at the very edge, fasciculate to spongy, soft, white to brown, sometimes brown with brighter api- ces; secondary tomentum present, arachnoid. Rhizines present, only on few lobes, whitish to brown, simple to branched, densely distributed. Cyphellae 1-20 per cm? towards the thallus center and 1-20 per cm? towards the margin, dispersed, rounded to irregular, urceolate with wide pore, erumpent to sessile, remaining below the level of the primary tomentum, with the margin elevated and involute, white to beige colored, with tomentum; pore up to 0.5 mm diam.; basal membrane scabrid, white, K+ yellowish, C-, KC-, P-. Medulla compact, white, K-, C-, KC-, P—. No substances detected by TLC. MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 267 Emilia Anna Ossowska et al.: ShortTitle a 3 : 7 aR" Figure 3. Morphology of Sticta kukwae (A, B, E holotype C LG3223, D LG3221, F LG3227) A upper surface B lower surface C, D lobes with sinuous margins and marginal phyllidia E lower tomentum with cyphellae and rhizines. Scale bars: 1 mm (A-F). Upper cortex paraplectenchymatous, up to 65 um thick, uniform, consisting of up to 7 cell layers with cells 5-10 um diam., their walls up to 1.5 um thick. Photobiont layer up to 130 um thick, its cells up to 15 um diam. Medulla up to 120 um thick, its hyphae 4 um broad, without crystals. Lower cortex para- plectenchymatous, up to 60 thick, with 7 cell layers; cells up to 10 um diam., their walls up to 2.5 um thick. Lower primary tomentum up to 400 um long, with cells resembling secondary tomentum and probably representing thalloconidia, simple or in fascicles formed of up to 20 hyphae, hyphae simple. Lower sec- ondary tomentum 70 mm long, simple to branched, moniliform. Cyphellae cav- ity up to 150 um deep; cells of basal membrane without or with single papillae. MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 268 Emilia Anna Ossowska et al.: ShortTitle Apothecia lecanorine (with algal layer below cortex), up to 250 um high, without distinct stipe; excipulum 150 um broad, without projecting hairs. Hymenium up to 75 um high; epihymenium 5 um high, orange-brown with gelatinous upper layer. Asci immature. Ascospores not observed. Habitat and distribution. Sticta kukwae is known from Bolivia and Peru. In Bo- livia, it was found saxicolous and was collected at a single locality in the Area Natural de Manejo Integrado Nacional Apolobamba in the Department La Paz, at an altitude of 4070 m. In Peru, it was also saxicolous and found in four localities in Puno, in a vegetation type of Roquedal, Matorral de Puna, at an altitude of 3850 m. Etymology. Named in honor of the lichenologist Martin Kukwa for his contri- bution to the taxonomy of lichens and lichenicolous fungi in Bolivia. Additional material examined. PERU. * Puno - Carabaya, Ollachea - Macusani (20 km of Macusani), in a vegetation type of Roquedal, Matorral de Puna, on rocks on the ground/close to the ground, 23 May 2012, N. Magain (LG3225, LG3227, LG3221 & LG3223). Notes. Sticta kukwae is another species in the S. weigelii morphodeme, along with the recently described S. andina B. Moncada, Licking & Sérus., S. scabrosa, and S. waikamoi Moncada & Ltcking. It differs from these species in the type of vegetative propagules and the presence of lobes with strongly sinuous margins, which have not been observed in the other species. Sticta weigelii s.str. and S. waikamoi produce isidia, and S. andina has isidia and phyllidia. Sticta scabrosa, as S. kukwae, produces phyllidia, but in this taxon, they are the same color as the thallus, whereas in the new species, they are blackish-brown. Both species can produce sparse apothecia, but in S. kukwae their margins are crenate and dark brown, whereas in S. scabrosa they are entire to very rarely shallowly crenate and in the same color as the thallus (Moncada et al. 2021b; Ossowska et al. 2022b). Sticta andina may also have apothecia, but they are abundant and with verrucose to crenate margins (Moncada et al. 2021a, b; Ossowska et al. 2022b). Another difference is found in the color of the lower surface, as S. andina has dark lower surface, in S. scabrosa it is yellow-brown, while in S. weigelii the color ranges from beige to dark brown, and in S. waikamoi it is dark brown (Moncada et al. 2020, 2021a, b; Ossowska et al. 2022b). The newly described species has a light brown lower surface. Sticta andina and S. scabrosa have a wide distribution (Moncada et al. 2021a, b; Kaasalainen et al. 2023). In contrast, S. weigelii was previously as- sumed to be widespread (Galloway 1994, 1997, 2006). However, recent research has shown that its distribution is probably limited to the Neotropics (Moncada et al. 2021b; Mercado-Diaz et al. 2023). All three taxa are also known from Bolivia (Ossowska 2021; Ossowska et al. 2022b). Sticta waikamoi is known from the Ha- waiian islands (Moncada et al. 2020, 2021a). Only S. weigelii has been reported from Peru (Ramos 2014), but without molecular evidence. In the phylogenetic tree (Fig. 1), the new species is closely related to S. um- bilicariiformis. However, it has many marginal pustules, which can sometimes make it appear sorediate; thalli is often quite large, and lobes are thick with wavy to foveolate margins. Additionally, the lower surface is cream-colored to brown and thickly tomentose. Sticta umbilicariiformis has been documented in East Africa, with a high probability of its occurrence in other regions as well (Magain and Sérusiaux 2015; Kaasalainen et al. 2023). The presence of lobes with sinuous margins is also a characteristic feature in the recently distinguished S. monlueckiorum Ossowska, Flakus & Rodr.-Flakus MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 269 Emilia Anna Ossowska et al.: ShortTitle from Bolivia. In S. monlueckiorum, the thallus is larger (up to 10 cm) and moder- ately branched, while the apothecia are laminal with hirsute margins and with- out vegetative propagules (Crous et al. 2023), whereas S. flakusiorum has a hirsute upper surface with abundant, submarginal apothecia and without vege- tative propagules. All three taxa differ also in the color of the lower surface and the density of the cyphellae. In S. monlueckiorum, the lower surface is beige to yellowish, and the cyphellae have a density of 41-60 per cm? towards the center and more than 100 towards the margins (Crous et al. 2023). In S. flakus- iorum, the lower surface is brown, and the cyphellae are 1-20 per cm? towards the center and 41-60 per cm? towards the margins, and in S. kukwae, 1-20 per cm? towards the thallus center and margins. The hyphae of primary tomentum of Sticta kukwae produce peculiar struc- tures that resemble budding conidia forming chains. Similar structures were found in the isidiate S. atlantica Magain & Sérus., S. fuliginoides Magain & Sérus., and S. fuliginosa by Magain and Sérusiaux (2015), who stated in the case of S. fuliginosa they can act as conidia. These cells in the mentioned spe- cies are very similar to cells of secondary tomentum in several Sticta species, and possibly both can play a role of conidia. Such spores thus can be named thalloconidia, which are on the other hand known mainly in several species of the genus Umbilicaria Hoffm. (Hestmark 1990, 1991, 1992), but also in some crustose lichens (e.g., Miriquidica nephaea (Sommerf.) P.F. Cannon, Protopar- melia leproloma (R. Sant.) Rambold & Poelt, Protoparmeliopsis peltata (DC.) Arup, Zhao Xin & Lumbsch, Rhizoplaca melanophthalma (DC.) Leuckert & Poelt, Sporastatia karakorina (Obermayer & Poelt) Davydov & Yakovch.) (Poelt and Obermayer 1990). However, the ultrastructural study of their development must be performed prior to the final change in the conception of their role. Acknowledgements We are grateful to the members of Herbario Nacional de Bolivia, Instituto of Herbario Nacional de Bolivia, Instituto de Ecologia, Universidad Mayor de San Andrés, La Paz, for the generous cooperation. We are also greatly indebted to Joseph Di Meglio for his helpful comments. This research received support from the SYNTHESYS Project (DE-TAF-8180) http://www.synthesys.info/, which is financed by the European Community Research Infrastructure Action under the FP7 “Capacities” Programme and the University of Gdansk, granted to EAO. Lichen samples were collected in Bolivia with the permission of the Ministerio de Media Ambiente y Agua and in cooperation with Herbario Nacio- nal de Bolivia (LPB), who in turn made specimens available to the Herbarium of the University of Gdansk and W. Szafer Institute of Botany, Polish Academy of Sciences in Krakow. We thank Eimy Rivas Plata, Daniel Ramos, Francois Lu- tzoni, and Jolanta Miadlikowska for organizing the field trip in Peru, which was funded by the National Science Foundation award DEB-1025930 REVSYS. Additional information Conflict of interest The authors have declared that no competing interests exist. MycoKeys 114: 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 270 Emilia Anna Ossowska et al.: ShortTitle Ethical statement No ethical statement was reported. Funding This research received support from the SYNTHESYS Project (DE-TAF-8180) http:// www.synthesys.info/, which is financed by the European Community Research Infra- structure Action under the FP7 “Capacities” Programme and the University of Gdansk, granted to EAO. The field trip in Peru was funded by the National Science Foundation award DEB-1025930 REVSYS. Laboratory work at the BGBM was partially supported by the Verein der Freunde des Botanischen Gartens. Funding for field and laboratory work to assemble the phylogenetic backbone for the genus Sticta was provided by the National Science Foundation award DEB-1354884 ("Collaborative Research: Evolution, Diversification, and Conservation of a Megadiverse Flagship Lichen Genus’). Author contributions Emilia Anna Ossowska: conceptualization, descriptions of new species, determination of species, molecular laboratory work and analyses, chromatographic analyses, manu- script writing, and editing; Bibiana Moncada: descriptions of new species, phylogenetic analyses, manuscript editing; Robert Lucking: phylogenetic analyses, manuscript writ- ing, and editing; Emmanuel Sérusiaux: molecular laboratory work and analyses, manu- script editing; Nicolas Magain: material collecting, molecular laboratory work and anal- yses, manuscript editing. Author ORCIDs Emilia Anna Ossowska ® https://orcid.org/0000-0002-1357-6071 Bibiana Moncada ® https://orcid.org/0000-0001-9984-2918 Robert Lticking © https://orcid.org/0000-0002-3431-4636 Emmanuel Sérusiaux © https://orcid.org/0000-0002-0456-0131 Nicolas Magain © https://orcid.org/0000-0001-5409-9518 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. References Aptroot A (2002) New and interesting lichens and lichenicolous fungi in Brazil. Fungal Diversity 9: 15-45. Consortium of Lichen Herbaria (2023) Consortium of Lichen Herbaria — building a Glob- al Consortium of Bryophytes and Lichens as keystones of cryptobiotic communities. 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Lichenologist (London, England) 55(5): 203-222. https://doi. org/10.1017/S0024282923000476 Supplementary material 1 Specimens of Sticta used in molecular analysis with locality, voucher information, GenBank accession numbers and list of references Authors: Emilia Anna Ossowska, Bibiana Moncada, Robert Licking, Emmanuel Sérusiaux, Nicolas Magain Data type: xlsx Explanation note: 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.114.139681.suppl1 MycoKeys 114; 259-276 (2025), DOI: 10.3897/mycokeys.114.139681 276