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Tephromela vinacea L.A.Santos, Aptroot, Lücking & M.Cáceres, sp. nov. (Figs 3E, F; 4C, D) Similar to Tephromela atra but corticolous and differing in the presence of a fatty acid, and in the wine-red pigment in the upper hymenium and epihymenium. HOLOTYPE. — Brazil. Pernambuco, Buíque, Parque Nacional Vale do Catimbau, alt. 745 m, 8°30’41”S, 37°16’37”W, on bark of tree, 13-15.VIII.2017, L.A. Santos s.n. (holo-, ISE [ISE46534]). ADDITIONAL SPECIMENS EXAMINED. — Brazil. Pernambuco, Buíque, Parque Nacional Vale do Catimbau, alt. 745 m, 8°29’49”S, 37°18’02”W, on bark of tree, 13-15.VIII.2017, L.A. Santos s.n. (ISE [ISE54172]); Mato Grosso do Sul, Fazenda Marambaia, Bonito, alt. 650 m, 20°58’S, 56°42’W, on wood, 30.X.2018, A. Aptroot 77088 (ABL, CGMS); Fazenda Santa Fé, Jardim, alt. 650 m, 21°32’S, 56°45’W, on bark of tree, 02.XI.2018, A. Aptroot 77478 (ABL, CGMS); Bonito, outskirts, near tower, alt. 475 m, 21°07’44”S, 56°30’41”W, on bark of tree, 09.XI.2018, A. Aptroot 78049 (CGMS). ETYMOLOGY. — The species is named after the wine-red pigmented gel that surrounds the paraphyses. CHEMISTRY. — Apothecia and medulla UV+ greenish white, C-, P-, K-. Alectoronic acid, atranorin and an unknown hydrophobic substance detected by TLC. This unidentified fatty acid has an RF value of 18 in solvent C. ECOLOGY AND DISTRIBUTION. — On exposed tree branch in Caatinga vegetation in NE Brazil and on trees and especially wooden poles in Cerrado in S Brazil; known only from Brazil, but probably occurring in adjacent countries. MYCOBANK. — MB 846316. GENBANK. — OP881896. DESCRIPTION Thallus crustose, whitish to pale ochraceous-grey, rimose to areolate, minutely verrucose, somewhat glossy, c. 0.1 mm thick. Prothallus black to greyish-white or absent. Photobiont an unicellular green alga, arranged in one layer below the cortex, algal layer 37-70 µm high. Apothecia always present, sessile, roundish, homogeneously distributed on the thallus, with somewhat unevenly crenate, sometimes lobate margin. Disk black, 0.5-2 mm in diam., flat to slightly convex, epruinose. Thalline margin concolorous with the thallus, c. 0.3 mm wide, raised above the level of the disk (100-125 µm). Proper exciple thickest at the base, becoming thinner at the edges, pale to intense yellow-brown, intensifying yellow in K. Exciple with crystals that after treatment with K partly or completely dissolve. Hymenium c. 63-100 µm thick, pale wine red. Epihymenium dark wine red, K+ intensifying wine red. Paraphyses simple, septate, 2-3 µm wide, with 4-6 µm wide hyaline apical cells, surrounded by wine red pigmented gel. Subhymenium 37.5-50 µm thick, brown. Hypothecium 62-87.5 µm thick, golden brown. Asci slightly clavate, 40-55 µm tall, 8-spored. Ascospores simple, hyaline, ellipsoid, 10-13 ×5-8 µm. Pycnidia black. Conidia rod-shape, 12.5-15 × 1 µm. DISCUSSION The new species superficially resembles Tephromela atra, but differs in secondary chemistry and is not closely related to the latter phylogenetically (Fig. 2). Fatty acids were first reported for the genus in Muggia et al. (2014). At the time, the authors did not describe the material as a new taxon and questioned the application of chemical characters in the delimitation of species within the genus. Subsequently, based on the presence of fatty acid, Cestaro et al. (2016) described T. pacifica. Kantvilas (2015) and Kantvilas & Elix (2017) also recently used chemical patterns to describe new species with unique chemistry. The metabolite found in our species is a fatty acid, as reported by Cestaro et al. 2016, but has a chemical standard, with a RF value of 18 in solvent C, different from that found by Cestaro et al. (2016) with a RF value of 50. Most similar to the new species are T. alectoronica, described from Australia but in the protologue also reported from Brazil, and T. rhizophorae Kalb, described from Brazil. Both lack fatty acids but contain traces of physodalic acid instead. The ascospores of T.rhizophorae are distinctly smaller (8-10×5-6 µm) whereas those of T. alectoronica are only slightly larger (12- 17 ×6-9 µm). Thus, without co-chromatography, T. vinacea L.A.Santos, Aptroot, Lücking & M.Cáceres, sp. nov. and T. alectoronica are difficult to distinguish. The specimen clustering as sister to T. vinacea L.A.Santos, Aptroot, Lücking & M.Cáceres, sp. nov., and here named T. aff. vinacea (Fig. 2) is of interest in this respect. It presents a UV + greenish white fluorescence, although alectoronic acid was not clearly detectable with TLC. It was collected at one of the localities where Kalb (2008) originally reported material identified as T. alectoronica (Chapada dos Guimarães, Mato Grosso). Thus, Kalb’s material might be conspecific with this sequenced specimen, which would imply that the Brazilian material of T. alectoronica is not that species but is represented by the clade here named T. aff. vinacea, quite distant from the Australian T. alectoronica (Fig. 2). However, given that the underlying specimen is rather small and the chemistry could not be determined with certainty, we refrain from any formal conclusions in this case., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on pages 18-20, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402, {"references":["MUGGIA L., PEREZ- ORTEGA S., FRYDAY A., SPRIBILLE T. & GRUBE M. 2014. - Global assessment of genetic variation and phenotypic plasticity in the lichen-forming species Tephromela atra. Fungal Diversity 64: 233 - 251. https: // doi. org / 10.1007 / s 13225 - 013 - 0271 - 4","CESTARO L., TONSBERG T. & MUGGIA L. 2016. - Phylogenetic data and chemical traits characterize a new species in the lichen genus Tephromela. Herzogia 29 (2): 383 - 402. https: // doi. org / 10.13158 / heia. 29.2.2016.383","KANTVILAS G. 2015. - Observations on the genus Tephromela (lichenised Ascomycetes) in Tasmania, with the description of a new species. Herzogia 28: 430 - 444. https: // doi. org / 10.13158 / heia. 28.2.2015.430","KANTVILAS G. & ELIX J. A. 2017. - Tephromela baudiniana sp. nov. (lichenised Ascomycetes) from Kangaroo Island. Swainsona 31: 27 - 30.","KALB K. 2008. - New or otherwise interesting Lichens. IV. Sauteria 15: 239 - 248."]}

Tephromela multireflexa Aptroot & M.F.Souza, sp. nov. (Fig. 3A, B) Similar to Tephromela atra, but corticolous and differing in the apothecium thalline margin containing lichexanthone in the cortex and α- collatolic acid in the medulla. HOLOTYPE. — Brazil. Mato Grosso, Cuiabá, Chapada dos Guimarães, Pousada do Parque private area, alt. 700 m, 15°26’50”S, 55°49’50”W, on bark, 12-19.IX.2020, A. Aptroot & M.F. Souza 81950 (holo-, CGMS; iso-, ABL). ETYMOLOGY. — The name refers to the two different UV-reflectant colours. CHEMISTRY. — Thallus UV-, C-, P-, K-, apothecium margin UV+ yellow, apothecium medulla UV+ bluish-green. With lichexanthone (apothecial margins) and α- collatolic acid detected by TLC. ECOLOGY AND DISTRIBUTION. — On exposed tree branch in Cerrado forest; known only from the type collection from Brazil. MYCOBANK. — MB 846314. DESCRIPTION Thallus crustose, mineral to greenish-grey, rimose to areolate, minutely verrucose, somewhat glossy, c. 0.3 mm thick, areoles 0.05-0.2 mm in diam. Prothallus absent. Photobiont an unicellular green alga, arranged in one layer below the cortex. Apothecia present, sessile, roundish to lobate, homogeneously distributed on the thallus. Disk black, 0.5- 2.5 mm in diam., flat to slightly convex, epruinose. Thalline margin concolorous with the thallus, glossy, c. 0.3 mm wide, raised above the level of the disk, minutely crenate. Proper exciple thickest at the base, becoming thinner at the edges, hyaline. Exciple with crystals that after treatment with K partly dissolve. Hymenium c. 90-180 µm thick, blue, colour in K unchanged. Subhymenium 50-70 µm thick, dark blue, in K dark purple. Epihymenium dark purple, pigment in K dissolving leaving the blue of the hymenium. Paraphyses simple, septate, 2-3 µm wide, with 4-6 µm wide hyaline apical cells, surrounded by lilac pigmented gel. Hypothecium 0-80 µm thick, pale yellow, intensifying orange yellow in K. Asci slightly clavate, 70-55 µm tall, 8-spored. Ascospores simple, hyaline, ellipsoid, 14-16×5-6.5 µm, wall 1 µm thick. Pycnidia not observed. DISCUSSION The substance α- collatolic acid is commonly occurring in the genus Tephromela, while lichexanthone is so far only reported from T. velloziae. In that species, and in two further species described below, lichexanthone occurs in the thallus and the apothecium margins, not only in the thallus. The new species is the first in the genus that combines both lichexanthone and α- collatolic acid., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on pages 16-18, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402

Tephromela xanthonica Guzmán-Guillermo, sp. nov. (Figs 3G, H; 4E, F) Similar to Tephromela velloziae, differing in its saxicolous ecology and its areolate thallus. HOLOTYPE. — Mexico. Veracruz, Municipality of Las Vigas de Ramírez, locality of Volcancillo, alt. 2300 m, 19°37’00.5”N, 97°04’01.0”W, on volcanic rock, IX.2020, Cárdenas-Mendoza s.n. (holo-, XALU [XALU1413]; para-, XALU [XALU1414, XALU1415, XALU1416, XALU1417]). ADDITIONAL SPECIMENS EXAMINED. — Brazil. Minas Gerais, Catas Altas, Caraça, alt. 1200-1400 m, 20°06’S, 43°29’W, on sandstone, 17-25.V.2021, L.A. Santos & Aptroot 52016, 52057 (ISE, CGMS). ETYMOLOGY. — The name refers to its chemistry. CHEMISTRY. — Thallus UV+ yellow, C-, P-, K-, apothecium margin UV+ yellow. With lichexanthone. ECOLOGY AND DISTRIBUTION. — On exposed volcanic rock associated with Pinus forest in Volcancillo, Veracruz, and on sandstone in Campo rupestre in Caraça, Minas Gerais, Brazil; known from Brazil and Mexico. MYCOBANK. — MB 846317. DESCRIPTION Thallus crustose, mineral to greenish-grey, rimose to verrucose, somewhat glossy, c. 0.3 mm thick, areoles 0.5-1 mm in diam. Prothallus absent. Photobiont an unicellular green alga, arranged in one layer below the cortex. Apothecia present, sessile, roundish to lobate when old, numerous on the center of the thallus. Disk black, 0.5-2.5 mm in diam., flat to slightly convex, epruinose. Thalline margin concolorous with the thallus, glossy, 0.15-0.18 mm wide, raised above the level of the disk, minutely crenate, with motes of white pruina when young. Proper exciple thickest at the base (50-60 µm), becoming thinner at the edges, intense yellowbrown. Hymenium c. 100-130 µm thick, pigment reddish purple, K+ purple. Subhymenium 80-90 µm thick, purple pigmented, dark purple in K. Epihymenium reddish-purple, pigment K+ purple. Paraphyses simple, septate, surrounded by dark red pigmented gel. Hypothecium 0-60 µm thick, with similar colorations as the hymenium. Asci slightly clavate, 65-55 µm tall, 8-spored. Ascospores simple, hyaline, ellipsoid, 9-13 × 5-6 µm, wall 1-2 µm thick. Pycnidia not observed. DISCUSSION Tephromela velloziae was described as containing lichexanthone and it also agrees in ascospore size (Kalb 1984). However, the isotypes Kalb 349 (B[B600157687, B600093712]) deposited in Herbarium Berolinense (digital material consulted in BiNHum 2021) and mentioned in the original description (Kalb 1984), differ morphologically and in ecology from the new species. They have a verrucose thallus without areoles, in contrast to the areolate thallus of T. xanthonica Guzmán-Guillermo, sp. nov. Also, T. velloziae grows on dead stems of a semi-woody herb in semi-arid Caatinga vegetation, whereas T. xanthonica Guzmán-Guillermo, sp. nov., is a saxicolous species, found growing on sandstone and on volcanic rock between species of Cladonia, Stereocaulon, Rhizocarpon, and other unidentified crustose lichens, in temperate to tropical montane ecosystem, including Pinus forest at 2250-2300 m in Mexico and campos rupestres at 1200-1400 m in Brazil., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on pages 20-21, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402, {"references":["KALB K. 1984. - Lichenes Neotropici ausgegeben von Klaus Kalb. Fasc. VIII (nos 301 - 350). Published by the author, Neumarkt / Opf.","BINHUM. 2021. - Biodiversitatsnetzwerk des Humboldt-Rings. Available athttps: // www. binhum. net / (accessed 13 April 2021)."]}

KEY TO SPECIES OF TEPHROMELA M.CHOISY IN BRAZIL The Brazilian material keying out here possibly does not represent T. alectoronica s.str. but perhaps T. aff. vinacea in our phylogenetic tree. 1. Saxicolous or muscicolous on rock or soil or lichenicolous on saxicolous Lecanora...................................... 2 — Corticolous or on Vellozia stems or lichenicolous on corticolous Dirinaria................................................ 10 2. Lichenicolous on saxicolous Lecanora........................................... T. campestricola (Nyl.) Rambold & Triebel — Saxicolous or muscicolous on rock or on soil............................................................................................... 3 3. Apothecia with constricted base; medulla UV+ greenish-white (alectoronic and α- collatolic acids) or medulla UV+ yellow (lichexanthone)........................................................................................................................ 4 — Apothecia margin not distinguishable from raised thallus parts (apothecia seemingly emarginate or immersed)................................................................................................................................................... 7 4. Medulla UV+ greenish-white (alectoronic and α- collatolic acid).............................. T. atra (Huds.) Hafellner — Medulla UV-negative; thallus and apothecium margin UV+ yellow (lichexanthone)................................... 5 5. Hymenium, epihymenium and subhymenium all equally reddish purple............................................................................................................................................................ T. xanthonica Guzmán-Guillermo, sp. nov. — Subhymenium reddish; epiphymenium bluish; hymenium hyaline or mottled bluish and brownish........... 6 6. Subhymenium reddish brown; epiphymenium and hymenium mottled bluish and brownish............................................................................................................... T. carassensis (Vain.) Aptroot & Lücking, comb. nov. — Subhymenium dark orangish brown; epiphymenium dark blue to black; hymenium hyaline............................................................................................................. T. epichlorina (Vain.) Aptroot & Lücking, comb. nov. 7. Apothecia (sub)convex, almost without thalline margin; thallus with black hypothallus visible between the areoles......................................................................................................................................................... 9 — Apothecia more or less flat, in raised thallus warts; thallus continuous to rimose......................................... 7 8. Ascospores 10-14 × 6-8 µm; medulla UV+ bluish-green (α- collatolic acid).............................................................................................................................................................................. T. buelliana (Müll. Arg.) Kalb — Ascospores 10-11 × 6-7 µm; medulla UV- (colensoic acid)................................................................................................................................................................................................... T. colensoica Rambold & Knoph 9. Ascospores 7-10 × 3.5-4.5 µm; medulla UV+ white (loxodellonic and glomelliferic acids)........................................................................................................................................................ T. matogrossensis Kalb & Elix — Ascospores 10-12 × 8-9 µm; medulla UV+ greenish-white (alectoronic and physodic acids).............................................................................................................................................................. T. immersa Kalb & Elix 10. Lichenicolous on corticolous Dirinaria...................................... T. cerasina (Müll. Arg.) Rambold & Triebel — Corticolous............................................................................................................................................... 11 11. Thallus UV-, but apothecial margins may be UV+ yellow; medulla UV- or UV+ greenish-white or UV+ bluishgreen......................................................................................................................................................... 12 — Thallus UV+ yellow (lichexanthone); ascospores 9-16 × 5-7.5 µm.............................................................. 16 12. Medulla UV- (no substances in medulla); ascospores long ellipsoid, 10-12 × 3-4 µm........................................................................................................................................................................ T. americana (Fée) Kalb — Medulla UV+ greenish-white or UV+ bluish-green; ascospores ellipsoid, at least 5 µm wide...................... 13 13. Apothecial margin UV+ yellow (lichexanthone), medulla UV+ bluish green (α- collatolic acid); fatty acids absend; ascospores 14-16 × 5-6.5 µm.................................................... T. multireflexa Aptroot & M.F.Souza, sp. nov. — Apothecial margin UV- or (medulla) UV+ greenish-white (alectoronic and/or physodic acids); fatty acids (TLC) present or absent....................................................................................................................................... 14 14. Ascospores 8-10 × 5-6 µm; in mangroves.......................................................................... T. rhizophorae Kalb — Ascospores 10-17 × 5-9 µm; in inland ecosystems...................................................................................... 15 15. Fatty acids present (TLC); ascospores 10-13 × 5-8 µm......................................................................................................................................................... T. vinacea L.A.Santos, Aptroot, Lücking & M.Cáceres, sp. nov. — Fatty acids absent (TLC); ascospores 12-17 × 6-9 µm.............................................. T. alectoronica Kalb s.lat. 16. On Vellozia stems; ascospores 9-12 × 5-7 µm........................................................................ T. velloziae Kalb — On tree bark; ascospores 14-16 × 6.5-7.5 µm........................................................................................................................................................................... T. obesimarginata L.A.Santos, Aptroot & M.Cáceres, sp. nov., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on page 23, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402

Tephromela obesimarginata L.A.Santos, Aptroot & M.Cáceres, sp. nov. (Figs 3C, D; 4A, B) Similar to Tephromela velloziae but corticolous and differing in the unevenly thickened thalline apothecium margin and the larger ascospores. HOLOTYPE. — Brazil. Minas Gerais, Catas Altas, Caraça, alt. 1200- 1400 m, 20°06’S, 43°29’W, on tree bark, 17-25.V.2021, L.A. Santos & A. Aptroot s.n. (holo-, ISE [ISE52321]; iso-, CGMS). ADDITIONAL SPECIMENS EXAMINED. — Brazil. Minas Gerais, Catas Altas, Caraça, alt. 1200-1400 m, 20°06’S, 43°29’W, on tree bark, 17-25.V.2021, L.A. Santos & A. Aptroot 52143, 52312, 52309 (ISE, CGMS). ETYMOLOGY. — The species is named after the irregularly thickened, plump, thalline apothecium margin. CHEMISTRY. — Thallus UV+ yellow, C-, P-, K-, apothecium margin UV+ yellow. With lichexanthone. ECOLOGY AND DISTRIBUTION. — On exposed trees in Campo rupestre vegetation in SE Brazil; known only from Brazil. MYCOBANK. — MB 846315. GENBANK. — OP881898. DESCRIPTION Thallus crustose, whitish to pale ochraceous-grey, rimose to areolate, verrucose,somewhat glossy, c. 0.1 mm thick.Prothallus absent. Photobiont an unicellular green alga, arranged in one layer below the cortex, algal layer c. 50 µm high. Apothecia always present, sessile, roundish, homogeneously distributed on the thallus, with unevenly crenate, irregularly thickened margin. Disk black, 0.7-2 mm in diam., flat to somewhat concave or saddle-shaped, epruinose. Thalline margin concolorous with the thallus, c. 0.2-0.6 mm wide, raised above the level of the disk (100-150 µm). Proper exciple thickest at the base, becoming thinner at the edges, pale to intense yellow-brown, intensifying yellow in K. Exciple with crystals that after treatment with K partly or completely dissolve. Hymenium c. 100-150 µm thick, violet, K+ greenish-grey. Epihymenium violet, K+ greenish-grey. Paraphyses simple, septate, 2-3 µm wide, with 3-5 µm wide hyaline apical cells, surrounded by violet pigmented gel.Hypothecium 100-150 µm thick, violet, K+ greenish-grey, with a narrow golden-brown band below the hymenium (25-50 µm thick). Asci slightly clavate, 50-75 µm tall, 8-spored. Ascospores simple, hyaline, ellipsoid, 14-16 × 6.5-7.5 µm. Pycnidia black. Conidia not observed. DISCUSSION The new species somewhat resembles Tephromela velloziae, but differs in the irregularly thickened apothecium margin, the larger ascospores (14-16×6.5-7.5 µm vs 9-12× 5-7 µm), and the corticolous habit., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on page 18, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402

Tephromela epichlorina (Vain.) Aptroot & Lücking, comb. nov. (Fig. 5C, D) Lecanora epichlorina Vain., Acta Societatis pro Fauna et Flora Fennica 7 (1): 91 (Vainio 1890). HOLOTYPE. — Brazil. Minas Gerais, Catas Altas, Caraça, alt. 1400- 1500 m, on moss on sandstone, 1885, E. Vainio distributed in Lichenes Brasilinses Exsiccati 164 (holo-, TUR [TUR-VAIN 5645]). ADDITIONAL SPECIMEN EXAMINED. — Brazil. Minas Gerais, Catas Altas, Caraça, on sandstone, 1997, Aptroot 41012 (ABL, SP). CHEMISTRY. — Thallus UV+ yellow, C-, P-, K-, apothecium margin UV+ yellow. With lichexanthone. ECOLOGY AND DISTRIBUTION. — On sandstone in Campo rupestre in Caraça, Minas Gerais, Brazil; known only from Brazil. MYCOBANK. — MB 846230. DESCRIPTION Thallus crustose, mineral to greenish-grey, rimose to verrucose, somewhat glossy, c. 0.3 mm thick, areoles 0.5-1 mm in diam. Prothallus absent. Apothecia present, sessile, roundish to lobate when old, numerous on the center of the thallus. Disk black, 0.5-1.5 mm in diam., flat, epruinose. Thalline margin concolorous with the thallus, glossy, 0.15-0.2 mm wide, raised above the level of the disk, crenate. Hymenium c. 100-150 µm thick, hyaline. Subhymenium c. 200 µm thick, dark orange brown. Epihymenium dark blue to black, c. 60 µm thick, pigment unchanged in K. Asci slightly clavate, 8-spored. Ascospores simple, hyaline, ellipsoid, 9-15 ×5-6 µm, wall 1-2 µm thick. Pycnidia not observed. DISCUSSION This species was described in Lecanora but on account of the anatomical and chemical characters belongs in Tephromela, as also suggested by an annotation label by Klaus Kalb. For differences with T. carassensis (Vain.) Aptroot & Lücking, comb. nov. and T. xanthonica Guzmán-Guillermo, sp. nov., see Discussion under T. carassensis (Vain.) Aptroot & Lücking, comb. nov. and in the key below., Published as part of Santos, Lidiane Alves dos, Aptroot, André, Souza, Maria Fernanda de, Lücking, Robert, Guzmán-Guillermo, Jorge & Cáceres, Marcela Eugenia da Silva, 2023, Four new species of Tephromela M. Choisy (Ascomycota, Tephromelataceae), three containing lichexanthone, from Brazil and Mexico, pp. 11-25 in Cryptogamie, Mycologie 20 (2) on page 22, DOI: 10.5252/cryptogamie-mycologie2023v44a2, http://zenodo.org/record/7815402, {"references":["VAINIO E. A. 1890. - Etude sur la classification et la morphologie des lichens du Bresil. I. Acta Societatis pro Fauna et Flora Fennica 7 (1): 1 - 247."]}

Tephromela carassensis (Vain.) Aptroot & Lücking, comb. nov. (Fig. 5A, B) Lecanora carassensis Vain., Acta Societatis pro Fauna et Flora Fennica 7 (1): 91 (Vainio 1890). HOLOTYPE. — Brazil. Minas Gerais, Catas Altas, Caraça, alt. 1400- 1500 m, on moss on sandstone, 1885, E. Vainio distributed in Lichenes Brasilinses Exsiccati 1572 (holo-, TUR [TUR-VAIN 5217]). ADDITIONAL SPECIMEN EXAMINED. — Brazil. Minas Gerais, Catas Altas, Caraça, on sandstone, 1997, Aptroot 40719 (ABL, SP). CHEMISTRY. — Thallus UV+ yellow, C-, P-, K-, apothecium margin UV+ yellow. With lichexanthone. ECOLOGY AND DISTRIBUTION. — On mosses on sandstone in Campo rupestre in Caraça, Minas Gerais, Brazil; known only from Brazil. MYCOBANK. — MB 846318. DESCRIPTION Thallus crustose, mineral to greenish-grey, rimose to verrucose, somewhat glossy, c. 0.3 mm thick, areoles 0.5-1 mm in diam. Prothallus absent. Apothecia present, sessile, roundish to lobate when old, numerous on the center of the thallus. Disk black, 0.5-1.5 mm in diam., flat, epruinose. Thalline margin concolorous with the thallus, internally with small crystals, glossy, 0.15-0.18 mm wide, raised above the level of the disk, crenate. Hymenium c. 100 µm thick, mottled bluish grey and pale brown. Subhymenium c. 200 µm thick, dark reddish brown. Epihymenium bluish grey, c. 60 µm thick, pigment unchanged in K. Asci slightly clavate, 8-spored. Ascospores simple, hyaline, ellipsoid, 9-12×5-6 µm, wall 1-2 µm thick. Pycnidia not observed. DISCUSSION This species was described in Lecanora but on account of the anatomical and chemical characters belongs in Tephromela, as also suggested by an annotation label by Klaus Kalb. It agrees in many aspects with T. epichlorina (Vain.) Aptroot & Lücking, comb. nov. (see below), and both were described from the same locality, but there are notably differences in the pigmentation patterns of the hymenium, epihymenium, and subhymenium. Both species differ from T. xanthonica Guzmán-Guillermo, sp. nov. also largely in apothecial pigment patterns (see key below).

Dirina paradoxa, Heterocyphelium triseptatum, Julella sublactea, Opegrapha aurantiaca, Peltula steppae, Sarcographina cyclospora y Schistophoron tenue son reportadas por primera vez para Colombia. Los géneros Dirina, Heterocyphelium Sarcographina y Schistophoron son reportados por primera vez para Colombia. Estos líquenes provienen principalmente de ecosistemas secos donde colecciones previas de hongos liquenizados han sido escasas. Dirina paradoxa, Heterocyphelium triseptatum, Julella sublactea, Opegrapha aurantiaca, Peltula steppae, Sarcographina cyclospora and Schistophoron tenue are reported for the first time in Colombia. The genera Dirina, Heterocyphelium Sarcographina and Schistophoron are reported for the first time in Colombia. These lichens mainly originate from dry ecosystems where previous collections of lichenized fungi have been uncommon.

Fissurina linoana Lücking, Moncada & G. Rodr. sp. nov. Mycobank MB 828114 (Fig. 3a). Diagnosis. Differing from Fissurina alligatorensis in the I-negative ascospores with thin walls and septa. Type. COLOMBIA. Atlántico: Piojó, Reserva Forestal Protectora (RFP) El Palomar; 10º45’47” N, 75º08’56” W, 200–600 m; 18 March 2016, R. Lücking, B. Moncada et al. 42977b (UNO, holotype). Description. Thallus corticolous, up to 2 cm diam., continuous, white-grey with a silvery shine, minutely uneven-verrucose; prothallus not observed. Thallus partially immersed in the periderm, in section 50–80 µm thick, ecorticate, dominated by the irregular photobiont layer, with large clusters of calcium oxalate crystals. Ascomata lirellate, more or less fissurine, unbranched to sparsely branched, immersederumpent, 0.5–1.5 mm long, 0.1–0.2 mm broad, 0.12–0.15 mm high; disc concealed; labia thin, entire, lighter than the thallus, along the slit with a thin, dark brown line, laterally covered by whitish thallus. Excipulum prosoplectenchymatous, 10–15 µm broad, yellowish, laterally covered by thallus including large clusters of crystals; periphysoids absent; hypothecium 10–15 µm high, light olive. Hymenium 90–100 µm high, clear; paraphyses unbranched, apically smooth. Asci 90–100 × 15–20 µm, oblong. Ascospores 8 per ascus, muriform, 15–20 × 9–12 µm, ellipsoid-oval, with thin septa and rectangular lumina, constricted at the median septum, hyaline, I–. Secondary chemistry: no sustances detected by TLC. Etymology. This new species is dedicated to Lino Olivares, one of the most knowledgeable and passionate empirical experts of the flora of the Colombian Caribbean dry forests and collaborator in the EcoSecos initiative and other conservation efforts in the region for 18 years (Castro-Vásquez et al. 2010, Rodríguez et al. 2012, Castellanos Castro and Newton 2015). ‐ Remarks. Currently, there are only two species known in Fissurina with a whitish, endoperidermal thallus, lacking substances, and muriform ascospores, namely F. alligatorensis Lendemer & R.C. Harris and F. ilicicola Lendemer & R.C. Harris (Lendemer and Harris 2014). The first agrees with the new species in thallus and ascoma morphology, as well as ascospore size, but has distoseptate, I+ strongly amyloid ascospores. The second one differs in the more distinct (hemithecioid) labia and the larger (24–32 × 12–14 µm), also distoseptate and I+ amyloid ascospores.

Graphis lurizana Lücking, Moncada & Celis sp. nov. Mycobank MB 828115 (Fig. 3b). Diagnosis. Differing from Graphis illinata in the apically exposed, black labia and the smaller ascospores. Type. COLOMBIA. Atlántico: Usiacurí, Distrito Regional de Manejo Integrado (DMI) Luriza; 10º44’22” N, 75º01’24” W, 100–200 m; 1 December 2015; R. Lücking, B. Moncada et al. 42922 (UNO, holotype; B, isotype). Description. Thallus corticolous, epiperidermal, up to 7 cm diam., continuous, white-grey, uneven, opaque; prothallus not observed. Thallus in section 100–150 µm thick, with thin, prosoplectenchymatous cortex and irregular photobiont layer, strongly encrusted with large clusters of calcium oxalate crystals. Ascomata lirellate, unbranched to sparsely branched, prominent, 1–3 mm long, 0.4–0.5 mm broad, 0.2–0.3 mm high; disc concealed; labia entire, apically exposed, black, with irregular, lateral thalline margin. Excipulum completely carbonized (thin at the base), 70–100 µm broad; hypothecium 20–30 µm high, hyaline. Hymenium 130–160 µm high, clear; paraphyses unbranched, apically smooth. Asci 120–130 × 25– 25 µm, oblong. Ascospores 1 per ascus, richly muriform, 80–110 × 25–30 µm, oblong, with slightly thickened septa and more or less rounded lumina, hyaline, I+ violet-blue. Secondary chemistry: no substances detected by TLC. Etymology. Graphis lurizana is dedicated to the Luriza community, who manages the Distrito Regional de Manejo Integrado (DMI) Luriza, the first protected area to be declared in the Atlántico Department (Molina-Acosta 2013). Remarks. Graphislurizana keysoutingroup 9 in the Graphis world key of Lücking et al. (2009a). There are four species in this group with large ascospores (> 80 µm), lacking secondary substances, and with elongate, prominent lirellae, namely G. acharii Fée, G. cleistomma Nyl., G. illinata Eschw., and G. subvernicosa Lücking. Graphis acharii and G. subvernicosa have (2–)4–6(–8) ascospores per ascus; in addition, in G. acharii the labia rather regularly become striate and the ascospores are longer (up to 170 µm), whereas in G. subvernicosa, the lirellae have only a basal thalline margin and the ascospores are narrower (15–20 µm broad). Graphis cleistomma and G. illinata agree with the new species in the singlespored asci; however, the first has shorter lirellae with an apically thin complete thalline margin, and larger ascospores (up to 180 × 40 µm), whereas the second has lirellae with an apically thick complete margin and slightly longer ascospores (up to 150 µm).

We present a first study of the diversity and community composition of lichens in seasonally dry tropical forest (DTF) remnants in the Atlántico department, Colombia. Lichens were sampled in two of the three protected areas of the department: Distrito de Manejo Integrado (DMI) Luriza and Reserva Forestal Protectora (RFP) El Palomar. The inventory revealed 61 species, including four new to science: Fissurina linoana Lücking, Moncada & G. Rodr. sp. nov., Graphis lurizana Lücking, Moncada & Celis sp. nov., G. mokanarum Lücking, Moncada & M.C. Martínez sp. nov., and Phaeographis galeanoae Lücking, Moncada & B. Salgado-N. sp. nov. Arthonia erupta and Coenogonium saepincola are new to South America, whereas thirteen species are recorded for Colombia for the first time. Further 37 species are new records for Atlántico, raising the total of species known from the department from 27 to 84. With 42 species at Luriza and 31 at El Palomar, species richness was comparable to that of other DTF sites in the Neotropics. Overlap in species composition between the two sites was remarkably low, with only twelve shared species (20 %), indicating a high level of heterogeneity. Biogeographical affinities lie with Central American DTF, which is in line with those of woody plants. These results underline the importance of the remaining fragments of DTF in Colombia in conserving partially unknown biodiversity and the necessity for their continuing conservation. RESUMEN Presentamos un primer estudio de la diversidad y composición de comunidades de líquenes en remanentes de bosque seco tropical (bs-T) en el departamento de Atlántico, Colombia. Se muestrearon líquenes en dos de las tres áreas protegidas del departamento: Distrito de Manejo Integrado (DMI) Luriza y Reserva Forestal Protectora (RFP) El Palomar. El inventario registró 61 especies, incluyendo cuatro especies nuevas para la ciencia: Fissurina linoana Lücking, Moncada & G. Rodr. sp. nov., Graphis lurizana Lücking, Moncada & Celis sp. nov., Graphis mokanarum Lücking, Moncada & M.C. Martínez sp. nov. y Phaeographis galeanoae Lücking, Moncada & B. Salgado-N. sp. nov. Arthonia erupta y Coenogonium saepincola son nuevos registros para América del Sur, mientras que trece especies son reportadas por primera vez para Colombia. Unas 37 especies adicionales son nuevos reportes para Atlántico, incrementando el total de especies conocidas para el departamento de 27 a 84. Con 42 especies en Luriza y 31 en El Palomar, la riqueza de especies fue comparable con la de otros sitios de bs-T en el Neotrópico. El traslape en la composición de especies entre los dos sitios es bajo, con solamente doce (20 %) compartidas, indicando un alto nivel de heterogeneidad. Las afinidades biogeográficas son mayores con los bs-T de Centroamérica, lo que coincide con las de plantas leñosas. Estos resultados enfatizan la importancia de los remanentes de bs-T en Colombia para la conservación de una biodiversidad poco conocida.

Sticta rhizinata Moncada & Lücking, spec. nov. (Fig. 7) MycoBank #801849 Differing from Sticta weigelii in the bryophilous growth habit and the long rhizines on the underside. Holotype: — COLOMBIA. Cundinamarca. Mun. La Calera, Parque Nacional Natural Chingaza, Monterrendondo; 4º 44' N, 73º 50' E; 3430 m; 13 Apr 2011, Moncada 4638 (UDBC; isotypes: COL, F). Primary photobiont cyanobacterial (Nostoc). Stipe absent. Thallus irregular to suborbicular, up to 15 cm diam., moderately branched, with 3–5 branches per 5 cm radius, branching anisotomous to pleurotomous; lobes laciniate, horizontal, adjacent to imbricate, involute, their apices rounded, involute, their margins crenate to sinuous, not thickened; lobe internodes (6–)8–13(–16) mm long, (6–)12–28(–32) mm broad; thallus coriaceous. Upper surface smooth to weakly foveolate or rugose–costillate, green-brown when fresh, redbrown and darkening towards the apices in the herbarium, shiny; surface glabrous, without papillae and pruina, but with irregular, yellowish maculae; marginal cilia absent. Apothecia not observed. Isidia abundant, predominantly marginal, aggregate, branched to coralloid, with rather long branches, vertically to obliquely oriented, up to 0.5 mm long and 0.5 mm broad, darker than the thallus, blackish brown to black, shiny, cylindrical to slightly flattened and becoming spathulate; basal stalk when present cylindrical, without cyphellae. Lower surface uneven, brown; primary tomentum dense up to the margin, thick but thinner towards the margin, spongy, rough, brown-black; secondary tomentum arachnoid, pale. Rhizines abundant, submarginal, aggregate, fibrillate and covered with secondary tomentum, brown-black, up to 10 mm long. Cyphellae 1–20 per cm 2 towards the thallus center and 61–100 per cm 2 towards the margin, scattered, round, urceolate with wide pore, erumpent to prominent, remaining below the level of the primary tomentum, with the margin elevated and involute, pale brown, without tomentum; pore (0.25–)0.4–0.8(–1.5) mm diam.; basal membrane pubescent, white, K + yellow, C –, KC–, P –. Medulla compact, cream-colored, K + yellow, C –, KC–, P –. No substances detected by TLC. Upper cortex paraplectenchymatous, 45–65 µm thick, formed by two different layers: upper layer consisting of 1–2 cell layers with cells 3–4 µm diam., their walls 1.2–2.5 µm thick and their lumina rounded to isodiametric, 1.2–2.5 µm diam.; lower layer consisting of 2–3 cell layers; cells 6–11 µm diam., their walls 1–1.2 µm thick and their lumina rounded to isodiametric, 5–10 µm diam. Photobiont layer 25–50 µm thick, its cells 10–20 µm diam. Medulla 55–175 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 25–40 µm thick, with 3–4 cell layers; cells 6–16 µm diam., their walls 1–1.5 µm thick. Hairs of lower primary tomentum 110–470 µm long, in fascicles of 12–20, branched hyphae, septate with their apices intertwined. Hairs of lower secondary tomentum 8–18 µm long, of single, branched, moniliform hyphae with free apices. Cyphellae cavity 30–150 µm deep; cells of basal membrane without papillae. Distribution and Ecology: —South American Andes (Colombia). Sticta rhizinata has been found between 2300 and 3720 m altitude in subandine forest and páramo regions in microhabitats with low to high light intensity. It occurs on the western slopes of the Cordilleras Central and Occidental. Typically it grows on the ground inbetween bryophytes of the genera Plagiochila, Frullania Raddi (1818), Metzgeria, Campylopus Brid. (Bridel 1819) and Dicranum Hedw. (Hedwig 1801), and lichens of the genera Hypotrachyna (Vain.) Hale (Hale 1974), Everniastrum Hale ex Sipman (Sipman 1986), and Peltigera Willd. (Willdenow 1787). Etymology: —The epithet refers to the very conspicuous rhizines of this species, quite different from other similar species in the Sticta weigelii complex. Remarks: —This species belongs in the Sticta weigelii complex, being characterized by a cyanobacterial photobiont and comparatively narrow, much branched lobes with marginal isidia (McDonald et al. 2003; Galloway 2006). Within that complex, S. rhizinata is well-characterized by its very narrow, laciniate lobes forming large and conspicuous rhizines on the underside, as well as its ecology, being typically found growing between bryophytes on the ground. Additional specimens examined (paratypes): — COLOMBIA. Boyacá: Mun. Aquitania, Páramo de los Pozos; 3115–3320 m; 26 Feb 2004, Medina & Castillo 10 (UPTC). Mun. Duitama, road from Duitama to Charalá; 3070 m; 23 Jul 1985, Escobar & Santa 213 (HUA). Mun. El Cocuy, Parque Nacional Natural El Cocuy, Alto de la Cueva; 3800 m; 21 Apr 2011, Fonseca & Martínez 224 (UDBC). Mun. Gachantivá, Vereda la Ciénaga, vía Cáscada La Honda; 2300 m; 18 May 2001, Moncada & Dávila 1846b (UDBC). Mun. Socha, Páramo de Pisba, road from Socha to La Punta, km 61.5, Alto de Granados; 3600 m; 10 Jun 1972, Cleef 4356 (B, COL, L). Cundinamarca. Alto de la Viga, Páramo de Cruz Verde, Camino Real towards Las Lagunas; 3450 m; 28 Nov 1974, Van den Hammen 3113 (B, COL). Parque Nacional Natural Sumapaz, Páramo de Chisacá, Santa Rosa, Bodegas de Sumapaz, Valle del Rio Colorados (Santa Rosita); 3470 m; 20 Aug 1976, Van den Hammen & Jaramillo-M. 4420 (B, COL). Parque Nacional Natural Sumapaz, Corregimiento Nazareth, summit to the east of Caseta Los Pinos; 3450 m; 11 May 2004, Moncada & Dávila 2178 (UDBC). Surroundings of Laguna de Chisacá; 4º 17' N, 74º 12 E; 3724 m; 15 Aug 2010, Moncada et al. 4063 (UDBC). Mun. Chipaque, Vereda Marilandia, vía Santuario; 2400 m; 8 Sep 2011, Moncada 4790, 4796, 4896 (UDBC). Mun. Choachí, El Verjón; PE Matarredonda; 04º 34' N, 74º 00' E; 2900–3220 m; 8 May 2010, Moncada 3163 (UDBC); ibid.; 3220 m; 23 Jan 2011, Moncada 4535 (UDBC). Mun. Guasca, Páramo de Guasca; 3350 m; 18 Aug 2011, Moncada 4766 (UDBC). Mun. La Calera, Parque Nacional Natural Chingaza, Monterrendondo; 4º 44' N, 73º 50' E; 3430 m; 13 Apr 2011, Moncada 4635b (UDBC). Mun. Supatá, Alto El Tablazo, Carretera Estación Radar to Supatá; 3400 m; 11–12 Sep 1984, Aguirre & Sipman 5262 (B, COL). Risaralda: Mun. Santa Rosa de Cabal, Vereda El Cedral, vía El Cedral to Laguna del Otún, 100 m of Finca Ceilán; 2300 m; 3 Sep 2003, Moncada & Dávila 2060 (UDBC)., Published as part of Moncada, Bibiana & Lücking, Robert, 2012, Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus, pp. 1-29 in Phytotaxa 74 (1) on pages 17-18, DOI: 10.11646/phytotaxa.74.1.1, http://zenodo.org/record/5066523, {"references":["Raddi, G. (1818) Jungermanniografia Etrusca. Memorie i Mathematica e di Fisica della Societa Italiana delle Scienze (Modena) 18: 14 - 56.","Bridel, S. (1819) Muscologia Recentiorum Supplementum 4. Gotha.","Hedwig, J. (1801) Species Muscorum Frondosorum. Paris & London.","Hale, M. E. Jr. (1974) Delimitation of the lichen genus Hypotrachyna (Vainio) Hale. Phytologia 28: 340 - 342.","Sipman, H. J. M. (1986) Notes on the lichen genus Everniastrum (Parmeliaceae). Mycotaxon 26: 235 - 251.","Willdenow, C. L. (1787) Florae Berolinensis Prodromus. Berlin.","McDonald, T., Miadlikowska, J. & Lutzoni, F. (2003) The lichen genus Sticta in the Great Smoky Mountains: a phylogenetic study of morphologial, chemical, and molecular data. The Bryologist 106: 61 - 79."]}

Sticta microcyphellata Moncada & Lücking, spec. nov. (Fig. 5) MycoBank #801847 Differing from Sticta ainoae in the thelotremoid and tiny cyphellae occurring in more than 120 per cm 2, the dark tomentum up to the margin; the tomentose-hirsute apothecial margin, and the yellow-cream, K+ wine red medulla. Holotype: — COLOMBIA. Norte de Santander: Parque Nacional Natural Tamá, trail from Quebrada La Pedrera to Paramito; 2640 m; 10 Sep 2000, Moncada & Dávila 1024 (UDBC; isotype: COL). Primary photobiont green (Dictyochloropsis). Stipe absent. Thallus irregular, up to 30 cm diam., moderately branched, with 3–5 branches per 5 cm radius, branching dichotomous to polytomous; lobes laciniate, horizontal to weakly subpendulous, interspaced to imbricate, plane to involute, their apices obtuse to truncate, plane to involute, their margins entire, weakly thickened; lobe internodes 12–30 mm long, 7–15 mm broad; thallus coriaceous. Upper surface smooth to weakly foveolate, bright green when fresh, pale yellowishbrown in the herbarium, matt; surface glabrous, with papillae towards the margin, without pruina, with irregular, white maculae; marginal cilia present, sparse, fasciculate, brown to black, up to 0.5 mm long. Vegetative propagules absent. Apothecia sparse, predominantly submarginal, dispersed, pedicellate, with slight invagination on the lower side, up to 4 mm diam.; disc red-brown, shiny; margin tomentose to weakly hirsute, pale brown. Lower surface uneven, pale to dark brown; primary tomentum dense up to the margin, thin, spongy to arachnoid, soft, dark brown; secondary tomentum absent. Rhizines scattered, submarginal, fibrillose, brown, up to 5 mm long. Cyphellae more than 100 per cm 2 towards the thallus center and 100–200 per cm 2 towards the margin, aggregate, rounded, thelotremoid, immersed to erumpent, remaining below the level of the primary tomentum, with the margin elevated and involute, cream-colored, without tomentum; pore 0.05–0.2 mm diam.; basal membrane pubescent, cream-colored to yellowish, K – to K + pale yellow, C –, KC– to KC+ pale yellow, P –. Medulla compact, cream-colored to yellowish, K + pink to wine-red, C –, KC–, P –. No substances detected by TLC. Upper cortex paraplectenchymatous, 35–45 µm thick, formed by two different layers: upper layer consisting of 1–2 cell layers with cells 2.5–4 µm diam., their walls 1.25 µm thick and their lumina rounded to isodiametric, 1.2–1.5 µm diam.; lower layer consisting of 3–4 cell layers; cells 6–11 µm diam., their walls 0.6–1.2 µm thick and their lumina rounded to isodiametric, 5–10 µm diam. Photobiont layer 20–30 µm thick, its cells 2.5–4 µm diam. Medulla 130–170 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 35–40 µm thick, with 3–4 cell layers; cells 6–14 µm diam., their walls 1.2–1.5 µm thick. Hairs of lower primary tomentum 70–200 µm long, in fascicles of 6–12, branched hyphae, septate with free apices. Cyphellae cavity 55–80 µm deep; cells of basal membrane without papillae. Apothecia biatorine, 500–700 µm high, with indistinct stipe; excipulum 100–120 µm broad, with tomentum. Hymenium 130–140 µm high; epihymenium 2.5–5 µm high, orange-brown, with gelatinous layer above. Ascospores 1–3-septate, fusiform, 30–40 × 5–7.5 µm, hyaline. Distribution and Ecology: —South American Andes (Colombia and Venezuela). The species is found between 1900 and 3700 m altitude in upper montane to subandine forests, in shaded to moderately exposed microsites on bark and sometimes on decaying organic matter. The liverwort genera Plagiochila and Radula Dumort. (Dumortier 1822) and the lichen Leptogium phyllocarpon (Pers.) Mont. (Montagne 1848) are commonly associated. Etymology: —Among tropical green-algal species with large thalli, this species is characterized by extremely small cyphellae. Remarks: —This new species is rather conspicuous due to its large, bright green thalli with rather broad, laciniate lobes, combined with very numerous, small, thelotremoid cyphellae. It is very similar to Sticta ainoae D. J. Galloway & J. Pickering (Galloway & Pickering 1990) from southern South America (type in BM revised), which has a marginally pale lower tomentum (often thin or absent towards the margin) and slightly larger, less numerous cyphellae, with the pore being 0.1–0.5 mm in diam. (not 0.05–0.3 as stated by Galloway & Pickering 1990). Other differences include the white, K– or K+ weakly ochraceous medulla, the glabrous apothecial margins, and the smaller ascospores in S. ainoae. Within Colombia, S. microcyphellata is the only green species with thelotremoid cyphellae. Additional specimens examined (paratypes): — COLOMBIA. Chocó: Mun. El Carmen de Atrato, Alto de Cortes; 2600 m; 28 Feb 1944, Core 297 (US). Huila: Mun. San Agustín, Páramo de las Papas, mountains and surroundings of Lagunas La Magdalena and Santiago; 3530–3630 m; 5–19 Sep 1985, Bischler 652 (COL). Norte de Santander: Mun. Toledo, Hoya de Samaria; 2100 m; 30 Oct 1941, Cuatrecasas et al. 12795 (COL). Parque Nacional Natural Tamá, Sector Orocué; 2800 m; 25 Apr 1981, Ruiz s.n. (COL). Santander: Mun. Charalá, Virolín, lower part of Cuchilla El Fara; 1820 m; 12 Nov 1984, Castro & Cruz 378 (COL). Valle del Cauca: Mun. El Silencio, Yanaconas; 1900–2200 m; 28 Feb 1944, Killip & García 33780 (COL). VENEZUELA. Táchira: Páramo de Tamá, base of Cobre Chiquito; 2400 m; 28 Mar 1979, López Figueiras & Hale 45614 (US). Villa Paez-Betania; 2550 m; 18 May 1967, Steyermark & Dunsterville 98372 (US); ibid., southeast of Santa Ana, Río Quinimarí; 2400 m; 10 Jan 1968, Steyermark & Dunsterville 100638 (US).

Sticta tunjensis Moncada & Lücking, spec. nov. (Fig. 10) MycoBank #801852 Differing from Sticta weigelii by the flabellate lobes with distinctly maculate surface, the presence of marginal cilia, and the papillate cells of the basal membrane of the cyphellae. Holotype: — COLOMBIA. Boyacá: Mun. Tunja, Vereda Barón Germania, El Malmo; 3065–3085 m; 17 Mar 2003, Alvaro et al. 474 (UPTC; isotype: UDBC). Primary photobiont cyanobacterial (Nostoc). Stipe absent. Thallus irregular, up to 5 cm diam., moderately branched, with 3–5 branches per 5 cm radius, branching pleurotomous; lobes flabellate, horizontal to subpendulous, imbricate, revolute to weakly canaliculate, their apices rounded to irregular in outline, involute, their margins crenate, not thickened; lobe internodes (3–)4–6(–8) mm long, (2–)3–8(–12) mm broad; thallus subcoriaceous, fragile. Upper surface smooth, dark green when fresh, pale ochraceous yellow to greenish grey in the herbarium, shiny; surface glabrous, without papillae and pruina, but with irregular, white to yellowish maculae; marginal cilia sparse or absent in oldest lobe portions, simple to fasciculate, white, up to 0.5 mm long. Apothecia not observed. Isidia abundant, predominantly marginal, aggregate, branched to coralloid, with long branches, horizontal to obliquely oriented, up to 0.5 mm long and 0.05 mm broad, blue-grey, matt, cylindrical; basal stalk when present cylindrical, without cyphellae. Lower surface smooth, cream-colored towards the apices and darker towards the center; primary tomentum irregular, absent towards the margin, thick, thinner towards the margin, fasciculate to spongy, soft, white to cream-colored; secondary tomentum pubescent, pale. Rhizines absent. Cyphellae 1–20 per cm 2 towards the thallus center and 41–60 per cm 2 towards the margin, scattered, rounded, urceolate with wide pore, erumpent to prominent, remaining below the level of the primary tomentum or emerging towards the margin, with the margin elevated and involute, white to cream-colored or dark brown towards the thallus center, without tomentum; pore (0.2–)0.3–0.5(–0.9) mm diam.; basal membrane pubescent, cream-colored, K + pale yellow, C –, KC–, P –. Medulla compact, white, K + pale yellow, C –, KC–, P –. No substances detected by TLC. Upper cortex paraplectenchymatous, 12–25 µm thick, uniform, consisting of 2–3 cell layers with cells 3–8 µm diam., their walls 0.6–2.5 µ m thick and their lumina rounded to isodiametric, 1.5–5 µm diam. Photobiont layer 20–35 µm thick, its cells 10–18 µm diam. Medulla 45–55 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 12–15 µm thick, with 1–2 cell layers; cells 3–7 µm diam., their walls 0.6–1 µm thick. Hairs of lower primary tomentum 470–560 µm long, in fascicles of more than 20, unbranched hyphae, septate with free apices. Hairs of lower secondary tomentum 5–12 µm long, of single, unbranched hyphae, septate with free apices. Cyphellae cavity 50–220 µm deep; cells of the basal membrane with one papilla. Distribution and Ecology: —South American Andes (Colombia); found between 1900 and 3080 m altitude, in (sub-)andine forest of the Cordillera Oriental, in microhabitats with medium light intensity; growing on bark and soil and usually associated with liverworts of the genus Plagiochila. Etymology: —Named after the type locality. Remarks: — Sticta tunjensis shares the general morphology with S. weigelii in having branched, cyanobacterial thalli with marginal isidia (McDonald et al. 2003; Galloway 2006). However, it can be distinguished by the green-grey rather than brown thallus color, the flabellate lobes, the distinctly maculate surface, the presence of marginal cilia, the paler lower tomentum, and the papillate cells of the basal membrane of the cyphellae. Phylogenetically, S. tunjensis is rather isolated and not closely related to any of the other species with S. weigelii morphology (Moncada et al. 2012a). Additional specimens examined (paratypes): — COLOMBIA. Boyacá: Mun. Villa de Leyva, SFF Iguaque; 2100–2900 m; 9 Jun 2001, Linares et al. 9699 (COL). Santander: Mun. Charalá, Virolín, creek trail from Finca La Araña to Cascada; 1950 m; 27 Apr 1984, Castro et al. 60 (COL)., Published as part of Moncada, Bibiana & Lücking, Robert, 2012, Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus, pp. 1-29 in Phytotaxa 74 (1) on pages 23-25, DOI: 10.11646/phytotaxa.74.1.1, http://zenodo.org/record/5066523, {"references":["McDonald, T., Miadlikowska, J. & Lutzoni, F. (2003) The lichen genus Sticta in the Great Smoky Mountains: a phylogenetic study of morphologial, chemical, and molecular data. The Bryologist 106: 61 - 79.","Moncada, B., Lucking, R., Parmnen, S. & Lumbsch, H. T. (2012 a) Sticta fuliginosa (Lobariaceae): species or morphotype? In: The 7 th IAL Symposium: Lichens: From Genome to Ecosystems in a Changing World. Abstracts: 159. Ramkhamhaeng University Press, Bangkok, Thailand."]}

Sticta arachnofuliginosa Moncada & Lücking, spec. nov. (Fig. 1) MycoBank #801843 Differing from Sticta fuliginosa in the larger lobes with faveolate, pubescent-arachnoid surface and the lack of papillae on the cells of the basal membrane of the cyphellae. Holotype: — COLOMBIA. Cundinamarca: Mun. Bogotá D. C., Surroundings of Laguna de Chisacá; 4º 17' N, 74º 12' W; 3734 m; Apr 2008, Moncada et al. 4007 (UDBC; isotypes: F, COL). Primary photobiont cyanobacterial (Nostoc). Stipe absent. Thallus orbicular to irregular in outline, up to 15 cm diam., sparsely branched, with 0–2 branches per 5 cm radius, branching polytomous; lobes suborbicular, horizontal, imbricate, plane to revolute, with rounded, revolute apices, margins entire to crenate, not thickened; lobe internodes 25–50 mm long, 25–60 mm broad; thallus coriaceous. Upper surface faveolate, grey to olive-brown when fresh, brownish grey in the herbarium, matt; surface pubescent, without papillae, pruina, and maculae; marginal cilia absent. Apothecia not observed. Isidia present, abundant, principally laminal, subaggregate, simple to branched or becoming coralloid, vertically oriented, up to 0.2 mm long and 0.05 mm broad, darker than the thallus, brown to blackish brown, shiny, granular to cylindrical; basal stalk when present cylindrical, without cyphellae. Lower surface uneven, white to cream-colored; primary tomentum dense up to the margin, thin to thick, hirsute to fasciculate, soft, white to cream-colored; secondary tomentum pubescent, white to cream-colored. Rhizines absent. Cyphellae scattered, 1–20 per cm 2 towards the thallus center and 21–40 per cm 2 towards the margin, rounded to irregular or becoming angular, plane, immersed, margin remaining below the level of the primary tomentum, white to cream-colored, with tomentum; pore 1–5 mm diam.; basal membrane pubescent, white, K – to K + pale yellow, C –, KC–, P –. Medulla lax, white, K – to K + pale yellow, C –, KC– to KC+ pale yellow, P –. No substances detected by TLC. Upper cortex paraplectenchymatous, 35–50 µm thick, uniform, consisting of 4–5 cell layers with cells 5–10 µ m diam., their walls 1.2–2.5 µm thick and their lumina rounded to isodiametric, 3–7 µm diam. Photobiont layer 35–60 µm thick, its cells 12–18 µm diam. Medulla 30–120 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 25–35 µm thick, with 2–3 cell layers; cells 4–10 µm diam., their walls 1.2–2.5 µm thick. Hairs of upper primary tomentum 130–160 µm long, in fascicles of up to six, branched hyphae with free, moniliform apices. Hairs of upper secondary tomentum 15–30 µm long, of single, branched, moniliform hyphae with free apices. Hairs of lower primary tomentum 350–480 µm long, in fascicles of 12–20, branched hyphae with free, moniliform apices. Hairs of lower secondary tomentum 20–50 µm long, of single, branched, moniliform hyphae with free apices. Cyphellae cavity 75–280 µm deep; cells of basal membrane without papillae. Distribution and Ecology: —Central American Cordilleras and South American Andes, as well as southern Chile (Guatemala to Chile). The species is found between 2300 and 4050 m altitude, in the páramo region and extending downwards into subandine and cool-temperate forests, usually in more or less exposed microsites. The Colombian records are from the Cordilleras Central and Occidental. The species grows mostly on bark but sometimes also on soil, usually associated with liverworts of the genera Metzgeria Raddi (1818), Lepicolea Dumort. (Dumortier 1835) and Plagiochila (Dumort.) Dumort. (Dumortier 1835), and the lichen Heterodermia circinalis (Zahlbr.) W. A. Weber (Weber 1981). Etymology: —The epithet refers to the superficial similarity with Sticta fuliginosa (Dicks.) Ach. (Acharius 2003) and the presence of a tomentum on the upper lobe surface. Remarks: —This is one of several species traditionally classified under the name Sticta fuliginosa (for an exact description of that species, see Suárez & Lücking 2012), sharing with the latter the broad, rounded, sparsely branched thallus lobes and laminal isidia, but differing in several morphological and anatomical details and being phylogenetically unrelated to S. fuliginosa s.str. (Moncada et al. 2012a). Thus, S. arachnofuliginosa is larger and more robust that S. fuliginosa and immediately set apart by the faveolate, tomentose upper lobe surface; in addition, the cyphellae are much larger and often becoming angular in outline and their basal membrane lacks papillae on the cells. Of all species with a S. fuliginosa gross morphology in Colombia, S. arachnofuliginosa appears to the most common and widespread (Moncada 2012), being in fact found throughout Central and South America. Additional specimens examined (paratypes): — GUATEMALA. Chimaltenango: Cerro de Tecpam; 4 Dec 1938, Standley 58725 (F, US). COLOMBIA. Boyacá: Mun. Arcabuco, 15 km from Arcabuco via Tunja; 2900 m; 11 Jul 1986, Sipman & Reyes 34392 (B, COL). Mun. El Cocuy, Parque Nacional Natural El Cocuy, Valle de las Lagunillas, Cabaña Sisuma; 6º 26' N, 72º 22' E; 3800 m; 19 Apr 2011, Fonseca & Martínez 27, 28, 62, 63 (UDBC). Parque Nacional Natural El Cocuy, Alto de la Cueva; 3800 m; 21 Apr 2011, Fonseca & Martínez 188a, 243 (UDBC). Mun. Villa de Leyva, SFF Iguaque, trail from ranger station vía Laguna de Iguaque; 2805 m; 24 May 2000, Moncada & Dávila 785 (UDBC). Caldas: Mun. Aranzazú, Vereda El Diamante, Finca El Porvenir 5º 15' N, 75º 26' E, 2600 m; 2010, Coca 570 (UDBC). Mun. Santa Rosa de Cabal, Nevado del Ruíz, northwest slope, near hotel of hot springs; 3500 m; 4 Feb 1979, Sipman & Valencia 10528 (COL, L). Cauca: Mun. Puracé, Volcán Puracé, near Pilimbalá; 3470 m; 5 Jan 1976, Cleef & Fernández-P. 493 (COL, L). Parque Nacional Natural Puracé, Pilimbalá, Laguna de San Rafael; 3300 m; 6 Oct 1984, Aguirre & Sipman 5979 (B, COL). Termales de San Juan; 3150 m; 14 Oct 1989, Cardona de Hollaender 992 (CUVC). Cundinamarca: Mun. Bogotá D. C., Parque Nacional Natural Sumapaz, Páramo de Chisacá, road from Usme to Nazareth, towards Laguna Grande.; 3700 m; 4 Sep 1984, Aguirre & Sipman 5004 (B, COL). Páramo de Chisacá, road towards Laguna Larga; 3500 m; 15 Jul 1986, Cardona de Hollaender et al. 40b (CUVC). Surroundings of Laguna de Chisacá; 4º 17' N, 74º 12' E, 3734 m; 8 Apr 2008, Beltrán et al. 51 (UDBC); ibid.; Ardila & Gómez 208 (UDBC), Moncada et al. 4027, 4034, 4049, 4129b, 4130, 4132, 4134b, 4139, 4149, 4227 (UDBC). Mun. Choachí, El Verjón. PE Matarredonda; 4º 33' N, 74º 00' E, 3220 m; 23 Jan 2011, Moncada 4607, 4618 (UDBC). Mun. Cogua, páramo between Cógua and San Cayetano, near Laguna Seca; 3685 m; 9 Nov 1972, Cleef 6139 (COL, L). Mun. Fómeque, Parque Nacional Natural Chingaza, road from Playa to Chuza; 3205 m; 1–4 Apr 1999, Dávila et al. 91 (UDBC). Parque Nacional Natural Chingaza, Río la Playa, Valle de los Frailejones; 3145 m; 1–4 May 1999, Moncada & Dávila 245b, 278, 302, 309, 325b (UDBC). Mun. Guasca, Páramo de Guasca; 3350 m; 18 Ago 2011, Moncada 4726 (UDBC); ibid.; 3200 m; 18 Ago 2011, Lücking 33342, 33346 (F, UDBC). Mun. La Calera, Páramo de Palacio; 3140 m; 22 Jul 1965, Merrill King et al. 482 (US). Zona de Amortiguación Parque Nacional Natural Chingaza, Vía del Ángulo towards Rincón del Oso; 3600 m; 7 Nov 2011, Moncada 4904, 4927 (UDBC). Mun. Villapinzón, Páramo de Villapinzón; 3300 m; 10 Sep 2002, Pinzón 1485 (COL). Huila: Mun. La Plata, Vereda La Candelaria, Rio La Candelaria waterfall; 2300 m; 16 Sep 1985, Aguirre & Sipman 6145 (B, COL). Norte de Santander: Mun. Toledo, Parque Nacional Natural Tamá, Sector El Paramito; 2800 m; 12 Sep 2000, Moncada & Dávila 1301, 1317 (UDBC). Risaralda: Mun. Pereira, Parque Nacional Natural Los Nevados, Laguna de Otún, northwest border; 3795 m; 16 Jan 1980, Boekhout 40 (COL). Mun. Santa Rosa de Cabal, Costado Occidental, Cordillera Central, 500 m south of Finca La Sierra; 3750 m; 1 Oct 1984, Aguirre & Sipman 5465 (B, COL). Tolima: Mun. Herveo, Parque Nacional Natural Los Nevados, Sector Brisas; 4050 m; 1–2 Jul 2002, Dávila 2643 (UDBC). VENEZUELA. Mérida: Sierra Nevada, Laguna Negra; 3500–3600 m; 5 Apr 1975, Hale & Figueiras 44556 (US). ECUADOR. Carchi: Valle de Maldonado, Carretera Tulcán-Maldonado; 3200 m; 18 May 1973, Nielsen et al. 5595c (US). CHILE. X. Región: Río Enco and roadside between Choshuenco and Enco; 39º 54' S, 72º 09' W, 400 m; Coppins et al. 8192 (BM). Parque Nacional Vicente Perez Rosales, Petrohue, Río Petrohue, near falls; 41º 08' S, 72º 27' W, 500 m; on rocks and in Nothofagus forest; 10 Dec 1986, Coppins et al. 8205 (BM). XI. Región: Laguna San Rafael; 46º 40' S, 74º 00' W; mature rainforest on hillside slopes above moraines of San Rafael Glacier; 28 Jan 1990, Galloway 9134 (BM). Laguna San Rafael National Park, vicinity of Puntilla del Cisne, around the hotel; 46º 38' S, 73º 51' W, 5–10 m; open scrubland with single Nothofagus and Embothrium, on dead scrub; 10 Nov 1997, Wedin 6052 (BM)., Published as part of Moncada, Bibiana & Lücking, Robert, 2012, Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus, pp. 1-29 in Phytotaxa 74 (1) on pages 4-6, DOI: 10.11646/phytotaxa.74.1.1, http://zenodo.org/record/5066523, {"references":["Raddi, G. (1818) Jungermanniografia Etrusca. Memorie i Mathematica e di Fisica della Societa Italiana delle Scienze (Modena) 18: 14 - 56.","Dumortier, B. C. (1835) Recueil d'Observations sur les Jungermanniacees: Revision des Genres. Blanquart, Tournay.","Weber, W. A. (1981) Lichenes Exsiccati distributed by the University of Colorado Museum, Boulder: Fascicles 1 - 15, nos. 1 - 600, 1961 - 1979. Mycotaxon 13: 85 - 104.","Suarez, A. & Lucking, R. (2012) Sticta viviana (lichenized Ascomycota: Peltigerales: Lobariaceae), a new species from Colombian paramos. Lichenologist (in press).","Moncada, B., Lucking, R., Parmnen, S. & Lumbsch, H. T. (2012 a) Sticta fuliginosa (Lobariaceae): species or morphotype? In: The 7 th IAL Symposium: Lichens: From Genome to Ecosystems in a Changing World. Abstracts: 159. Ramkhamhaeng University Press, Bangkok, Thailand."]}

Sticta isidiokunthii Moncada & Lücking, spec. nov. (Fig. 3) MycoBank #801845 Differing from Sticta kunthii in the presence of marginal isidia. Holotype: — COLOMBIA. Cundinamarca: Mun. La Calera, Parque Nacional Natural Chingaza, Monterrendondo; 4º 44' N, 73º 51' W; 3430 m; 13 Apr 2011, Moncada 4630 (UDBC; isotypes: COL, F). Primary photobiont cyanobacterial (Nostoc). Stipe absent. Thallus irregular to suborbicular, up to 15 cm diam., moderately branched, with 3–5 branches per 5 cm radius, branching polytomous; lobes laciniate to flabellate, horizontal to subpendulous, imbricate, involute to weakly canaliculate, with their apices irregular and involute and their margins crenate to lacerate, not thickened; lobe internodes (5–)8–25(–35) mm long, (5–)10–20(–30) mm broad; thallus coriaceous. Upper surface pitted to rugose-foveolate towards the center, dark green when fresh, greyish brown with darker apices in the herbarium, shiny; surface glabrous, without papillae and pruina but with irregular, scattered, white maculae; marginal cilia absent. Apothecia sparse, principally laminal, dispersed, subpedicellate, with pronounced invagination on lower side, up to 2.5 mm diam.; disc orange-brown to red-brown, shiny; margin hirsute, with white to brown hairs. Isidia abundant, predominantly marginal, aggregate, branched to coralloid, vertically to obliquely oriented, up to 0.5 mm long and 0.5 mm broad, darker than the thallus, greenish brown to dark brown, shiny, granular to cylindrical or very slightly flattened; basal stalk when present flattened, without cyphellae. Lower surface uneven, beige to dark brown towards the center; primary tomentum dense, thick but thinner towards the margin, spongy, soft, beige; secondary tomentum absent. Rhizines absent. Cyphellae 1–10 per cm 2 towards the thallus center and 21–40 per cm 2 towards the margin, scattered, irregular, urceolate with wide pore, erumpent to sessile, remaining below the level of the primary tomentum, with the margin elevated and involute, cream-colored, without tomentum; pore (0.25–)0.5–1(–2) mm diam.; basal membrane pubescent, white, K – to K + pale yellow, C –, KC–, P –. Medulla compact, white, K + ochraceous yellow, C –, KC–, P –. No substances detected by TLC. Upper cortex paraplectenchymatous, 25–50 µm thick, uniform, consisting of 2–3 cell layers with cells 8–19 µm diam., their walls 0.6–1.5 µm thick and their lumina rounded to isodiametric, 6–16 µm diam. Photobiont layer 25–50 µm thick, its cells 10–20 µm diam. Medulla 50–150 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 20–25 µm thick, with 2 cell layers; cells 8–14 µm diam., their walls 1.2–1.5 µm thick. Hairs of lower primary tomentum 125–450 µm long, in fascicles of 12–20, unbranched hyphae, septate with free apices. Cyphellae cavity 100–130 µm deep; cells of basal membrane without papillae. Apothecia biatorine, 400–500 µm high, with distinct stipe; excipulum 100–130 µm broad, laterally with projecting hairs. Hymenium 70–80 µm high; epihymenium 2.5–5 µm high, orangebrown, without gelatinous upper layer. Ascospores not observed. Distribution and Ecology: —South American Andes (Colombia and Bolivia), found between 2290 and 3600 m altitude, in subandine forest and páramo regions, on bark of shrubs and treelets in more or less exposed microhabitats; usually associated with liverworts of the genera Metzgeria and Microlejeunea Stephani (1888) and lichens of the genera Leptogium (Ach.) Gray (Gray 1821), Erioderma Fée (Fée 1825), and Sticta. Etymology: —This species resembles Sticta kunthii Hook f. (Kunth 1822) in thallus morphology but produces isidia. Remarks: —This new species has pitted thallus lobes and a general thallus morphology as in Sticta kunthii and its relatives (Galloway 1995), but differs from those species in the marginal isidia, whereas apothecia are very rare. Sticta isidiokunthii is not the isidiate counterpart of S. kunthii s.str., but instead forms its own clade (Moncada et al. 2012a). Because of the marginal isidia, S. isidiokunthii could be confused with S. weigelii and similar species (McDonald et al. 2003; Galloway 2006), but differs from those in the pitted lobe surface and the pale underside. Additional specimens examined (paratypes): — COLOMBIA. Boyacá: Mun. El Cocuy, Parque Nacional Natural El Cocuy, Alto de la Cueva; 3800 m; 21 Apr 2011, Fonseca & Martínez 225b, 247 (UDBC). Mun. Villa de Leyva, SFF Iguaque, surroundings of the administrative building; 2805 m; 24 May 2000, Moncada & Dávila 844 (UDBC). Cundinamarca: Mun. Bogotá D. C., Parque Nacional Natural Sumapaz, Páramo de Chisacá, Santa Rosa, Bodegas de Sumapaz, Valle del Rio Colorados (Santa Rosita); 3460 m; 20 Aug 1976, Van der Hammen & Jaramillo-M. 4354 (B, COL). Mun. Choachí, El Verjón, PE Matarredonda; 4º 33' N, 74º 00' E; 3220 m; 23 Jan 2011, Moncada 4587 (UDBC). Mun. Guasca, Páramo de Guasca; 3350 m; 18 Aug 2011, Moncada 4760, 4761, 4764 (UDBC); ibid.; Lücking 33302, 33320, 33330 (F, UDBC). Mun. La Calera, Parque Nacional Natural Chingaza, Monterredondo; 4º 44' N, 73º 50' E; 3430 m; 13 Apr 2011, Moncada 4624, 4629, 4631, 4632, 4633, 4634, 4643 (UDBC). Zona de Amortiguación Parque Nacional Natural Chingaza, Vía del Ángulo towards Rincón del Oso; 3600 m; 7 Nov 2011, Moncada 4959, 4969 (UDBC). Mun. San Antonio del Tequendama, PN Chicaque; 2290 m; 23 May 1996, Moncada & Dávila 19b (UDBC). Norte de Santander: Mun. Toledo, Parque Nacional Natural Tamá, Sector Orocué, surroundings of administrative building; 2490 m; 11 Sep 2000, Moncada & Dávila 1047 (UDBC); ibid.; 2650 m; 12 Sep 2000, Moncada & Dávila 1203 (UDBC). Tolima: Unknown locality; 1844, Goudot s.n. (PC). Mun. Santa Isabel, Camino Real between La Bodega and El Ochoral; 2940 m; 16 Feb 1980, Valencia & Boekhout 191to (COL). BOLIVIA. La Paz: Mapiri; Jan 1893, Bang 1754 (US; as Sticta querzicans var. peruviana)., Published as part of Moncada, Bibiana & Lücking, Robert, 2012, Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus, pp. 1-29 in Phytotaxa 74 (1) on pages 9-11, DOI: 10.11646/phytotaxa.74.1.1, http://zenodo.org/record/5066523, {"references":["Stephani, F. (1888) Westindische Hepaticae. Hedwigia 27: 276 - 302.","Gray, S. F. (1821) A Natural Arrangement of British Plants according to each other, as pointed out by Jussieu, De Candolle, Brown etc. with an Introduction to Botany. London.","Fee, A. L. A. (1825) Essai sur les Cryptogames des Ecorces Exotiques Officinales. Paris.","Kunth, C. S. (1822) Synopsis plantarum, quas in itinere circa plagas Orbis Novi colleg. Humboldt et Bonpland. Paris.","Galloway, D. J., Stenroos, S. & Ferraro, L. I. (1995) Flora Criptogamica de Tierra del Fuego. Lichenes Peltigerales: Lobariaceae y Stictaceae, Vol. 6, Fasc. 6. Consejo Nacional de Investigaciones Cientificas y Tecnicas de la Republica Argentina, Buenos Aires.","Moncada, B., Lucking, R., Parmnen, S. & Lumbsch, H. T. (2012 a) Sticta fuliginosa (Lobariaceae): species or morphotype? In: The 7 th IAL Symposium: Lichens: From Genome to Ecosystems in a Changing World. Abstracts: 159. Ramkhamhaeng University Press, Bangkok, Thailand.","McDonald, T., Miadlikowska, J. & Lutzoni, F. (2003) The lichen genus Sticta in the Great Smoky Mountains: a phylogenetic study of morphologial, chemical, and molecular data. The Bryologist 106: 61 - 79."]}

Sticta papillata Moncada & Lücking, spec. nov. (Fig. 6) MycoBank #801848 Differing from Sticta dilatata in the shorter stipe and entire apothecial margin. Holotype: — COLOMBIA Cundinamarca: Mun. Villapinzón, Páramo de Villapinzón; 3200 m; 1 Oct 2010, Alfonso 3 (UDBC). Primary photobiont cyanobacterial (Nostoc). Stipe present but indistinct, up to 5 mm long. Thallus palmate to irregular in outline, up to 10 cm diam., moderately branched, with 3–5 branches per 5 cm radius, branching polytomous to anisotomous; lobes laciniate to flabellate, ascending to almost vertical, adjacent to imbricate, involute to weakly canaliculate, their apices rounded, involute, their margins entire to crenate, not thickened; lobe internodes (5–)6–10(–12) mm long, (5–)8–12(–15) mm broad; thallus coriaceous. Upper surface smooth, dark green when fresh, greenish brown in the herbarium, shiny; surface glabrous, without papillae and pruina, but with abundant, irregular, cream-colored maculae; marginal cilia abundant, penicillate to agglutinate, dark brown, up to 1.5 mm long. Vegetative propagules absent. Apothecia principally laminal, dispersed, subpedicellate, with pronounced invagination on lower side, up to 1 mm diam.; disc red-brown, matt; margin entire, cream-colored. Lower surface smooth to uneven, cream-colored to pale brown towards the center; primary tomentum dense but sparse towards the margin, thick, becoming thin towards the margin, spongy, soft, dark brown; secondary tomentum present but sometimes indistinct, arachnoid, pale. Rhizines absent. Cyphellae 1–10 per cm 2 towards the thallus center and 41–60 per cm 2 towards the margin, scattered, rounded, plane to cupuliform, erumpent to sessile, remaining below the level of the primary tomentum, with the margin elevated to weakly involute, white, without tomentum; pore (0.8–)1–1.25(–1.8) mm diam.; basal membrane pubescent, white, K+ pale yellow, C–, KC–, P–. Medulla compact, white, K+ yellow, C–, KC–, P–. No substances detected by TLC. Upper cortex paraplectenchymatous, 12–35 µm thick, uniform, consisting of 3–5 cell layers with cells 3–10 µm diam., their walls 1.2–2.5 µm thick and their lumina rounded to isodiametric, 1.5–7 µm diam. Photobiont layer 20–70 µm thick, its cells 10–18 µm diam. Medulla 20–70 µm thick, its hyphae 2.5 µm broad, without crystals. Lower cortex paraplectenchymatous, 8–18 µm thick, with 2–3 cell layers; cells 2.5–6.5 µm diam., their walls 0.6–1.2 µm thick. Hairs of lower primary tomentum 120–720 µm long, in fascicles of more than 20, unbranched hyphae, septate with intertwined apices. Hairs of lower secondary tomentum 5–10 µm long, of single, unbranched hyphae, septate with free apices. Cyphellae cavity 80–170 µm deep; cells of the basal membrane with 4–6 papillae. Distribution and Ecology: —South American Andes (Colombia); found within a comparatively narrow altitudinal range of 2720–3200 m in (sub-)andine forests of the Cordilleras Occidental and Oriental, in moderately exposed microhabitats; growing on bark of trees and shrubs, associated with liverworts of the genera Radula and Metzgeria and lichens of the genus Leptogium, among others. Etymology: —Named after the distinct papillae on the cells of the basal membrane of the cyphellae; this was the first species in which we rediscovered this character after Vainio (1890) had initially described it for Sticta laevis (Nyl.) Vain. (Vainio 1890). Remarks: — Sticta papillata belongs in a group of various species with pedunculate, much branched thallus having rather narrow, distinctly ciliate lobes that produce very large and conspicuous cyphellae with more than one papillae per cell on their basal membrane. It is most similar to S. dilatata (Nyl.) Vain. (Vainio 1913), but has narrower lobes, entire instead of crenulate apothecial margins, and a less distinct stipe. It should be mentioned here that the name S. dilatata is misapplied in the literature. Zahlbruckner (1925) assumed that Vainio (1913) had based his combination on S. laciniata var. dilatata Nyl., a taxon described from Colombia and later renamed S. granatensis by Nylander (1874). However, Vainio (1913) based his combination on a very different lichen, S. tomentosa var. dilatata Nyl., also described from Colombia (Nylander 1860). Additional specimens examined (paratypes): — COLOMBIA. Cundinamarca: Mun. Guasca, Vereda La Concepción, basin of the Tunjo; 3000 m; 20 Jun 1989, Sánchez & Linares 1159 (COL). Mun. Guasca, Páramo de Guasca; 3200 m; 18 Ago 2011, Lücking 33370 (F, UDBC). Nariño: Mun. Piedrancha, La Planada, San Isidro, south of Ricaurte (road from Pasto to Tumaco); 1750 m; 2 Jun 1993, Sipman et al. 32871 (B, COL). Norte de Santander: Mun. Toledo, Parque Nacional Natural Tamá, trail from Quebrada la Pedrera to Paramito; 2680 m; 10 Sep 2000, Moncada & Dávila 988 (UDBC); ibid., Sector El Paramito; 2720 m; 12 Sep 2000, Moncada & Dávila 1274 (UDBC)., Published as part of Moncada, Bibiana & Lücking, Robert, 2012, Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus, pp. 1-29 in Phytotaxa 74 (1) on pages 15-17, DOI: 10.11646/phytotaxa.74.1.1, http://zenodo.org/record/5066523, {"references":["Vainio, E. A. (1890) Etude sur la classification et la morphologie des lichens du Bresil, I. Acta Societas pro Fauna et Flora Fennica 7: V - XXIX, 1 - 247.","Vainio, E. A. (1913) Lichenes Insularum Philippinarum II. Philippine Journal of Science C, Botany, 8: 99 - 137.","Zahlbruckner, A. (1925) Catalogus Lichenum Universalis 3. Borntraeger, Leipzig.","Nylander, W. (1874) Animadversiones circa Spruce Lichenes Amazonicos et Andinos. Flora 57: 70 - 73.","Nylander, W. (1860) Synopsis Methodica Lichenum Omnium Hucusque Cognitorum Praemissa Introductione Lingua Gallica Tractata. Martinet, Paris."]}

Moncada, Bibiana, Lücking, Robert (2012): Ten new species of Sticta and counting: Colombia as a hot spot for unrecognized diversification in a conspicuous macrolichen genus. Phytotaxa 74 (1): 1-29, DOI: 10.11646/phytotaxa.74.1.1, URL: http://biotaxa.org/Phytotaxa/article/view/phytotaxa.74.1.1