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This is the printable version of an interactive identification tool for 100 species of lichens commonly occurring in Thailand, prepared on the occasion of a workshop held in Bangkok in June 2017. Of course, this guide cannot be used to identify all lichens found in the Country, but it may be useful as a tutorial in workshops and courses for students and beginners, who will be asked to identify only the species which are included in the key. In this way, they will learn the basic terminology, and will make the first steps in lichen identification. The dichotomous key is also available in several stand-alone versions: printable, CD-Rom, and for mobile devices via the free app KeyToNature (Android and iOS). The key has been created using program FRIDA at the Department of Life Sciences of the University of Trieste (Italy), and will be further tested and implemented at the Department of Biology of the Ramkhamhaeng University, Bangkok, to encompass a broader set of species.

4. Pyxine coccifera (Fée) Nyl. , Mém. Soc. Impér. Sci. Nat. Cherbourg 5: 108 (1857). Parmelia coccifera Fée, Essai cryptog. écorces officin.: 126 (1825). Holotype:—[without locality] ad corticem Alcornocae (G!). For further synonyms see Kalb (1987). Thallus corticolous, very rarely saxicolous, 2–7 cm wide, adnate to loosely adnate. Lobes radiating, discrete to ± weakly contiguous, plane to convex, but often slightly concave towards the tips, 0.4–1.2 mm wide, subrotund at the apices. Upper surface grey to yellowish grey or dull yellow, pruina punctiform, sparse at the lobe tips or totally absent; polysidiangia and isidia absent. Pseudocyphellae scarlet, marginal and laminal, subreticulate, often developing into soralia. Soralia marginal and laminal, orbicular to linear, ± becoming excavate, producing bright red and grey granular soredia. Medulla pale yellow to creamy in upper part; lower medulla white; scarlet below the pseudocyphellae. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia very rare, obscurascens - type, 0.5–1.5 mm wide; disc epruinose or weakly pale grey-pruinose. Internal stipe distinct, pale yellow in the upper part, white below, K-, P-, C-, KC-. Ascospores 14–18 × 6–8 μm. Pycnidia not seen. Chemistry: Cortex K+ yellow, UV-; medulla K-, C-, KC-, P-; red-pigmented medulla K+ purple, C+ purple-brown, KC+ violet, P-; containing atranorin, chiodectonic acid, methyl pyxinate, methyl 3- Omethylpyxinate, 25-acetoxy-20,24-epoxydammerane-3-one, 25-acetoxy-20,24-epoxydammerane-3β- ol and norstictic acid in the hypothecium only (Elix 2009). Distribution and habitat:—Very common on bark and rock in coniferous, dry dipterocarp, mixed deciduous and shrub forests. Also in Australia, Central and South America and Africa. Notes:— Pyxine coccifera is well characterized by the vivid red-pigmented soralia and pseudocyphellae. Selected material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on the bark of Dipterocarpus obtusifolius in dry dipterocarp forest, 800 m, 27 June 1994, K . Boonpragob et al. 1860 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Khon Substation, on rocks in dry evergreen forest, 700 m, 17°06'43'' N, 101°37'51'' E, 21 May 2009, S . Meesim & K. Buaruang MSPL 148 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on rocks in dry dipterocarp forest, 397 m, 17°20'57'' N, 101°33'32'' E, 28 July 2009, S . Meesim & K. Buaruang MSPL 320 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Khon Substation, on the bark of Shorea obtusa in dry dipterocarp forest, 700 m, 17°14'32'' N, 101°18'28'' E, 10 November 2009, K . Buaruang & S. Meesim MSPL 506, 731 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, trail to Pha Kob, Lon Tae Forest Ranger Station, on the bark of an unidentified tree in coniferous forest, 1245 m, 17°12'00'' N, 101°33'48'' E, 12 November 2009, S . Meesim & K. Buaruang MSPL 604, 607 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, in front of Pra Tum Nak, on rocks in shrub forest, 850 m, 14°26' N, 101°42' E, 18 June 1999, P . Mongkolsuk 15097 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, on the bark of Dipterocarpus obtusifolius in dry dipterocarp forest, 700 m, 14°18' N, 101°37' E, 20 February 2000, P . Mongkolsuk 17356 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on pages 40-41, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}

Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun, Kalb, Klaus (2012): The lichen family Physciaceae in Thailand-I. The genus Pyxine. Phytotaxa 59 (1): 32-54, DOI: 10.11646/phytotaxa.59.1.2, URL: http://biotaxa.org/Phytotaxa/article/view/phytotaxa.59.1.2

6. Pyxine cognata Stirt. , Proc. Royal Phil. Soc. Glasgow 11: 311 (1879). Holotype:— INDIA. Nilgherries [Nilgiri]; A. Watt s. n. (BM!).— Pyxine berteriana var. himalaica D. D. Awasthi, Phytomorphology 30: 366 (1982). Holotype:— INDIA. Almora district; Uttar Pradesh, on way to Kasardevi, ca. 1930 m, on bark, 12 June 1956, D. D. Awasthi 3476 (LWG!). Thallus corticolous, 2–6 cm wide, adnate to tightly adnate. Lobes radiating, discrete to ± contiguous, plane to slightly concave, in older thalli becoming slightly convex in the centre, 0.3–1.0 mm wide, subrotund to truncate at the apices. Upper surface white to whitish grey or grey-brown, pruinose in extended patches near the apices, the pruina ± glistening; polysidiangia, isidia and soralia absent. Pseudocyphellae not prominent, usually retricted to the margins, rarely laminal or becoming subreticulate, often obscured by spreading pruina. Medulla orange in upper part; lower medulla white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia common, obscurascens - type, 0.3–1.0 mm wide; disc epruinose. Internal stipe distinct, upper part orange-red, K + pale red, P-, C-, KC-, lower part much paler or white, K-, P-, C-, KC-. Ascospores (13–) 16–20 × 6–7 μm. Pycnidia not seen. Chemistry: Cortex K-, UV + yellow; medulla K-, C-, KC-, P-; containing lichexanthone (major), terpenes (major or minor), unknown pigment (minor or trace). Distribution and habitat:—Uncommon on bark in dry dipterocarp, dry evergreen and mixed deciduous forests. Also in Australia, China, India, Java and South America; not yet known from Africa. Notes:— Pyxine cognata is characterized by the cortical lichexanthone, the absence of vegetative propagules, the partly orange medulla and apothecia of the obscuracens - type with an orange-red, K+ pale red internal stipe. Pyxine berteriana is similar, but it has broader, sparsely pruinose lobes and a medulla that is partly yellow. The two species contain very different arrays of terpenes (Kalb, 1987, pl. 11). Material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on Melia azedarach, hill evergreen forest, 1170 m, 30 June 1994, K . Boonpragob 2278 (RAMK).— Chiang Mai; Queen Sirikit Botanic Garden, on Stereospermum neuranthum, dry dipterocarp forest, 730 m, 18°53'54'' N, 98°51'38'' E, 30 June 1994, K . Boonpragob 3582 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Pak Dang Forest Ranger Station, on the bark of an unidentified tree in dry evergreen forest, 750 m, 17°10'42'' N, 101°21'48'' E, 22 May 2009, S . Meesim 259 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Ho Forest Ranger Station, on the bark of Tamarindus indica in mixed deciduous forest, 569 m, 17°07'13'' N, 101°34'30'' E, 29 July 2009, S . Meesim 338 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on the bark of an unidentified tree in mixed deciduous forest, 397 m, 17°20'49'' N, 101°33'29'' E, 28 July 2009, S . Meesim 344 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on the bark of Saraca indica in mixed deciduous forest, 397 m, 17°20'49'' N, 101°33'29'' E, 28 July 2009, S . Meesim 347, 348 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, on the bark of Roystonea regia in secondary forest, 385 m, 14°33' N, 101°40' E, 15 January 2000, N . Homchantara 16269 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 42, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}

8. Pyxine cylindrica Kashiw ., Bull. Natl Sci. Mus. , Tokyo, B, 3 (2): 66 (1977). Holotype:— PAPUA NEW GUINEA. Morobe District; about 3 km W of Lae, coconut tree plantation, H. Kashiwadani 12382 (TNS!). Thallus 2–5 cm wide, adnate, subdichotomously lobate. Lobes radiating, discrete to ± imbricate, plane to slightly convex, 0.3–0.6 mm wide, subrotund to truncate at the apices. Upper surface white to yellowish white or yellowish grey, sparsely pruinose at the lobe tips or epruinose; polysidiangia and soralia absent. Pseudocyphellae distinct, marginal and laminal, irregular or very rarely becoming reticulate. Isidia often along the margins, but also laminal, cylindrical, simple or sparingly branched, 0.3–0.8 mm tall, ca. 0.1 mm wide. Medulla uniformly white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia and pycnidia not seen. Chemistry: Cortex K + yellow, C-, KC-, P + yellow, UV –; medulla K + yellow → red, C-, P + orange; containing atranorin (minor), chloroatranorin (minor), norstictic acid (major), testacein (minor), unknown terpenes (minor). Distribution and habitat:—Rare on bark in dry dipterocarp forest. Also in Australia, South and East Asia and Papua New Guinea. Notes:—This lichen is characterized by the cylindrical isidia, the white medulla and the presence of atranorin, norstictic acid and testacein. Pyxine keralensis Awasthi is morphologically similar, but it has a yellow-orange medulla and it lacks norstictic acid and testacein. Material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on Syzygium claviflora in an open, dry dipterocarp forest, 870 m, 18°54'33'' N, 98°51'17'' E, 18 March 2008, K . Kalb (hb. Kalb 36807, 36815, 36836)— Chiang Mai; Queen Sirikit Botanic Garden, on Holgarna kurzii in an open, dry dipterocarp forest, 730 m, 18°53'54'' N, 98°51'38'' E, 17 January 1995, K . Boonpragob 3934 (RAMK 17630)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 44, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495

16. Pyxine retirugella Nyl. , Ann. Sci. Nat. ; Bot., 4, 11: 240 (1859). Holotype:—NOUKAHIVA. [Marquesas Islands, Nuku Hiva] ad cortices, D. E. S. A. Jardin s. n.; (H-NYL 31789!)— Pyxine consocians Vain., The Philipp. Journ. Sci., C, 8: 109 (1913). Holotype:— PHILIPPINES, Comiran Island, Sulu Sea, ad corticem arboris frondosae, September 1910, E. D. Merrill 7167 p.p. (TUR-VAIN. 8706!). Thallus 2–5 cm wide, adnate, subdichotomously lobate. Lobes irregular, discrete to contiguous or imbricate, plane to weakly convex but often slightly concave towards the tips, 0.3–1.0 mm wide, subrotund at the apices. Upper surface whitish to pale greenish grey or yellow-grey, pruinose towards the lobe tips or epruinose; isidia absent. Pseudocyphellae marginal and laminal, irregular, linear or becoming reticulate. Polysidiangia laminal and marginal, scattered or clustered, nodular to subcylindrical, short, ± branched, pustulate, bursting apically to produce granular soredia. Medulla uniformly white or cream. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia rare, obscurascens - type, 0.5–2.5 mm wide; disc grey-pruinose when immature. Internal stipe distinct, white to pale brown, K-, P-. Ascospores 17–22 × 6–9 µm. Pycnidia not seen. Chemistry: Cortex K + yellow, C-, KC-, P + yellow, UV-; medulla K + yellow → red, C-, P + orange; containing atranorin (minor), chloroatranorin (minor), norstictic acid (major), testacein (minor), unknown terpenes (minor). Distribution and habitat:—Occurs on bark, rarely on rocks, in hill evergreen, mangrove and mixed deciduous forests. Also in Africa, Australia, Asia, Christmas Island (Indian Ocean) and several Pacific islands. Notes:—This lichen is characterized by the adnate thalli, polysidiangia, white or cream medulla, cortical atranorin and obscurascens - type apothecia with a white to pale brown internal stipe. Pyxine physciaeformis is morphologically similar, but it contains cortical lichexanthone and has physciaeformis - type apothecia. P. boonpragobiana is morphologically also very similar, but differs by a different suite of terpenes. Material from Thailand examined:— Chiang Mai; Sunku, on twig of an unidentified tree in hill evergreen forest, 1400 m, 18°48' N, 98°58' E, 19 January 1999, P. Mongkolsuk 014376 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Ho Forest Ranger Station, on the bark of an unidentified tree in mixed deciduous forest, 569 m, 17°06'41'' N, 101°38'36'' E, 29 July 2009, S. Meesim & K. Buaruang MSPL 403 (RAMK).— Trat; Muang district, near Ban Nam Chieo, on Rhizophora apiculata in a ± disturbed mangrove forest, 3 m, 12°10'25'' N, 102°28'37'' E, 25 February 2011, K. Kalb (hb. Kalb 38858)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on pages 49-50, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495

14. Pyxine profallax Kalb , Bibl. Lichenol. 99: 243 (2009). Holotype: PAPUA NEW GUINEA. Central District; around Woitape, ca. 1600 m, 19 October 1975, Kurokawa 9309 (TNS!). Thallus corticolous, 3–5 cm wide, adnate. Lobes radiating, discrete to ± imbricate, plane to slightly concave, 0.8–1.0 mm wide, subrotund to truncate at the apices. Upper surface whitish to grey or brownish grey, sparsely pruinose at the lobe tips or epruinose; polysidiangia, isidia and soralia absent. Pseudocyphellae distinct, marginal and laminal, irregular, becoming reticulate on the upper surface. Medulla uniformly white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia common, obscurascens - type, 0.5–1.0 mm wide; disc black, not pruinose. Internal stipe distinct, white, K-, P-. Ascospores 16–22 × 6–8 μm. Pycnidia laminal, immersed to slightly sessile, black; conidia bacilliform, 3–4 × 1 µm. Chemistry: Cortex K+ yellow, C-, KC-, P+ yellow, UV-; medulla K+ yellow → red, C-, P+ orange; containing atranorin (minor), chloroatranorin (minor), norstictic acid (major), testacein (submajor), unknown terpenes showing the same array as P. fallax (minor). Distribution and habitat:—A very rare species, corticolous in a lower montane scrub. Also in Papua New Guinea (holotype). Notes:— Pyxine profallax is characterized by the adnate thallus, the absence of soredia, isidia and polysidiangia, the uniformly white medulla with norstictic acid, cortical atranorin, and obscurascens - type apothecia. P. philippina and P. schmidtii are very similar as both lack vegetative propagules and contain additional testacein, but the latter lacks norstictic acid and the former contains a different array of terpenes. Material from Thailand examined:— Loei; Phu Luang Wildlife Sanctuary, Lan Suriyan, on bark in a lower montane scrub, 1473 m, 17°16'44'' N, 31°07'02'' E, 25 June 2008, S. Meesim 29 (RAMK 17620)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on pages 48-49, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495

18. Pyxine sorediata (Ach.) Mont. , in Ramón de la Sagra, Hist. phys., polit. nat. Cuba, Bot. Plant. Cellul. 2: 188 (1842). Type:— Lecidea sorediata Ach., Syn. meth. lich.: 54 (1814). Lectotype (selected by Kalb 1987: 74 [as holotype]):—in America septentrionale, Müh/eⁿberg; (H-ACH 378!, isolectotypes: S!, UPS!).— Physcia glaucovirescens Nyl., Syn. meth. lich. 1(2): 419 (1860) = Pyxine glaucovirescens (Nyl.) Aptroot, Crypt., Bryol. Lichenol. 9: 146 (1988). Lectotype (selected by Aptroot 1988b: 146):— AUSTRALIA. Queensland; Morton [Moreton] Bay, Verreaux 1846; (PC n.v., fide A. Aptroot, loc. cit.; isolectotype:—(H). For further synonymy see Kalb (1987). Thallus corticolous as well as saxicolous, 3–8 (–20) cm wide, adnate to loosely adnate. Lobes radiating, contiguous to imbricate, plane to slightly concave, 0.6–1.3 mm wide, subrotund at the apices. Upper surface yellowish grey to brownish grey, lead-grey or blue-grey, distinctly pruinose; pruina punctiform towards lobe apices; polysidiangia and isidia absent. Pseudocyphellae very distinct at the margins, often grey-pruinose and becoming reticulate. Soralia initially marginal, developing from fissures, then laminal and orbicular, occasionally becoming corticate and developing into pseudoisidia; soredia granular, dirty white or grey. Medulla lemon-yellow above; lower part yellow-brown or yellow-orange. Lower surface black in the centre, paler towards the margin; rhizines ± dense, black to blue-black, furcate. Apothecia rare, obscurascens - type, 0.5–1.4 mm wide; disc epruinose. Internal stipe distinct; upper part dark orange, K + red, P-, C-, KC-; lower part whitish, K-, P-, C-, KC-. Ascospores 14–19 × 6–8 µm. Pycnidia rare, laminal immersed in small warts in thallus; conidiospores bacilliform, 3–4 × 1 µm. Chemistry: Cortex K + yellow or K-, C-, KC-, P + pale yellow or P-, UV –; medulla K-, C-, P-; containing atranorin (minor, or usually trace), chloroatranorin (trace), unknown terpenes (major), unknown pigment (minor). Distribution and habitat:—A rare species on rocks and bark in coniferous and dry dipterocarp forests. Also in Europe, North, Central and South America, Africa, Australia, Asia and the Pacific islands. Notes:—This species is characterized by the coarse, dirty white to grey, granular soredia, the absence of isidia and polysidiangia, the yellow to yellow-orange medulla, the prominent marginal pseudocyphellae and the presence of cortical atranorin. Pyxine endochrysina Nyl. is superficially similar, but it is esorediate and has nodular to squamulose or lobulate, marginal isidia. Material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on the trunk of Syzygium claviflora in dry dipterocarp forest, 750 m, 18°53'54'' N, 98°51'38'' E, 1 July 1994, K . Boonpragob et al. 2667 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Lon Tae Forest Ranger Station, Pha Kop, on the bark of an unidentified tree in coniferous forest, 1343 m, 17°11'13'' N, 101°33'58'' E, 12 November 2009, S . Meesim & K. Buaruang 599 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 51, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89.","Aptroot, A. (1988 b) Lichens of Madagascar: The Pyxinaceae (syn. Physciaceae). Cryptogamie, Bryologie et Lichenologie 9: 141 - 147."]}

Pyxine farinosa Kashiw., Bull. Natl Sci. Mus. , Tokyo, B, 3 (2): 67 (1977). Holotype:— MICRONESIA. Caroline Islands; Yap Island, 6 August 1939, F. Fujikawa 52 (TNS!). For further synonymy see Elix (2009). Thallus 2–5 cm wide, adnate. Lobes radiating, discrete to imbricate, plane to slightly convex, 0.3–1.2 mm wide, subtruncate at the apices. Upper surface white to yellowish white or yellowish grey, sparsely pruinose at the lobe tips or epruinose; polysidiangia and isidia absent. Pseudocyphellae ± distinct, marginal and laminal, irregular or becoming reticulate. Soralia laminal or occasionally apical, distinctly elevated, orbicular, subglobose; soredia farinose, white to yellow-white. Medulla upper part orange to orange-red, lower part white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia rare, obscurascens - type, 0.5–2.0 mm wide; disc epruinose. Internal stipe distinct, white, K-, P-. Ascospores 13–17 × 6–8 µm. Pycnidia not seen. Chemistry: Cortex K+ yellow, C-, KC-, P+ yellow, UV-; medulla K-, C-, KC-, P-; containing atranorin (minor), chloroatranorin (minor), unknown terpenes (minor), unknown pigment (minor). Distribution and habitat:—Corticolous (especially on palm trees) in coastal forests up to 300 m in South and East Asia, Australia, Papua New Guinea, Philippines, Polynesia and Micronesia. Notes:—This species is characterized by the white to yellowish white or yellowish grey upper surface with laminal, elevated, orbicular soralia, the orange to orange-red upper part of the medulla, a colourless stipe and the presence of atranorin and terpenes. Pyxine fallax is similar, but it has a uniformly white medulla, and it contains norstictic acid and testacein. For differences from P. daedalea see key., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 46, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495

Pyxine berteriana (Fée) Imshaug, Trans. Amer. Microsc. Soc. 76: 254 (1957). Type:— Circinaria berteriana Fée, Essai cryptog. écorces officin.: 128 (1825). Holotype:—Insula Martinicensi [Martinique], ad corticem Quassiae excelsae (G!) — Pyxine meissneri Tuck. ex Nyl., Ann. Sci. Nat.; Bot., sér. 4, 11: 205 (1859). Lectotype (selected by Imshaug 1957: 254):— Cuba, C. Wright, Wright, Lich. ins. Cubae exs. 95 (FH!) = Pyxine cocoës var. meissneri (Tuck. ex Nyl.) Tuck., Proc. Amer. Acad. Arts Sci. 12: 166 (1877).— Pyxine meissneri var. rinodinoides Vain., Ann. Acad. Sci. Fenn. A (6): 69. Holotype:—[Lesser Antilles] Insula S. Jan, ad corticem arboris prope Coral-Bay (Caroline), 22 February 1906, Raunkiaer 443 (C, fide Jungbluth 2010). Thallus corticolous, 5–9 cm wide, adnate to tightly adnate. Lobes radiating, discrete to ± contiguous, plane to convex, but often slightly concave towards the tips, 0.3–1.3 mm wide, subrotund at the apices. Upper surface white to light grey, greenish grey or yellow-grey, sparsely pruinose at the lobe tips (pruina punctiform) or epruinose; polysidiangia, isidia and soredia absent. Pseudocyphellae distinct, marginal and laminal, subreticulate. Medulla upper part: (pale) yellow, lower part: white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia common, cocoës - type, 0.5–2.0 mm wide; disc epruinose. Internal stipe distinct, white, K-, P-, C-, KC-. Ascospores 15–20 × 6–7 μm. Pycnidia rare, immersed in small warts on thallus with an exposed black ostiolum; conidia 4–5 × 1–1.1 µ m. Chemistry: Cortex K-, UV + yellow; medulla upper part K+ distinctly orange to yellow, C-, KC-, P+ orange, lower part: all spot tests negative; containing lichexanthone (major), terpenes (major or minor), unknown pigment (minor or trace). Distribution and habitat:—Pantropical and common in North, Central and South America, Africa, Australia, New Caledonia and the Galapagos Islands. Notes:—This species is characterized by the presence of cortical lichexanthone, the absence of vegetative propagules, the yellow upper medulla (white below) and apothecia of the cocoës- type and a white internal stipe. It is often confused with P. australiensis (Kalb 1994:61), but that species has narrower lobes, a uniformly yellow medulla and apothecia of the obscurascens - type. Also similar is the Neotropical P. endolutea Kalb (Kalb 1987: 49), but this species is readily separated by the presence of atranorin (UV-). Although this species has not yet been reported for Thailand, we have added a description for convenience. Most of the herbarium material which was labelled P. berteriana was revised to P. cognata which is easily separated by its orange medulla (upper part) and obscurascens - type apothecia., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on pages 37-38, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Imshaug, H. A. (1957) The lichen genus Pyxine in North and Middle America. Transactions of the American Microscopical Society 76: 246 - 269.","Jungbluth, P. (2010) Estudos taxonomicos em Physcia (Schreb.) Michx. e Pyxine Fr. (Physciaceae, Ascomycota). PhD thesis (unpublished), Sao Paulo, 227 pp.","Kalb, K. (1994) Pyxine species from Australia. Herzogia 10: 61 - 69.","Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}

5. Pyxine cocoës (Sw.) Nyl. , Mém. Soc. Imp. Sci. Nat. Cherbourg 5: 108 (1857). Type:— Lichen cocoës Sw., Nova gen. spec. plant.: 146 (1788);— Coccocarpia pellita var. cocoës (Sw.) Zahlbr., Cat. lich. univ. 3: 286 (1925). Holotype:— Jamaica, on Cocos, Swartz (S!; isotype:—H-ACH 379!). Thallus usually corticolous, rarely also saxicolous, 1–3 (–10) cm wide, adnate. Lobes radiating, discrete to ± contiguous, plane to slightly convex or concave, 0.4–0.8 mm wide, subrotund to truncate at the apices. Upper surface white to yellowish white, pale yellow-brown or grey, patchily pruinose; pruina glistening; polysidiangia and isidia absent. Pseudocyphellae usually restricted to the margins, rarely laminal or becoming reticulately confluent, developing into soralia with age. Soralia marginal and laminal, orbicular to linear, ± coalescing into extensive patches; soredia granular. Medulla uniformly white. Lower surface black in the centre, paler towards the periphery; rhizines ± dense, furcate. Apothecia rare, cocoës - type, 0.4–1.4 mm wide; disc epruinose. Internal stipe distinct, reddish brown in the upper part and K + purple, P-; lower part white, K-, P-. Ascospores 15–18 × 6–7 μm. Conidia bacilliform, 3–4 × ca. 1 μm. Chemistry: Cortex K-, UV + yellow; medulla K-, C-, KC-, P-; containing lichexanthone (major), ± unknown terpenes (traces). Distribution and habitat:—This very common, mainly pantropical lichen grows on rock, wood and bark in dry dipterocarp and mixed deciduous forests. Notes:— Pyxine cocoës is characterized by the cortical lichexanthone, marginal pseudocyphellae developing into coalescing soralia with rather coarsely granular soredia, the white medulla and apothecia of the cocoës - type with an internal stipe that is reddish brown (K+ purple) in the upper part and the patchy pruina covering large parts of the lobe tips. Selected material from Thailand examined:— Chiang Mai; Mae Rim, along a big pond called ' Huay Tueng tao Reservoir', ca. 6 km NNW of Chiang Mai, on bark in a dry dipterocarp forest, 360 m, 18°52'11'' N, 98°56'28'' E, 16 March 2008, K . Kalb (hb. Kalb 37099).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on rocks in mixed deciduous forest, 397 m, 17°20'49'' N, 101°33'29'' E, 28 July 2009, S . Meesim & K. Buaruang MSPL 308, 309 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Ho Forest Ranger Station, on the trunk of Bambusa bambos in mixed deciduous forest, 569 m, 17°07'13'' N, 101°34'30'' E, 29 July 2009, S . Meesim & K. Buaruang 427 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, Khao Noi, on the bark of Wrightia lecomtei in dry dipterocarp forest, 300 m, 14°10' N, 101°30' E, 2 April 1998, P . Mongkolsuk 12917 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, the trail "Kang Kao" cave Khao Lookchang site, on the bark of Pterospermum aceritolium in mixed deciduous forest, 1100 m, 14°32' N, 101°40' E, 6 April 1999, P . Mongkolsuk 13781 (RAMK).— Chonburi; Panutnikom district, Tung Kwang, on the trunk of Phyllanthus acidus in plantation, 50 m, 13 April 2001, P . Mongkolsuk 30000 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 41, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495

13. Pyxine philippina Vain. , The Philipp. Journ. Sci., C, 8 (2): 110 (1913). Lectotype (selected by Kashiwadani 1977: 163):— PHILIPPINES. Luzon; Benguet, ad corticem arborum frondosarum, May 1911, E. D. Merrill 7934 (TUR-VAIN 8707!).— Pyxine glaucescens Vain. The Philipp. Journ. Sci., C, 8: 109 (1913). Holotype: PHILIPPINES. Comiran Island, Sulu Sea, ad corticem arboris frondosae, September 1910, E. D. Merrill s.n. (TUR-VAIN 8705!). Thallus corticolous, 2–5 cm wide, adnate to loosely adnate. Lobes radiating or irregular, contiguous to imbricate, plane to partly convex, 0.5–1.4 mm wide, subrotund at the apices. Upper surface whitish to pale greenish grey or yellow-grey, faintly pruinose towards the lobe tips or epruinose; polysidiangia, isidia and soralia absent. Pseudocyphellae marginal and laminal, irregular, linear or becoming reticulate. Medulla uniformly white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia common, obscurascens - type, 1–3 mm wide; disc black, not pruinose. Internal stipe distinct, white to pale brown, K-, P-. Ascospores 15–23 × 6–8 µm. Pycnidia not seen. Chemistry: Cortex K+ yellow, C-, KC-, P+ yellow, UV-; medulla K+ yellow → red, C-, P+ orange; containing atranorin (minor), chloroatranorin (minor), norstictic acid (major), testacein (minor), unknown terpenes showing the same array as P. retirugella Nyl. (minor). Distribution and habitat:—Very rare on bark in a mixed deciduous forest. Also in Australia, Asia, Africa, South America and the Pacific (Hawaiian Islands and Micronesia). Notes:— Pyxine philippina is characterized by the adnate thallus, the absence of soredia, isidia and polysidiangia, the uniformly white medulla, cortical atranorin and obscurascens - type apothecia with an internal stipe that is brownish in the upper part. P. schmidtii Vain., which also produces testacein, is morphologically very similar in lacking any vegetative propagules, but can easily be distinguished by the absence of norstictic acid in the medulla. P. profallax is morphologically also very similar, but this species differs by a different suite of terpenes. Material from Thailand examined:— Pitsanulok; Phu Hin Rongkla National Park, air raid shelter point, on the bark of unidentified tree in mixed deciduous forest, 560 m, 18 September 2006, S. Meesim 21506 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 48, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kashiwadani, H. (1977) The genus Pyxine (Lichens) in Papua New Guinea. Bulletin of the National Science Museum, Tokyo, B, 3 (2): 63 - 70."]}

19. Pyxine subcinerea Stirt. , Trans. Proc. New Zealand Inst. 30: 397 (1898). Holotype:— AUSTRALIA. Queensland; [locality unknown], F. M. Bailey 22; (BM!). For further synonymy see Kalb (1987). Thallus 3–8 cm wide, adnate to loosely adnate, subdichotomously lobate. Lobes radiating, contiguous to imbricate, plane but often slightly concave towards the tips, 0.3–0.7 (–1.0) mm wide, subrotund to subtruncate at the apices. Upper surface yellowish grey to grey to brownish grey or olive-grey, pruinose; pruina densely punctiform towards the lobe tips or forming larger plates within the thallus, often glistening; polysidiangia and isidia absent. Pseudocyphellae distinct at the margins, often spreading laminally, rarely reticulate. Soralia marginal or submarginal and linear, then laminal and punctiform, often coalescing; soredia farinose. Medulla lemon yellow to yellow above, very thin below, white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, furcate. Apothecia common, obscurascens - type, 0.3–1.5 mm wide; disc epruinose. Internal stipe indistinct; upper part red-brown, K + purple, P-; lower part white, K-, P-. Ascospores 13–20 × 6–8 µm. Pycnidia rare, laminal immersed in small warts in thallus; conidiospores bacilliform, 3–4 × 1 µm. Chemistry: Cortex K-, UV + yellow; medulla upper part K-, C-, KC-, P + orange, lower part K-, C-, KC-, P-; containing lichexanthone (major), terpenes (major or minor), unknown pigment (minor or trace). Distribution and habitat:—Occurs on rock, wood and bark in dry dipterocarp, lower montane and mixed deciduous forests. This mainly pantropical species often extends into subtropical and temperate regions. Notes:— Pyxine subcinerea is characterized by the marginal soralia with farinose soredia, the yellow medulla, obscurascens - type apothecia and the presence of lichexanthone in the upper cortex. Pyxine meissnerina is very similar in its overall habitus, but this species is readily separated by a P- upper part of the medulla and a UV- cortex which contains atranorin. Material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on Quercus mespilifolioides in an open, dry dipterocarp forest, 570 m, 18°53'54'' N, 98°51'38'' E, 1 July 1994, P . Mongkolsuk 1473 (RAMK 17619, 17633, 17634).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on the trunk of Saraca indica in mixed deciduous forest, 397 m, 17°20'49'' N, 101°33'29'' E, 28 July 2009, S . Meesim 355 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Luang Wildlife Research Station, on the bark of an unidentified tree in lower montane rainforest, 923 m, 17°20'34'' N, 101°30'26'' E, 12 November 2008, S . Meesim & K. Buaruang 186 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Phu Ho Forest Ranger Station, on the bark of an unidentified tree in lower montane rainforest, 923 m, 17°20'34'' N, 101°30'26'' E, 12 November 2009, S . Meesim & K. Buaruang MS 0186 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Nam Chan Forest Ranger Station, on the trunk of Saraca indica in mixed deciduous forest, 397 m, 17°20'49'' N, 101°33'29'' E, 28 July 2009, S . Meesim & K. Buaruang MSPL 355 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on pages 51-52, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}

12. Pyxine petricola Nyl. , in J. M. Crombie, Journ. Bot. 14: 263 (1876). Lectotype (designated here):—ISLAND OF RODRIGUEZ, 9 December 1874, I. B. Balfour 2391 (Transit of Venus Expedition) (BM!, isolectotypes H-NYL 31754! and H-NYL 2336, insertum post mortem). — Pyxine subvelata Stirt., Trans. Proc. New Zealand Inst. 30: 396 (1898). Holotype:— AUSTRALIA. Queensland; Jimbour, June 1895, F. M. Bailey s.n. (BM!). For further synonymy see Kalb (1987). Thallus corticolous or saxicolous, 2–6 cm wide, adnate to loosely adnate. Lobes radiating or irregular, contiguous to imbricate, plane to concave, rarely convex, 0.7–1.2 mm wide, subrotund to truncate at the apices. Upper surface white to pale grey, greenish grey or yellowish grey, pruinose in patches or seldom only punctiform, matt or glossy; polysidiangia, soralia and isidia absent. Pseudocyphellae ± distinct, marginal and laminal, usually restricted to the peripheral parts of the lobes, rarely becoming reticulate. Medulla uniformly white. Lower surface black in the centre, paler towards the margin; rhizines dense, short, simple or ± furcate. Apothecia cocoës - type, 0.5–1.5 mm wide; disc epruinose. Internal stipe distinct; upper part rose to orangered, K+ purple, P-; lower part white, K-, P-. Ascospores 14–20 × 5–9 µm. Conidia bacilliform, 3–4 × 1 µm. Chemistry: Cortex K-, UV+ yellow; medulla K-, C-, KC-, P-; containing lichexanthone (major), terpenes (major or minor). Distribution and habitat:—Occurs on rock and bark in dry dipterocarp and mixed deciduous forests. Also in Australia, Asia, Africa, South America and the Pacific (Hawaiian Islands and Micronesia). Notes:— Pyxine petricola is characterized by the adnate to loosely adnate thallus, the absence of soredia, isidia and polysidiangia, the white medulla, cortical lichexanthone and cocoës - type apothecia with an internal stipe that is reddish in the upper part. Material from Thailand examined:— Loei; Phu Luang Wildlife Sanctuary, Phu Ho Forest Ranger Station, on the bark of Sandoricum koetjape in mixed deciduous forest, 569 m, 17°07'13'' N, 101°34'30'' E, 29 July 2009, MSPL 375 (RAMK).— Sakonnakon; Ratchapat Sakonnakon, on the trunk of Mangifera indica in dry dipterocarp forest, 300 m, 19 March 1998, W . Khumtim 11889 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 47, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}

7. Pyxine coralligera Malme , Bih. Kongl. Svensk. vetensk.-akad. handl. 23, afd. 3 (13): 40 (1897). Type:— BRAZIL. Mato Grosso; Serra da Chapada, near São Jeronymo, ad rupes apricas, 3 June 1894, G. O. A. Malme [Exped. Prim. Regnell. Lichens 2749c]; lectotype selected by Swinscow & Krog 1975a: 53. (S!). For synonymy see Kalb (1987).— Pyxine palniensis D. D. Awasthi, Phytomorphology 30 (4): 374 (1982). Holotype: INDIA. Tamil Nadu; Palni hills, Perumalmalai area near Perumal peak, on rock, 2000 m, 17 December 1970, D. D. Awasthi 70.1092 (LWU!). Thallus corticolous and saxicolous, 3–10 cm wide, loosely adnate. Lobes radiating, discrete to ± imbricate, plane to slightly convex, 0.5–1.0 mm wide, subrotund to truncate at the apices. Upper surface pale to dark grey, weakly pruinose towards the lobe tips or epruinose; true isidia and soralia absent. Pseudocyphellae laminal, irregular to linear or becoming ± reticulate subapically. Polysidiangia mainly laminal, occasionally marginal, scattered or clustered, nodular to subcylindrical or clavate, short, ± branched, pustulate, bursting apically to release corticate fragments but no soredia. Medulla (upper part) creamy yellow to yellow-orange or yellow-brown, lower part white. Lower surface black in the centre, paler towards the margin; rhizines ± dense, simple or irregularly branched. Apothecia rare, obscurascens type, 0.5–1.0 mm wide; disc epruinose. Internal stipe distinct, pale yellow to yellow-orange or yellow-brown, K + greenish, P + dirty orange-brown. Ascospores 16–20 × 5–7 μm. Pycnidia not seen. Chemistry: Cortex K + yellow, C-, KC-, P + yellow, UV-; medulla upper part: K-, C-, P + orange, lower part: all spot tests negative; containing atranorin (minor), chloroatranorin (minor), testacein (major), unknown terpenes (minor). Distribution and habitat:—A pantropical species, very common on bark and rocks, in tropical rainforests, dry dipterocarp forests and lower montane scrubs. Also in Australia, Africa, Asia, South America, Cuba and Papua New Guinea. Notes:— P. coralligera is characterized by the polysidiangia, cortical atranorin, the pale yellow to yelloworange or yellow-brown medulla, obscurascens - type apothecia, an internal stipe that is concolorous with the medulla, and by the presence of medullary testacein and a characteristic array of terpenes. It is morphologically similar to P. retirugella, but the latter differs in containing additional norstictic acid and a different suite of terpenes. P. daedalea is morphologically similar and shows the same array of terpenes on TLC plates. But it is easily distinguished by the presence of laminal soralia. Co-chromatography of this species on TLC plates in solvents A, B' and C together with the morphologically identical P. palniensis shows that the two species have an identical array of terpenes. The only difference is a very low concentration of testacein in the latter so that no P-reaction could be observed. Material from Thailand examined:— Chiang Mai; Queen Sirikit Botanic Garden, on the bark of Castanopsis costata in dry dipterocarp forest forest, 630 m, 18°50' N, 98°46' E, 26 January 1994, K . Boonpragob et al. 719, 731 (RAMK).— Chiang Mai; Queen Sirikit Botanic Garden, on the bark of Syzygium claviflora, dry dipterocarp forest, 750 m, 1 July 1994, K . Boonpragob 2670, 2671, 2689 (RAMK).— Chiang Mai; medicinal garden, on bark in a ± open Cinchona plantation near Doi Suthep-Pui National Park, 1085 m, 18°48'22'' N, 98°54'53'' E, 17 March 2008, K . Kalb (hb. Kalb 36780, 36788).— Loei; Phu Luang Wildlife Sanctuary, a way from in front of Head Quarter of Phu Luang Wildlife Sanctuary 700 m, on the bark of an unidentified tree in tropical rainforest, 700 m, 17°20'18'' N, 101°30'32'' E, 20 May 2009, S . Meesim & K. Buaruang MSPL 080 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, a way from in front of Head Quarter of Phu Luang Wildlife Sanctuary 500 m, on the rocks in dry dipterocarp forest, 700 m, 17°14'33'' N, 101°18'26'' E, 21 May 2009, S . Meesim & K. Buaruang MSPL 119 (RAMK).— Loei; Phu Luang Wildlife Sanctuary, Helicopter Apron, on the rocks in lower montane scrub, 1487 m, 17°16'23'' N, 101°31'08'' E, 25 Much 2009, S . Meesim 86 (RAMK).— Loei; Nahaew National Park, on the rocks in hill evergreen forest, 1260 m, 12 July 1995, P . Mongkolsuk et al. 4218 (RAMK).— Loei; Nahaew National Park, on the rocks in dry dipterocarp forest, 900 m, 5 November 1995, P . Mongkolsuk et al. 6616, 6618 (RAMK).— Loei; Nahaew National Park, on rocks in dry dipterocarp forest, 670 m, 6 November 1995, P . Mongkolsuk et al. 6767 (RAMK)— Nakhon Ratchasima; Khao Yai National Park, Former Golf Course, on the bark of an unidentified tree in shrub forest, 750 m, 14°25' N, 101°23' E, 5 December 1999, P . Mongkolsuk 15842 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, Former Golf Course, on the bark of an unidentified tree in tropical rainforest, 750 m, 14°25' N, 101°23' E, 5 December 1999, P . Mongkolsuk 15990 (RAMK).— Nakhon Ratchasima; Khao Yai National Park, the trail kilometer no. 33 "Nong Pakchee", on the bark of an unidentified tree in tropical rainforest, 800 m, 14°20' N, 101°39' E, 23 May 2000, P . Mongkolsuk et al. 15996, 16028, 19511 (RAMK)., Published as part of Mongkolsuk, Pachara, Meesim, Sanya, Poengsungnoen, Vasun & Kalb, Klaus, 2012, The lichen family Physciaceae in Thailand-I. The genus Pyxine, pp. 32-54 in Phytotaxa 59 (1) on page 43, DOI: 10.11646/phytotaxa.59.1.2, http://zenodo.org/record/5061495, {"references":["Swinscow, T. D. V. & Krog, H. (1975 a) The genus Pyxine in East Africa. Norwegian Journal of Botany 22: 43 - 68.","Kalb, K. (1987) Brasilianische Flechten. 1. Die Gattung Pyxine. Bibliotheca Lichenologica 24: 1 - 89."]}