The Species

Graphis bungartzii Barcenas-Peña, Lücking, Herrera-Campos & R. Miranda sp. nov.

MycoBank No.: MB804537

Differing from Graphis elegans in the erumpent lirellae with lateral thalline margin and the smaller ascospores, from G. filiformis in the labia becoming striate, and from G. schiffneri in the laterally carbonized excipulum.

Type: Mexico, Jalisco, Municipio La Huerta, Cuixmala-Chamela Biosphere Reserve, Chamela Biological Station (UNAM), near Hornitos stream, 19°30′56·1″N, 105°02′7·2″W, 59 m, seasonally tropical dry forest, on bark of Croton sp., June–July 2011, Barcenas Peña 1247 (MEXU—holotype); same locality and date, on bark of Thouinia sp., Barcenas Peña 1248, 1249 (MEXU—paratypes).

(Fig. 2 A–D)

Fig. 2. Graphis bungartzii (holotype). A, habit and flexuose, branched, erumpent lirellae; B, concealed disc; striate labia, and thick lateral thalline margin; C, section through a lirella, laterally carbonized excipulum, and not inspersed hymenium; D, mature transversely septate ascospore. Scales: A & B=1 mm; C=100 µm; D=20 µm.

Thallus corticolous, 2–5 cm diam., up to 150 µm thick, continuous; surface smooth, pale greenish grey; prothallus absent. Thallus in section c. 30 µm with cartilaginous upper cortex, irregular algal layer, and large clusters of calcium oxalate crystals.

Apothecia lirelliform, flexuose, branched, erumpent, with lateral thalline margin, 1·5–3·5 mm long, 0·1–0·2 mm wide, 0·03–0·05 mm high; disc concealed; proper margin thin, labia striate, black; thalline margin thick, of the same colour as the thallus. Excipulum entire, laterally carbonized, 15–45 µm wide, black, basally orange-brown; laterally covered by corticated algiferous thallus including large clusters of calcium oxalate crystals; hypothecium prosoplectenchymatous, 20–27 µm high, pale orange-brown; hymenium 90–110 µm high, colourless, not inspersed, paraphyses hyaline, unbranched; epithecium granulose, 5·0–7·5 µm high, brown. Asci fusiform, 70–90×10–20 µm. Ascospores 8 per ascus, ellipsoid to fusiform, transversely 5–11-septate, 25–38×5–8 µm, 3–4 times as long as wide, colourless.

Chemistry (TLC)

Norstictic and connorstictic acids (thallus K+ yellow turning red, in sections with yellow outflow with precipitating red needles).

Etymology

This new species is dedicated to Frank Bungartz for his support and encouragement offered to Alejandrina Barcenas-Peña and Ricardo Miranda González, and his interest in Mexican lichens.

Distribution and ecology

The type locality, at an elevation of 59 m, is characterized by a tree mixture of the transition between dry forest and the more humid and taller semi-deciduous forest at arroyos (Lott et al. Reference Lott, Bullock and Solís-Magallanes1987). In this locality, three specimens of the new species were found growing on the bark of Croton sp. and Thouinia paucidentata. The additional specimens cited were collected in different sites, between 2 and 13·5 km from the type locality in dry forest at an altitude of 50–300 m. Although morphologically G. bungartzii could be placed in the G. duplicata group, its known ecology contrasts with that of most other species in that group, which have typically been reported from higher elevations (750–1500 m) and from evergreen rainforests with a slight to distinct dry season (Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008).

Remarks

Currently there is only one other species known with labia becoming striate, a laterally carbonized excipulum, clear hymenium, small, transversely septate ascospores, and the chemical combination of norstictic and connorstictic acids, viz. Graphis elegans (Sm.) Ach. (Lücking et al. Reference Lücking, Archer and Aptroot2009b ). That species differs from the new one in the erumpent to prominent lirellae lacking a thalline margin and the larger, thick-walled ascospores with 9–13 septa, although it shares the same secondary substances. Graphis filiformis Adaw. & Makahija is also similar to Graphis bungartzii, but differs in having longer, irregularly to radiately branched lirellae with entire labia, and it lacks connorstictic acid. The eastern paleotropical Graphis schiffneri Zahlbr. agrees with the new species in most aspects but has a completely carbonized excipulum, thicker thalline margin, and only norstictic acid as the main secondary substance.

Specimens examined. Mexico: Jalisco: Municipio La Huerta, Cuixmala-Chamela Biosphere Reserve, Chamela Biological Station (UNAM), 100 m E from station buildings, between Chachalacas trail and Camino Antiguo Sur dirt road, 19°29′51·1″N, 105°02′30·2″W, 68 m, seasonally tropical dry forest, on bark of Thouinia paucidentata, 2009, Miranda 1088; 300 m W from Tejón trail at 600 m sign, 19°30′11·4″N, 105°02′53·8″W, 41 m, seasonally tropical dry forest, on bark of Thouinia paucidentata, 2010, Miranda 1774; Ejido Gargollo, S of Chamela Biological Station (UNAM), 19°24′17·9″N, 104°58′56·5″W, 57 m, seasonally tropical dry forest, on bark of Thouinia paucidentata, 2010, Miranda 4266 (all MEXU).

Graphis chamelensis Barcenas-Peña & Lücking sp. nov.

MycoBank No.: MB804538

Differing from Graphis emersa in the longer, partly branched lirellae, and from G. conferta in the norstictic acid chemistry.

Type: Mexico, Jalisco, Municipio La Huerta, Cuixmala-Chamela Biosphere Reserve, Chamela Biological Station (UNAM), near Hornitos stream, 19°30′52·6″N, 105°02′05·5″W, 59 m, seasonally tropical dry forest- semideciduous arroyo forest transition, on bark of Forchhammeria pallida, June–July 2011, Barcenas Peña 1239 (MEXU—holotype).

(Fig. 3A–D)

Fig. 3. Graphis chamelensis (holotype). A, habit and prominent lirellae; B, concealed disc; labia entire, and basal to lateral thalline margin; C, section through a lirella, entire, completely carbonized excipulum and not inspersed hymenium; D, mature transversely septate ascospore. Scales: A & B=1 mm; C=100 µm; D=20 µm.

Thallus corticolous, 0·5–1·0 cm diam., 40–80 µm thick, continuous; surface smooth, pale greenish grey; prothallus absent. Thallus in section with thick cartilaginous upper cortex, irregular algal layer, and large clusters of calcium oxalate crystals.

Apothecia lirelliform, flexuose, partly branched, prominent, with basal to lateral thalline margin, 1–3 mm long, 0·2–0·4 mm wide, 0·25–0·30 mm high; disc concealed; proper margin thick, labia entire, black. Excipulum entire, completely carbonized, 50–70 µm wide, black, basal internal parts brown; hypothecium prosoplectenchymatous, 20–30 µm high, colourless to pale yellowish; hymenium 80–100 µm high, colourless, clear, paraphyses unbranched; epithecium granulose, 10–25 µm high, olive-brown. Asci fusiform, 70–90×15–25 µm. Ascospores 4–8 per ascus, oblong, transversely 7–10-septate, 25–35×7–10 µm, 2–4 times as long as wide, colourless.

Chemistry (TLC)

Norstictic acid (thallus K+ yellow turning red, in sections with yellow outflow with precipitating red needles).

Distribution and ecology

The type material of Graphis chamelensis was collected in the same transitional vegetation as G. bungartzii, but on the bark of Forchhammeria pallida. The additional specimens were found growing on Bursera heteresthes and Guettarda elliptica in three different sites in dry forest at 48–95 m, 1–2 km from the type locality. Although G. chamelensis was found on a higher number of different phorophytes than G. bungartzii, it seems to grow in a more limited area, probably requiring more specific microclimate conditions. Although the species forms part of the G. nuda group, its ecological preferences contrast with those of other species in that group, which are more typically found at high altitudes (Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008).

Remarks

Graphis chamelensis is similar to G. emersa Müll. Arg., from which it differs mainly by the presence of longer, partly branched prominent lirellae with a sometimes basally developed thalline margin. Both species have the same chemistry (Lücking et al. Reference Lücking, Archer and Aptroot2009b ). Graphis conferta Zenker is morphologically like G. chamelensis; however, it has sessile lirellae without a thalline margin, slightly longer ascospores, and no secondary substances.

Additional specimens examined. Mexico: Jalisco: Municipio La Huerta, Cuixmala-Chamela Biosphere Reserve, Chamela Biological Station (UNAM), near Tejón trail, 19°30′25·6″N, 105°02′32·8″W, 48 m, seasonally tropical dry forest, on bark of Forchhammeria pallida, 2011, Barcenas Peña 2000; 19°30′29·3″N, 105°02′29·8″W, 83 m, on bark of Bursera heteresthes, Barcenas Peña 2001; 19°30′34·4″N, 105°02′21″W, 95 m, on bark of Guettarda elliptica, Barcenas Peña 2002 (all MEXU).

Graphis rosae-emiliae Barcenas-Peña & Lücking sp. nov.

MycoBank No.: MB804539

Differing from Graphis syzygii in the larger ascospores and the lack of secondary substances, and from G. aurita in the inspersed hymenium and larger ascospores.

Type: Mexico, Veracruz, Municipio San Andrés Tuxtla, Los Tuxtlas Biosphere Reserve, Los Tuxtlas Tropical Biology Station, near Laguna Zacatal, 18°35′07·1″N, 95°05′23·4″W, 242 m, lowland rainforest, on bark of Orthion oblanceolatum, May 2011, Barcenas Peña 1235 (MEXU—holotype); same locality and date, on bark of unidentified phorophyte, Barcenas Peña 1254 (MEXU—paratype).

(Fig. 4A–D)

Fig. 4. Graphis rosae-emiliae (holotype). A, habit and flexuose, unbranched, erumpent lirellae; B, concealed disc; labia striate, lateral thalline margin; C, section through a lirella, completely carbonized excipulum, inspersed hymenium, and calcium oxalate crystals visible with polarized light; D, mature transversely septate ascospore. Scales: A & B=1 mm; C=100 µm; D=20 µm.

Thallus corticate, 1–3 cm diam., 120–180 µm thick, continuous; surface smooth, greenish grey; prothallus absent. Thallus in section with cartilaginous upper cortex, irregular algal layer, and large clusters of calcium oxalate crystals.

Apothecia lirelliform, flexuose, unbranched, erumpent, with lateral thalline margin, 1–3 mm long, 0·25 mm wide, 0·1 mm high; disc concealed; proper margin thick, labia striate, black. Excipulum completely carbonized, 70–90 µm wide, black; hypothecium prosoplectenchymatous, 20–25 µm high, colourless; hymenium 80–90 µm high, colourless, inspersed (type A according to Lücking Reference Lücking2009), paraphyses unbranched; epithecium granulose, 10–15 µm high, colourless to pale yellowish. Asci fusiform, 70–90×10–20 µm. Ascospores 8 per ascus, oblong, transversely 9–13-septate, 45–55×5–10 µm, 5–9 times as long as wide, colourless.

Chemistry (TLC)

No substances detected (all spot tests negative).

Distribution and ecology

The type locality is characterized by an exuberant tropical rainforest with high humidity, at an elevation of 242 m along the east side of San Martín Tuxtla volcano. The type material of G. rosae-emiliae was found on the bark of canopy branches of Orthion oblanceolatum. The additional specimen was also found in the canopy, in a different locality and on a different date, 21 km from the type locality, at 345 m, on the northern slope of the Santa Marta Volcano.

Remarks

Graphis rosae-emiliae is characterized by its unbranched, erumpent lirellae with striate labia and lateral thalline margin, the completely carbonized excipulum, and the inspersed hymenium (type A), as well as transversely septate ascospores and lack of secondary metabolites. Other species with a similar lirella type, inspersed hymenium, and transversely septate ascospores are G. leucaenae Aptroot, G. inspersolongula Aptroot, and G. syzygii Aptroot (Lücking et al. Reference Lücking, Archer and Aptroot2009b ). Both G. leucaenae and G. inspersolongula have much larger ascospores and produce norstictic acid. Graphis syzygii Aptroot differs in its smaller ascospores and the presence of stictic acid. Finally, G. cinerea Fée shares the same chemistry as G. rosae-emiliae but it has very large ascospores (100–200×18–30 µm) and a different type of inspersion, among other morphological differences. Morphologically and chemically similar is G. aurita Eschw., but it lacks inspersion and its ascospores are smaller.

Specimen examined. Mexico: Veracruz: Municipio Catemaco, Los Tuxtlas Biosphere Reserve, Selva del Marinero, Ejido Adolfo López Mateos, 18°26′0·5″N, 94°57′44·1″W, 345 m, lowland rainforest, on bark of unidentified phorophyte, 2009, Barcenas Peña 1170 (MEXU).

Updates to the world key to species of Graphis

The publication by Lücking et al. (Reference Lücking, Archer and Aptroot2009b ) of a first global working key to the genus Graphis, as redefined by Staiger (Reference Staiger2002) and Lücking (Reference Lücking2009), has boosted the discovery and description of new species. Including the three species introduced here, no less than 41 species have been described or reinstated in the four years following publication of the key, representing an increase of 12·4% over the 330 species treated in the key. Of these, six are from North America (Lendemer Reference Lendemer2010; Lücking et al. Reference Lücking, Seavey, Common, Beeching, Breuss, Buck, Crane, Hodges, Hodkinson and Lay2011; Seavey & Seavey Reference Seavey and Seavey2011), four from Mexico (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011; this paper), one from Central America (Costa Rica: Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012), nine from South America (Peru: Rivas Plata & Lücking Reference Rivas Plata and Lücking2013; Brazil: Dal-Forno & Eliasaro Reference Dal-Forno and Eliasaro2010; Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011; Cáceres et al. Reference Cáceres, Santos Vieira, Santos De Jesus and Lücking2012), three from Europe (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011), 16 from Asia (India: Jagadeesh Ram & Sinha Reference Jagadeesh Ram and Sinha2009; Chitale et al. Reference Chitale, Makhija and Sharma2011; Sharma & Khadilkar Reference Sharma and Khadilkar2011; Sri Lanka: Weerakoon et al. Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbsch2012; Cambodia: Nakanishi et al. Reference Nakanishi, Kashiwadani, Futagami and Moon2010; Moon et al. Reference Moon, Nakanishi, Futagami and Kashiwadani2011; South Korea: Joshi et al. Reference Joshi, Lücking, Yamamoto, Wang, Koh and Hur2010; Moon et al. Reference Moon, Nakanishi and Kashiwandani2012; China: Jia & Wei Reference Jia and Wei2009; Jia Reference Jia2011; Indonesia: see key entries below), one from Fiji (Lücking et al. Reference Lücking, Lumbsch, Von Konrat and Naikatini2010), and one pantropical species (this paper: see key entries below). We expect that this trend will continue in the near future; for the Graphis scripta group alone, Lücking (Reference Lücking2012) predicted nearly 50 missing species based on the data from the published key. Some nomenclatural and taxonomic changes were also proposed, the scope of which are summarized below.

Since the publication of a complete, updated key would go beyond this paper and would also be futile in the light of possibly many more species being described in the near future, for each of the newly established species or nomenclatural or taxonomic changes, we provide a detailed entry based on the key published previously (Lücking et al. Reference Lücking, Archer and Aptroot2009b ). For the moment, we apply the concept of Graphis sensu Staiger (Reference Staiger2002) and Lücking (Reference Lücking2009), although it has been shown that this encompasses two genera (Rivas Plata et al. Reference Rivas Plata, Hernández, Lücking, Staiger, Kalb and Cáceres2011). However, at present the second genus, which has tentatively been named Allographa (Rivas Plata et al. Reference Rivas Plata, Hernández, Lücking, Staiger, Kalb and Cáceres2011), cannot be named formally until the type species of Hemithecium has been sequenced. The reason for this is that the name Allographa is invalid and must be validated, but the type species of Hemithecium potentially belongs in that clade, providing a valid name that would then have priority.

One species, Graphis parvicarpa B. O. Sharma & Khadilkar (Sharma & Khadilkar Reference Sharma and Khadilkar2011), turned out to belong in the genus Pallidogramme. The apical carbonization, as shown in the excellent illustrations in the protologue, is not part of the excipulum but of the epithecium. The type of hymenial inspersion agrees with Pallidogramme, and nearly hyaline ascospores are often found in that genus. We therefore propose the following combination:

Pallidogramme parvicarpum (B. O. Sharma & Khadilkar) Lücking comb. nov.

MycoBank No.: MB804862

Graphis parvicarpa B. O. Sharma & Khadilkar, Lichenologist 43: 487 (2011).

Names that were validated or changed in nomenclature or taxonomic concept are:

• Graphis arbusculaeformis (Vain.) Lücking (Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012); validated.

• Graphis betulina (Pers.) Ach. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011); replaces G. scripta (L.) Ach. auct. p.p.

• Graphis caribica Lücking (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011); validated.

• Graphis inustuloides Lücking (Lücking & McCune Reference Lücking and McCune2012); replaces G. britannica sensu Staiger auct.

• Graphis macrocarpa (Pers.) Röhl. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011); replaces G. scripta (L.) Ach. auct. p.p.

• Graphis norvestitoides Sutjaritturakan (Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012); validated and taxonomy corrected (see key below, Group 18, couplet 3 and Group 20, couplet 1).

• Graphis pedunculata Bungartz & Aptroot (Bungartz et al. Reference Bungartz, Lücking and Aptroot2010); validated.

• Graphis pulverulenta (Pers.) Ach. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011); replaces G. scripta (L.) Ach. auct. p.p.

• Graphis rongklaensis Sutjaritturakan (Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012); validated.

The following are updates to the key entries in the world key to Graphis (Lücking et al. Reference Lücking, Archer and Aptroot2009b ), for 41 newly described or reinstated species from 2009 to 2013:

• Group 1, couplet 10: Excipulum laterally carbonized; hymenium clear; norstictic acid; ascospores muriform; 50–100×20–30 µm; USA. ... Graphis hodgesiana Lendemer (Lendemer Reference Lendemer2010)

• Group 1, couplet 10: Excipulum laterally carbonized; hymenium clear; norstictic acid; ascospores submuriform; 20–35×10–15 µm; USA. ... Graphis tamiamiensis Lendemer (Lendemer Reference Lendemer2010)

• Group 2, couplet 2: 2-methoxy-psoromic acid; thallus sorediate; ascospores 30–35×7–8 µm; Cambodia ... Graphis taneina M. Nakan. et al. (Moon et al. Reference Moon, Nakanishi, Futagami and Kashiwadani2011)

• Group 3, couplet 3: Lirellae prominent, with complete thalline margin, elongate and irregularly branched; India ... Graphis elevativerrucosa Chitale et al. ... (Chitale et al. Reference Chitale, Makhija and Sharma2011)

• Group 4, couplet 5: Lirellae prominent to sessile, with thick lateral thalline margin, very short and unbranched (dussii-morph); norstictic and salazinic acids; China ... Graphis paradussii Z. F. Jia (Jia Reference Jia2011)

• Group 4, couplet 35: Disc exposed, non-pruinose; lirellae with rounded ends; Europe ... Graphis macrocarpa (Pers.) Röhl. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011)

• Group 4, couplet 35: Disc exposed, pruinose or weakly pruinose; lirellae with acute ends and conspicuously thick lateral thalline margin; Europe ... Graphis betulina (Pers.) Ach. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011)

• Group 4, couplet 35: Disc exposed, pruinose; lirellae with acute ends and thin lateral thalline margin; Europe ... Graphis pulverulenta (Pers.) Ach. (Neuwirth & Aptroot Reference Neuwirth and Aptroot2011)

• Group 4, couplet 41: Graphis scripta (L.) Ach. s. str. would key out under G. pinicola Zahlbr. according to the characters given by Neuwirth & Aptroot (Reference Neuwirth and Aptroot2011); possibly the collections of G. pinicola from subtropical regions (with lateral thalline margin) represent G. scripta s. str., whereas tropical material (with apically thin complete thalline margin) would have to be named G. guineensis C. W. Dodge; this issue needs further study.

• Group 6, couplet 12: Disc exposed, non-pruinose; South Korea ... Graphis jejuensis K. H. Moon et al. (Moon et al. Reference Moon, Nakanishi and Kashiwandani2012)

  [In the original description (Moon et al. Reference Moon, Nakanishi and Kashiwandani2012), this species is given as carbonized apically or in the upper half, and thus far only G. pertricosa (Kremp.) A. W. Archer is known with such a character combination, differing in the muriform ascospores and norstictic acid; however, the illustrations provided by Moon et al. (Reference Moon, Nakanishi and Kashiwandani2012) suggest a laterally carbonized excipulum]

• Group 6, couplet 12: Disc concealed; labia non-pruinose; lirellae immersed in distinct, raised, pseudostromatic areas; USA ... Graphis elevata Seavey & J. Seavey (Seavey & Seavey Reference Seavey and Seavey2011)

• Group 7, couplet 6: Ascospores 70–100×15–20 µm; lirellae erumpent, with thick lateral thalline margin (subserpentina morph); Fiji ... Graphis collinsiae Lücking & Lumbsch (Lücking et al. Reference Lücking, Lumbsch, Von Konrat and Naikatini2010)

• Group 8, couplet 7: Lirellae prominent to sessile, with thick lateral thalline margin, very short and unbranched (dussii-morph); norstictic and salazinic acids; China ... Graphis paradussii Z. F. Jia (Jia Reference Jia2011)

• Group 8, couplet 10: Lirellae prominent, with basal thalline margin, irregularly branched; ascospores 35–65 µm long; USA ... Graphis brittoniae Seavey & J. Seavey (Seavey & Seavey Reference Seavey and Seavey2011)

• Group 8, couplet 15: Lirellae erumpent to prominent, lacking or with basal thalline margin; short and sparsely branched; thallus distinctly verrucose to almost isidiate; China ... Graphis guangdongensis Z. F. Jia & J. C. Wei (Jia & Wei Reference Jia and Wei2009)

• Group 8, couplet 17: Lirellae erumpent; thallus verrucose; Brazil ... Graphis invisibilis Dal-Forno & Eliasaro (Dal-Forno & Eliasaro Reference Dal-Forno and Eliasaro2010)

• Group 8, couplet 18: Lirellae stellately branched and clustered into pseudostromata; thallus isidiate; Brazil ... Graphis itatiaiensis Nelsen et al. (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011)

• Group 8, couplet 20: Thallus green; lirellae 1–5 mm long, lacking basal margin, with grey-black labia; Mexico ... Graphis marusae B. Peña & Lücking (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011)

• Group 8, couplet 29: Lichexanthone; disc exposed (handelii morph); South Korea ... Graphis flavopalmicola Y. Joshi et al. (Joshi et al. Reference Joshi, Lücking, Yamamoto, Wang, Koh and Hur2010)

• Group 8, couplet 36: Lirellae prominent, with basal to lateral thalline margin, elongate and irregularly branched; Mexico ... Graphis chamelensis Barcenas-Peña & Lücking (this paper)

• Group 8, couplet 42: Lirellae elongate to very long, prominent; thallus sorediate; Brazil ... Graphis lueckingii Dal-Forno & Eliasaro (Dal-Forno & Eliasaro Reference Dal-Forno and Eliasaro2010)

• Group 8, couplet 51: Lirellae irregularly branched, with apically thin complete margin; ascospores regularly 3-septate, 15–20×5–6 µm; Peru ... Graphis pitmanii Rivas Plata & Lücking (Rivas Plata & Lücking Reference Rivas Plata and Lücking2013)

• Group 8, couplet 51: Lirellae irregularly branched but not radiate, with very thin complete margin; ascospores 7–11-septate, 35–55×10–12 µm; Brazil ... Graphis paranaensis Dal-Forno & Eliasaro (Dal-Forno & Eliasaro Reference Dal-Forno and Eliasaro2010)

• Group 10, couplet 6: Lirellae prominent, with thick lateral thalline margin (marginata morph); ascospores 50–70 µm long; Sri Lanka ... Graphis srilankensis Weerakoon et al. (Weerakoon et al. Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbsch2012)

• Group 10, couplet 13: Lirellae prominent, with basal to lateral thalline margin and exposed disc; Peru ... Graphis apertoinspersa Rivas Plata & Lücking (Rivas Plata & Lücking Reference Rivas Plata and Lücking2013)

• Group 13, couplet 1: 2-methoxy-psoromic acid; thallus sorediate; ascospores 30–35×7–8 µm; Cambodia ... Graphis taneina M. Nakan. et al. (Moon et al. Reference Moon, Nakanishi, Futagami and Kashiwadani2011)

• Group 13, couplet 7: Ascospores medium-sized to large, 50–95×7–9 µm, 8 per ascus; labia non-pruinose; lirellae erumpent to prominent, lacking or with basal thalline margin, very long and irregularly to radiately branched; India ... Graphis neoraensis Jagadesh & G. P. Sinha (Jagadeesh Ram & Sinha Reference Jagadeesh Ram and Sinha2009)

• Group 13, couplets 8 and 9: Thallus distinctly verrucose but not isidiate; lirellae with verrucose lateral thalline margin; Sri Lanka ... Graphis allugallenensis Weerakoon et al. (Weerakoon et al. Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbsch2012)

• Group 13, couplet 14: Thallus white-grey; lirellae erumpent, labia non-pruinose; ascospores 40–60 µm long; USA. ... Graphis appendiculata Common & Lücking (Lücking et al. Reference Lücking, Seavey, Common, Beeching, Breuss, Buck, Crane, Hodges, Hodkinson and Lay2011)

• Group 13, couplet 20: Lirellae with lateral thalline margin; thallus white-grey; pantropical ... Graphis subtenella Müll. Arg.

• Group 13, couplet 20: Lirellae with basal thalline margin; thallus green-grey; Indonesia ... Graphis chlorotica A. Massal.

  [Revision of type material of G. chlorotica revealed that it differs morphologically from G. subtenella, which is why the latter is removed from synonymy and accepted as a separate species]

• Group 14, couplet 6: Ascospores medium-sized, 45–65×12–16 µm; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella morph); India ... Graphis panhalensis (Patw. & C. R. Kulk.) Chitale et al. (Chitale et al. Reference Chitale, Makhija and Sharma2011)

  [This species is close to G. parilis Kremp. but appears to differ by the narrower ascospores, a feature that needs to be evaluated with more material]

• Group 15, couplet 5: Ascospores small, 25–38×5–8 µm, 5–11-septate, not conspicuously thick-walled; lirellae erumpent, with lateral thalline margin; Mexico ... Graphis bungartzii Barcenas-Peña et al. (this paper)

• Group 16, couplet 1: Norstictic acid; lirellae prominent, with complete thalline margin (acharii morph); ascospores 70–95×25–35 µm; Cambodia ... Graphis cambodiensis M. Nakan. et al. (Nakanishi et al. Reference Nakanishi, Kashiwadani, Futagami and Moon2010)

• Group 16, couplet 1: Stictic acid; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella morph); ascospores 25–42×13–17 µm; India ... Graphis maharashtrana Chitale et al. (Chitale et al. Reference Chitale, Makhija and Sharma2011)

• Group 17, couplet 7: Ascospores very large (120–150 µm long); lirellae very short to rounded, prominent, with complete thalline margin; Brazil ... Graphis brachylirellata M. Cáceres & Lücking (Cáceres et al. Reference Cáceres, Santos Vieira, Santos De Jesus and Lücking2012)

• Group 17, couplet 11: Lirellae erumpent, lacking or with basal thalline margin; Sri Lanka ... Graphis dotalugensis Weerakoon et al. (Weerakoon et al. Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbsch2012)

• Group 17, couplet 15: Lirellae prominent, lacking thalline margin but labia grey-black; thallus green; Mexico ... Graphis marusae B. Peña & Lücking (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011)

• Group 18, couplet 3: Graphis norvestitoides Sutjaritturakan is incorrectly keyed out here since it has an inspersed hymenium (see below, Group 20, couplet 1); the correct species to be keyed out here is; Costa Rica ... Graphis pseudoaquilonia Lücking (Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012)

• Group 18, couplet 6: Ascospores 4–6 per ascus, 25–30×10–13 µm; norstictic acid; lirellae erumpent, with basal thalline margin; Brazil ... Graphis archeri Dal-Forno & Eliasaro (Dal-Forno & Eliasaro Reference Dal-Forno and Eliasaro2010)

• Group 19, couplet 1: Ascospores small, 30–40×7–8 µm; norstictic and stictic acids; lirellae erumpent from pseudostromatic areas; USA ... Graphis hinnulea Seavey & J. Seavey (Seavey & Seavey Reference Seavey and Seavey2011)

• Group 19, couplet 1: Ascospores small to medium-sized, 45–55×5–10 µm; no substances; lirellae erumpent with lateral thalline margin; Mexico ... Graphis rosae-emiliae Barcenas-Peña & Lücking (this paper)

• Group 19, couplet 2: Stictic acid; ascospores 80–100×15–20 µm; lirellae prominent, with complete thalline margin; Brazil ... Graphis cerradensis Marcelli et al. (Lumbsch et al. Reference Lumbsch, Ahti, Altermann, Amo De Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011)

• Group 20, couplet 1: Ascospores terminally muriform; norstictic acid; ascospores 80–100×15–20 µm; Thailand ... Graphis norvestitoides Sutjaritturakan (Lücking et al. Reference Lücking, Sutjaritturakan and Kalb2012)

• Group 20, couplet 2: Thallus ecorticate; labia pruinose; Sri Lanka ... Graphis knucklensis Weerakoon et al. (Weerakoon et al. Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbsch2012)