Introduction
In the recently published world-wide key to the genus Graphis (Lücking et al. Reference Lücking, Archer and Aptroot2009), two names were published without a valid description and indication of their status: Graphis norvestitoides Sutjaritturakan and G. rongklaensis Sutjaritturakan. In addition, the combination Graphis arbusculiformis was not validly published. These three names are validated herein. In addition, the new species G. pseudoaquilonia Lücking is described, correcting a taxonomic error in the concept of G. norvestitoides.
Graphis arbusculiformis (Vain.) Lücking comb. et stat. nov.
MycoBank No: MB563806
Graphis subdisserpens f. arbusculaeformis Vain., Bot. Tidsskr. 29: 364 (1909); type: Thailand, Schmidt s. n. (TUR-Vainio 27563!—holotypus).
Graphis norvestitoides Sutjaritturakan sp. nov.
MycoBank No.: MB563807
Differing from Graphis vestitoides in the inspersed hymenium and the presence of norstictic acid.
Graphis norvestitoides Sutjaritturakan in Lücking et al., Lichenologist 41: 430 (2009) [nom. inval.]; type: Thailand, Nakhon Ratchasima Province, Khao Yai National Park, forest behind former heliport, 700 m, on unidentified tree trunks in secondary forest, 1999, Sutjaritturakan 12763 (RAMK—holotypus).
(Fig. 1)

Fig. 1. Habit and anatomy of new species. A, Graphis norvestitoides, habit; B, G. pseudoaquilonia, habit; C–E, G. norvestitoides, section through lirella (C), ascus (D) and ascospores (E). Scales: A & B=2 mm; C=100 µm; D=30 µm; E=20 µm.
Thallus corticolous, 3–7 cm diam., 70–120 µm thick, continuous; surface smooth to uneven, pale green-grey to white-grey; prothallus absent except when bordered by other lichens. Section with prosoplectenchymatous upper cortex 20–30 µm thick, irregular algal layer 20–40 µm thick and indistinct medulla 10–40 µm thick and penetrating into the periderm; algal layer with large clusters of calcium oxalate crystals.
Lirellae straight to curved, sparsely branched, prominent, with apically thin complete thalline margin, 1–10 mm long, 0·5–0·7 mm wide, 0·3–0·5 mm high; disc concealed; labia thick, becoming finely striate, appearing dark grey to greyish black with white lines in between striae; thalline margin laterally thick, with algae and clusters of calcium oxalate crystals, white-grey, apically thin, lacking algae and crystals (cortex only), dark grey. Excipulum apically crenulate, completely carbonized, 80–150 µm thick, black; laterally covered by corticate algiferous thallus including clusters of crystals in basal parts; hypothecium prosoplectenchymatous, 15–25 µm high, colourless. Hymenium 130–180 µm high, colourless, strongly inspersed, inspersion dissolving in K; epithecium granulose, 5–15 µm high, olive-brown; paraphyses unbranched; asci fusiform, 120–160×20–30 µm. Ascospores 4–8 per ascus, oblong, terminally muriform with 15–21 transverse septa and 1–2 longitudinal septa in the terminal segments, 70–100×12–15 µm, 5–7 times as long as wide, colourless, septa I+ violet-blue.
Secondary chemistry. Norstictic acid (surface UV−; medulla C−, P+ orange; section K+ yellow efflux forming red, needle-shaped crystals).
Notes. This new species is characterized by lirellae of the acharii-morph (Lücking Reference Lücking2009), being prominent, covered with a thick thalline layer laterally and a thin layer apically so that the upper part of the labia appears dark grey, and having striate labia. In addition, the species has a strongly inspersed hymenium of type B (Lücking Reference Lücking2009), characteristic of the G. cinerea group, terminally muriform ascospores, and norstictic acid. The name was included as a nomen nudum in the world-wide key to Graphis (Lücking et al. Reference Lücking, Archer and Aptroot2009: 430) but erroneously keyed out in Group 18 as having a non-inspersed hymenium and larger ascospores. The reason for this error is that the material studied at that time included two different taxa, which was overlooked. The material from Thailand indicated as original material and here designated as type of G. norvestitoides has an inspersed hymenium and smaller ascospores (70–100 µm long), whereas the material mentioned from the Neotropics, which would key out precisely in Group 18, couplet 3(2), has a clear hymenium and larger ascospores (80–170 µm long). This material is described below as a new species, G. pseudoaquilonia. Instead, G. norvestitoides as described here would key out in Group 20, couplet 1, together with G. subflexibilis Lücking & Chaves, which also has an inspersed hymenium and terminally muriform ascospores, but lacks secondary substances and the ascospores are larger (80–140×10–20 µm).
Graphis pseudoaquilonia Lücking sp. nov.
MycoBank No.: MB563808
Differing from Graphis vestitoides in the presence of norstictic acid and from G. norvestitoides in the clear hymenium.
Typus: Costa Rica, Limón, Gandoca-Manzanillo Wildlife Refuge, Manzanillo Section (La Amistad Caribe Conservation Area), Manzanillo, 82°53′ W, 9°38′ N, sea level, lowland coastal moist forest zone, disturbed coastal vegetation with Cocos nucifera, Coccoloba uvifera, on bark (lower stem), March 2004, Lücking 17098a (F—holotypus, USJ—isotypus). Heredia, La Selva Protection Zone, La Selva Biological Station (Cordillera Volcánica Central Conservation Area), 84°00′ W, 10°25′ N, 50–100 m, March 2004, Lücking 17076a (F, INB—paratypi).
(Fig. 1)
Thallus corticolous, 3–7 cm diam., 70–150 µm thick, continuous; surface smooth to uneven, pale green-grey to white-grey; prothallus absent except when bordered by other lichens. Section with prosoplectenchymatous upper cortex 20–30 µm thick, irregular algal layer 20–40 µm thick and indistinct medulla 10–40 µm thick and penetrating into the periderm; algal layer with large clusters of calcium oxalate crystals.
Lirellae straight to curved, sparsely branched, prominent, with apically thin complete thalline margin, 1–10 mm long, 0·5–0·7 mm wide, 0·3–0·5 mm high; disc concealed; labia thick, becoming finely striate, appearing dark grey to greyish black with white lines in between striae; thalline margin laterally thick, with algal cells and clusters of calcium oxalate crystals, white-grey, apically thin, lacking algal cells and crystals (cortex only), dark grey. Excipulum apically crenulate, completely carbonized, 120–170 µm thick, black; laterally covered by corticate algiferous thallus including clusters of crystals in basal parts; hypothecium prosoplectenchymatous, 15–25 µm high, colourless. Hymenium 150–250 µm high, colourless, clear; epithecium granulose, 5–15 µm high, olive-brown; paraphyses unbranched; asci fusiform, 140–230×35–50 µm. Ascospores 2–6 per ascus, oblong, terminally muriform with 15–21 transverse septa and 1–2 longitudinal septa in the terminal segments, 80–170×20–30 µm, 5–6 times as long as wide, colourless, septa I+ violet-blue.
Secondary chemistry. Norstictic acid (surface UV−; medulla C−, P+ orange; section K+ yellow efflux forming red, needle-shaped crystals).
Notes. This new species corresponds to the concept of the key couplet 3(2) in Group 18 of the world key to Graphis (Lücking et al. Reference Lücking, Archer and Aptroot2009), in having lirellae with striate labia and completely carbonized excipulum, a clear hymenium, terminally muriform ascospores, and norstictic acid. As in other species of this group, specimens with first-generation lirellae have invariably entire labia but the striations eventually form in older specimens. The new species is most similar to G. vestitoides Fink, which differs in the lack of secondary substances. Graphis norvestitoides, originally keyed out there, has an inspersed hymenium and smaller ascospores. Both taxa were misidentified with Graphis aquilonia (A. W. Archer) Staiger (Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008, Reference Lücking, Archer and Aptroot2009), but that species has labia that remain entire and are covered by a thick, white thallus layer up to the top. All these species share the terminally muriform ascospores but can be distinguished as follows:
1 No substances (section K−) ... G. vestitoides
Norstictic acid (section K+ yellow efflux forming red, needle-shaped crystals) ... 2
2(1) Hymenium strongly inspersed ... G. norvestitoides
Hymenium clear ... 3
3(2) Labia remaining entire, covered by thick, white-grey thallus layer up to the top; ascospores 70–110 µm long ... G. aquilonia
Labia becoming striate, covered by apically thin thallus layer and therefore top of labia appearing dark grey; ascospores 80–170 µm long ... G. pseudoaquilonia
Graphis rongklaensis Sutjaritturakan nom. nov.
MycoBank No.: MB563809
Graphis rongklaensis Sutjaritturakan in Lücking et al., Lichenologist 41: 398 (2009) [nom. inval.]; Graphina vestita M. Nakan., Kashiw. & K. H. Moon, Bull. Natn. Sci. Mus., Tokyo, Ser. B, 27(2, 3): 53 (2001); type: Thailand, Moon & Nakanishi 2295 (TNS—holotypus; RAMK—isotypus!).
Notes. This new name is introduced because the necessary combination of the epithet vestita in Graphis is blocked by Graphis vestita Fr. [Syst. Orb. Veg. (Lundae) 1: 288 (1825)]. A further, later homonym is Graphis vestita Fée [Bull. Soc. Bot. Fr. 21: 26 (1874)], a species of unknown taxonomic status as the type material has not yet been studied.
The material was studied in the frame of the NSF-funded projects Ticolichen (DEB 0206125 to The Field Museum; PI Robert Lücking), Phylogeny and Taxonomy of Ostropalean Fungi, with Emphasis on the Lichen-forming Thelotremataceae (DEB 0516116 to The Field Museum; PI H. T. Lumbsch; Co-PI R. Lücking), Neotropical Epiphytic Microlichens – An Innovative Inventory of a Highly Diverse yet Little Known Group of Symbiotic Organisms (DEB 715660 to The Field Museum; PI R. Lücking), and ATM – Assembling a Taxonomic Monograph: The Lichen Family Graphidaceae (DEB-1025861 to The Field Museum; PI T. Lumbsch, CoPI R. Lücking).