Key Characters

The taxonomy of the genus Graphis sensu Staiger (Reference Staiger2002) was revised in a separate paper (Lücking Reference Lücking2009, this volume). The key is based on the results of that study and uses the following suite of morphological, anatomical, and chemical characters to delimit species.

Thallus colour. Most species have a white-grey thallus due to the presence of large clusters of calcium oxalate crystals in and above the photobiont layer. A few taxa have an olive-green colour, similar to Phaeographis and related genera, because the crystal clusters are located below the photobiont layer (Lücking Reference Lücking2009, this volume). In species with high concentrations of norstictic or stictic acid, the thallus might appear pale yellowish. Discoloration in herbarium material is common and due to improper drying which causes decomposition of chlorophyll (yellow to yellow-brown colour of originally white-grey or olive-green thalli) or secondary substances (red-brown colour in specimens containing norstictic or salazinic acid). In such cases, a hand-cut section of the thallus helps to identify the thallus type and natural colour (Fig. 1A–B).

Fig. 1. Morphological and anatomical features in Graphis. A, G. acharii with photobiont layer below calcium oxalate crystal clusters; B, G. caribica with photobiont later above calcium oxalate crystal clusters; C–D, G. isidiata and G. patwardhanii with isidia; E, G. crebra with exposed, pruinose discs; F, G. caesiella with pruinose labia. Scale = 1 mm.

Thallus cortex. Species with a corticate thallus have a compact, slightly glossy surface, whereas ecorticate taxa show a matt, rough or farinose thallus under the dissecting microscope. If present, a cortex is identified as a thin but distinct, hyaline layer of compacted, parallel hyphae in hand-cut sections under the compound microscope (Fig. 1A–B).

Vegetative propagules. Only four species have isidia and one (G. sorediosa) has soralia. Isidia are usually numerous in G. isidiata (Fig. 1C), G. isidiza, and G. patwardhanii (Fig. 1D), but scattered and easily overlooked in G. stellata, so careful inspection of the thallus surface is required.

Lirella emergence. Lirella emergence can be categorized into four states (Fig. 2): 1) immersed (upper part of hymenium more or less level with thallus surface, labia flush or slightly emergent; Fig. 2A); 2) erumpent (upper part of hymenium above, but lower part below thallus level; lirellae with gently sloping margins; Fig. 2B); 3) prominent (hymenium fully above thallus level; lirellae with steeply sloping margins; Fig. 2C & D); 4) sessile (lirellae basally constricted; Fig. 2E–F). Variation of lirella emergence on a single thallus depends mostly on age, with the lirellae in most cases starting out as fissures in the thallus. Lirella emergence is best observed in transverse sections of mature lirellae towards the thallus centre under the dissecting microscope or, with some experience, from surface view, using a light source at an angle of 45 degrees (light sources at higher angles will make the lirellae appear flatter than they actually are). For mature lirellae, we accepted a variation range over two states, i.e. immersed to erumpent, erumpent to prominent, and prominent to sessile. For a given species, the degree of lirella emergence given in the key is the state observed in most specimens (>75%); or the actual range is given.

Fig. 2. Morphological and anatomical features in Graphis. A, G. symplecta with immersed lirellae; B, G. lineola with erumpent lirellae; C, G. vestitoides with prominent lirellae; D, G. tumidula with prominent lirellae; E, G. plagiocarpa with sessile lirellae; F, G. ruiziana with sessile lirellae. Scale = 1 mm.

Thalline margin of lirellae. Because of the carbonized excipulum, the labia in Graphis species are black. However, in many species they are covered by a thalline margin, leaving only the upper part of the labia exposed or covering them completely (Fig. 3). A thalline margin is absent when the black labia are erumpent to sessile and the black colour reaches down to the thallus level (Fig. 3A); by default, immersed lirellae must have at least a lateral thalline margin. A basal thalline margin is developed when it covers less than half of the overall height of the lirellae (Fig. 3B), whereas a lateral thalline margin usually is at level with the height of the lirellae and only leaves the upper, horizontal part of the labia exposed and black (Fig. 3C & D). Many species have an apically thin complete thalline margin: the lateral part of the thalline margin is thick and contains a photobiont layer, whereas the upper part consists of a thin, hyaline cortex only; in this case the upper part of the labium appears dark grey(Fig. 3E). In species with an apically thick complete thalline margin, the margin is uniformly composed of cortex and photobiont layer up to the top and the black excipulum surface is more or less invisible (Fig. 3F). The nature of the thalline margin varies with age: even in species with eventually prominent lirellae lacking a thalline margin, young lirellae (or the tips of the lirellae) first appear as immersed fissures or narrow black lines with a thalline margin. Thus, the nature of the thalline margin should be observed in mature lirellae towards the centre of the thallus.

Fig. 3. Morphological and anatomical features in Graphis. A, G. geraensis lacking a thalline margin; B, G. consimilis with a basal thalline margin; C, G. myrtacea with a lateral thalline margin; D, G. flavens with a thick lateral thalline margin; E, G. argentata with an apically thin complete thalline margin; F, G. illinata with an apically thick complete thalline margin. Scale = 1 mm.

Relative length and branching of lirellae. We have identified six different lirellae shape patterns, i.e. relative length and branching (Fig. 4): 1) round and unbranched (more or less as long as wide; Fig. 4A), 2) very short and unbranched (up to 2 mm long and 3–10 times as long as wide in the longest lirellae; Fig. 4B), 3) short and sparsely branched (up to 3 mm long and 10–15 times as long as broad in the longest lirellae; Fig. 4C), 4) elongate and irregularly branched (5–10 mm long and 15–30 times as long as broad in the longest lirellae; Fig. 4D), 5) very long and radiately branched (usually over 10 mm long and consisting of one or a few lirellae coveringmost of the thallus; Fig. 4E), and 6) short and stellately branched (usually up to 3 mm long and consisting of several to many clusters of stellately branched lirellae; Fig. 4F). Within a given species, variation usually covers one to two different stages, for example, the lirellae can be short to elongate and sparsely to irregularly branched; however, species with round or very short and unbranched lirellae, as well as those with radiately or stellately branched lirellae, vary very little in lirella shape, even in cases where a large number of specimens has been studied (e.g., G. mexicana, G. ruiziana, G. dendrogramma). The degree of branching in a given species also varies depending on the group (Lücking Reference Lücking2009, this volume): in species with striate labia lacking a thalline margin (striatula group), variation in relative length and branching is much greater than in species with entire labia and lateral thalline margins (scripta and subserpentina groups = ‘Eugraphis’), and this is reflected in our species concept.

Fig. 4. Morphological and anatomical features in Graphis. A, G. mexicana with round, unbranched lirellae; B, G. cleistomma with very short, unbranched lirellae; C, G. librata with short, sparsely branched lirellae; D, G. furcata with elongate, irregularly branched lirellae; E, G. arbusculaeformis with very long, radiately branched lirellae; F, G. astrolirellata with short, stellately branched lirellae. Scale = 1 mm.

Disc exposure. Species of Graphis often have the disc more or less exposed when fully hydrated, but in dry conditions the disc is concealed. Yet, a few species have an exposed disc even in dry condition (Fig. 1E); these are almost exclusively found in G. scripta and relatives, which underlines the taxonomic importance of this feature (Lücking Reference Lücking2009, this volume).

Labia and disc pruinosity. Many species exhibit pruinose labia or discs when exposed (Fig. 1E–F). Variation of this character is little understood, but most of the material available indicates that this feature is species specific. The pruina is composed of a layer of crystals and dead hyphal material, differing from the thick pruina characteristic of Carbacanthographis and Dyplolabia, composed of a layer of hyphae inspersed with crystals or pigment (Staiger Reference Staiger2002) and not found in Graphis s. str.

Labium striation. Labium striation (Fig. 5A–B) has been used as an important taxonomic feature in the past (Wirth & Hale Reference Wirth and Hale1963, Reference Wirth and Hale1978) and was also taken up by Staiger (Reference Staiger2002). However, since labium striation seems to be connected to the formation of new hymenia (Staiger Reference Staiger2002), it appears that entire versus striate labia represent developmental stages of the same species rather than a taxonomically important character. On the other hand, there are many taxa in which a striate stage is unknown, i.e. they either never form new hymenia or have other strategies to produce new ascospores. Also, there are many striate taxa in which a non-striate stage is unknown and in which even the younger lirellae at the thallus margin are already striate. Thus, while labium striation alone cannot be used to distinguish species, this character clearly is of taxonomic value. Labium striation is best observed in surface view under the dissecting microscope, in combination with a hand-cut section under the compound microscope; in a few species, thin lines of pruina can resemble striation but the excipulum will appear entire in sections.

Fig. 5. Morphological and anatomical features in Graphis. A, G. rustica with entire labia; B, G. angustata with striate labia; C, G. glaucescens with an apically carbonized excipulum; D. G. pavoniana with a laterally carbonized excipulum; E, G. rhizocola with a completely carbonized excipulum; F. G. gomezii with a completely carbonized excipulum (basal part of excipulum merged with substratum). Scale = 1 mm.

Excipulum carbonization. Graphis is characterized by a partially to completely carbonized excipulum (Fig. 5C–F). Following Redinger (Reference Redinger1935) and Wirth & Hale (Reference Wirth and Hale1963, Reference Wirth and Hale1978), Staiger (Reference Staiger2002) distinguished several groups characterized by their degree of excipulum carbonization. It is not yet well-understood whether the thick carbonized basal excipulum in certain species of Graphis is homologous with the excipulum in other genera or, at least in part, represents a carbonized hypothecium, as in Leiorreuma, Sarcographa, and Thecaria (Staiger Reference Staiger2002). The best way to assess excipulum carbonization is by the observation of hand-cut sections under the dissecting microscope at a high magnification against a white background. Particularly in species of the striatula group, the laterally carbonized excipulum may converge basally to appear almost complete; in these cases, one should inspect several mature lirellae and opt for completely carbonized only if the excipulum is clearly contiguous basally in the majority of the sections.

Hymenium inspersion. Hymenium inspersion is another character that was rarely used in the taxonomy of Graphis, but turned out to be relevant at the species level (Staiger Reference Staiger2002; Lücking et al. Reference Lücking and Rivas Plata2008; Lücking Reference Lücking2009, this volume). The importance of this character is underlined by the fact that two different types of inspersion occur within the genus and that these types are largely restricted to three species groups. Several species related to G. scripta and G. insulana have hymenia inspersed with scattered, small oil droplets lining the paraphyses that more or less persist in KOH but do not obstruct the view of asci and ascospores (type A; Fig. 6A). Other species, related to G. acharii, have a strong and dense hymenial inspersion of larger, irregular droplets completely obstructing the view of asci and ascospores but easily dissolving in KOH (type B; Fig. 6B).

Fig. 6. Morphological and anatomical features in Graphis. A, G. insulana with type A inspersion (asci and ascospores easily visible); B. G. argentata with type B inspersion (asci and ascospores barely visible); C, G. rhizocola with ascospores with lens-shaped lumina; D. G. insulana with ascospores with nearly rectangular lumina; E, G. vestitoides with biocellate ascospore (terminally muriform) and gelatinous caps; F, G. elegans with thick-walled ascospores. Scale = 10 μm.

Ascospores. Species of Graphis typically have hyaline, I+ violet-blue, distoseptate ascospores with lens-shaped lumina (Fig. 6C). The iodine reaction is concentrated in the endospore and usually strong, giving a violet-blue to violet-brown colour (Staiger Reference Staiger2002), except at the tips where the lumina tend to be smaller with less endospore. In a few species (e.g., G. chrysocarpa, G. mucronata, G. pittieri), the ascospores become grey-brown. Some variation is found in the degree of the endospore thickenings, especially in taxa with muriform ascospores (Fig. 6D). ‘Biocellate’ ascospores, such as in G. vestitoides, are transversely septate in the middle but muriform at both ends (Fig. 6E). Correlation with other characters, as well as examination of ascospore ontogeny in species with muriform ascospores (which do not feature a terminally muriform stage), show that terminally muriform ascospores do not represent an ontogenetic stage but are species-specific. Many taxa feature gelatinous sheaths or caps (Fig. 6E), but this character has been little explored and depends on the age of the specimen and how it was preserved. Thick-walled ascospores are found in G. elegans and a few other species (Fig. 6F).

For proper interpretation of ascospore characters, care should be taken to exclude taxonomically uninformative infraspecific and ontogenetic variation. Character states change during ascospore development, for which endospore development is a good indicator: in young ascospores, the endospore is thin and the iodine reaction weak. Mature ascospores are often (slightly) constricted at the septa, while young ascopores are not. In old, post-mature ascospores, the endospore disintegrates and the iodine reaction disappears, and often the ascospores shrivel. The number of ascospores per ascus should be carefully assessed observing several asci from different lirellae. Ascopores are often discharged prematurely or their number is difficult to assess within the ascus. If there are more than four ascospores per ascus, the actual number is most probably eight.

Ascospore size and its taxonomic relevance is a particularly difficult character. Based on the study of indivdual thalli, as well as size variation in well-delimited species, we have generally accepted a size range of about 100%, i.e. the upper limit being about twice as large as the lower limit (e.g., 25–50 μm, 50–100 μm, 100–200 μm). Species with otherwise identical characters but different ascospores, so-called sporomorphs (Wirth & Hale Reference Wirth and Hale1978) occur frequently in Graphis. Analysis of hundreds of species and thousands of specimens revealed certain preferred ascospore size classes, i.e. size ranges that occur more frequently than others: very small (15–25 μm), small (25–45 μm; most ascospores 30–40 μm), medium (45–80 μm; most ascospores 50–70 μm), large (80–150 μm; most ascospores 100–130 μm), and very large (150–300 μm; most ascospores 180–250 μm). Ascospore size should be measured in mature ascospores only, i.e. with the endospore fully developed, and to assess size range, one should measure at least ten ascospores in a given specimen, from different asci and preferably from different lirellae. In species with a variable number of ascospores per ascus, ascospore size might vary more than usual and one should refer to the larger ascospores unless asci with few, large ascospores are an exception in a specimen. Ascospore size should be measured in water; KOH will result in an increase of size ranges by more or less 10–20%.

Secondary chemistry. Most species of Graphis lack secondary substances (Staiger Reference Staiger2002; Archer Reference Archer2006; Lücking et al., Reference Lücking and Rivas Plata2008; Lücking Reference Lücking2009, this volume). Species with a simple chemistry with norstictic or stictic acids and accessory substances are particularly common in relatives of G. scripta and G. insulana (‘Eugraphis’). Salazinic and protocetraric acids are comparatively rare. A few species, especially from the Indian subcontinent, have a combination of up to three of these acids. The concentration of such species in India, also seen in the related genus Hemithecium (Adawadkar & Makhija Reference Adawadkar and Makhija2005; Makhija et al. Reference Makhija, Dube, Adawadkar and Chitale2005a, Reference Makhija and Adawadkarb), does not appear to be an artifact, as we have seen authentic TLC plates of some of these species, but might reflect a peculiarity of the evolution and palaeogeography of Graphis and relatives. Lichexanthone is found in less than a handful of taxa. Yellow to orange anthraquinones are restricted to a few species and usually occur as pruina on the surface of the labia. Isohypocrelline, a dark red, K+ green perlene quinone widespread within the family, has so far been found only in two species of Graphis.

Secondary substances in Graphis are identified either by TLC or spot tests. Most users are familiar with TLC and spot test techniques to routinely identify lichen substances (e.g., Walker & James Reference Walker and James1980; Lumbsch Reference Lumbsch, Kranner, Beckett and Varma2002), but in crustose lichens the reliable application of spot tests is more difficult and there are contradictions as to colour reactions in the literature. Unfortunately, certain rare substances such as hirtifructic and hypostictic acid cannot be identified by spot tests, and TLC is needed in doubtful cases. TLC is also necessary in material or groups (e.g., G. scripta and G. insulana groups) that may contain more than one major substance, especially if collected in the Eastern Palaeotropics. Microchemical reactions should be performed under the microscope on hand-cut sections of thallus and lirellae using a 10–20% KOH solution (to test for the presence of norstictic, salazinic, or stictic acid) and under the dissecting microscope using Steiner's PD solution (to test for the presence of protocetraric acid). A persistently pale to bright yellow efflux without formation of crystals indicates stictic acid, whereas a bright yellow efflux eventually producing red, needle-shaped crystals (immediately or up to 60 seconds later) indicates norstictic (large crystals) or salazinic acid (small crystals). In some cases, the bark substratum reacts with a yellow efflux to KOH, but the colour is more orange-yellow and only appears on the lower (bark) side of the thallus; with some experience, the characteristic stictic-yellow can be readily recognized.

Lirellae morphs. In order to facilitate recognition of lirella morphology, a number of different lirella morphs have been defined that combine lirella emergence, thalline margin, relative length, and branching (Lücking Reference Lücking2009; this volume). In addition to anatomical and chemical characters, one should use the key to morphs below and compare a given specimen to these morphs (Fig. 7–12) before entering the main key.

1. 1 Lirellae lacking thalline margin or with basal thalline margin only ... 2

   Lirellae with lateral to complete thalline margin ... 11

2. 2 (1) Labia entire ... 3

   Labia striate ... 7

3. 3 (2) Lirellae very short and unbranched ... 4

   Lirellae short to elongate and sparsely to irregularly or stellately to radiately branched ... 5

4. 4 (3) Lirellae erumpent ... subregularis-morph (Fig. 7A)

   Lirellae prominent to sessile ... nuda-morph (Fig. 7C)

5. 5 (3) Lirellae prominent to sessile, radiately branched ... slendrae-morph

   Lirellae erumpent to prominent, sparsely to irregularly or stellately branched ... 6

6. 6 (5) Lirellae stellately branched ... geraensis-morph (Fig. 7B)

   Lirellae sparsely to irregularly branched ... hossei-morph (Fig. 7D)

   

   Fig. 7. Lirella morphs in Graphis. A, subregularis-morph (G. subregularis); B, geraensis-morph (G. geraensis); C, nuda-morph (G. ruiziana); D, hossei-morph (G. analoga); E, granulocarpa-morph (G. granulocarpa); F, striatula-morph (G. striatula). Scale = 1 mm.

   

   Fig. 8. Lirella morphs in Graphis. A, lineola-morph (G. lineola); B, deserpens-morph (G. deserpens); C, centrifuga-morph (G. arbusculaeformis); D, coarctata-morph (G. cervinonigra); E, handelii-morph (G. handelii); F, scripta-morph (G. scripta). Scale = 1 mm.

   

   Fig. 9. Lirella morphs in Graphis. A, caesiella-morph (G. caesiella); B, dendrogramma-morph (G. dendrogramma); C, hyphosa-morph (G. hyphosa); D, glaucescens-morph (G. glaucescens); E, tenella-morph (G. myrtacea); F, asterizans-morph (G. asterizans). Scale = 1 mm.

   

   Fig. 10. Lirella morphs in Graphis. A, dussii-morph (G. plagiocarpa); B, subserpentina-morph (G. japonica); C, marginata-morph (G. rustica); D. farinulenta-morph (G. farinulenta); E, lumbricina-morph (G. lumbricina); F, albotecta-morph (G. albotecta). Scale = 1 mm.

   

   Fig. 11. Lirella morphs in Graphis. A, globosa-morph (G. mexicana); B, cleistomma-morph (G. cleistomma); C, illinata-morph (G. illinata); D, consanguinea-morph (G. consanguinea); E, nudaeformis-morph (G. nudaeformis); F, granulosa-morph (G. granulosa). Scale = 1 mm.

   

   Fig. 12. Lirella morphs in Graphis. A, negrosina-morph (G. negrosina); B, rhizocola-morph (G. rhizocola); C, multisulcata-morph (G. multisulcata); D, subradiata-morph (G. subradiata); E, symplecta-morph (G. symplecta); F, acharii-morph (G. acharii). Scale = 1 mm.

7. 7 (2) Lirellae with yellow or orange pruina ... chrysocarpa-morph

   Lirellae lacking (pigmented) pruina ... 8

8. 8 (7) Lirellae very short and unbranched, prominent to sessile ... ... granulocarpa-morph (Fig. 7E)

   Lirellae short to very long and sparsely to radiately branched, erumpent to prominent ... 9

9. 9 (8) Lirellae sparsely to irregularly branched, erumpent to prominent ... ... striatula-morph (Fig. 7F)

   Lirellae radiately or stellately branched, erumpent ... 10

10. 10 (9) Lirellae short, stellately branched ... hunanensis-morph

    Lirellae elongate to very long, radiately branched ... sorediosa-morph

11. 11 (1) Lirellae with lateral thalline margin (upper part of labia black) ... 12

    Lirellae with complete thalline margin (upper part of labia dark grey or invisible) ... 33

12. 12 (11) Labia entire ... 13

    Labia striate ... 28

13. 13 (12) Lirellae immersed to erumpent ... 14

    Lirellae prominent to sessile ... 25

14. 14 (13) Disc exposed ... 15

    Disc concealed ... 16

15. 15 (14) Disc pruinose ... scripta-morph (Fig. 8F)

    Disc non-pruinose ... handelii-morph (Fig. 8E)

16. 16 (14) Labia pruinose ... 17

    Labia non-pruinose ... 20

17. 17 (16) Lirellae radiately or stellately branched ... 18

    Lirellae sparsely to irregularly branched ... 19

18. 18 (17) Lirellae very long and radiately branched ... dendrogramma-morph (Fig. 9B)

    Lirellae short and stellately branched ... hyphosa-morph (Fig. 9C)

19. 19 (17) Thallus ecorticate ... glaucescens-morph (Fig. 9D)

    Thallus corticate ... caesiella-morph (Fig. 9A)

20. 20 (16) Thalline margin thick, bulging ... 21

    Thalline margin thin, not bulging ... 22

21. 21 (20) Lirellae very long, radiately branched, disc slightly exposed ... longiramea-morph

    Lirellae short to elongate and sparsely to irregularly branched, disc concealed ... ... subserpentina-morph (Fig. 10B)

22. 22 (20) Lirellae radiately or stellately branched ... 23

    Lirellae sparsely to irregularly branched ... 24

23. 23 (22) Lirellae very long and radiately branched ... centrifuga-morph (Fig. 8C)

    Lirellae short and stellately branched ... coarctata-morph (Fig. 8D)

24. 24 (22) Lirellae short and sparsely branched ... lineola-morph (Fig. 8A)

    Lirellae elongate and irregularly branched ... deserpens-morph (Fig. 8B)

25. 25 (13) Lirellae stellately branched, lateral margin thin ... stellata-morph

    Lirellae unbranched to irregularly branched, lateral margin usually thick ... 26

26. 26 (26) Labia pruinose ... farinulenta-morph (Fig. 10D)

    Labia non-pruinose ... 27

27. 27 (26) Lirellae very short and unbranched, usually sessile ... dussii-morph (Fig. 10A)

    Lirellae short to elongate and sparsely to irregularly branched, usually prominent ... marginata-morph (Fig. 10C)

28. 28 (12) Lirellae prominent ... 29

    Lirellae immersed to erumpent ... 30

29. 29 (28) Lirellae stellately branched ... asterizans-morph (Fig. 9F)

    Lirellae sparsely to irregularly branched ... celata-morph

30. 30 (28) Labia pruinose ... 31

    Labia non-pruinose ... 32

31. 31 (30) Thallus ecorticate ... glaucescens-morph

    Thallus corticate ... chloroalba-morph

32. 32 (30) Lirellae very long and radiately branched ... dichotoma-morph

    Lirellae short to elongate and sparsely to irregularly branched ... ... tenella-morph (Fig. 9E)

33. 33 (11) Labia entire ... 34

    Labia striate ... 42

34. 34 (33) Lirellae round to very short, unbranched ... 35

    Lirellae short to very long, sparsely to radiately branched ... 36

35. 35 (34) Lirellae round ... globosa-morph (Fig. 11A)

    Lirellae very short ... cleistomma-morph (Fig. 11B)

36. 36 (34) Lirellae radiately or stellately branched ... 37

    Lirellae sparsely to irregularly branched ... 38

37. 37 (36) Lirellae very long and radiately branched ... anguilliradians-morph

    Lirellae short and stellately branched ... evirescens-morph

38. 38 (36) Lirellae sessile, thalline margin partly flaking off ... ... nudaeformis-morph (Fig. 11E)

    Lirellae erumpent to prominent, thalline margin persistent ... 39

39. 39 (38) Thalline margin apically thin (upper part of labia dark grey) ... 40

    Thalline margin apically thick (labia more or less invisible) ... 41

40. 40 (39) Lirellae erumpent ... negrosina-morph (Fig. 12A)

    Lirellae prominent ... rhizocola-morph (Fig. 12B)

41. 41 (39) Lirellae erumpent ... subserpentina-morph (Fig. 10B)

    Lirellae prominent ... illinata-morph (Fig. 11C)

42. 42 (33) Lirellae radiately branched ... subradiata-morph (Fig. 12D)

    Lirellae unbranched to irregularly branched ... 43

43. 43 (42) Lirellae very short, unbranched ... 44

    Lirellae short to elongate, sparsely to irregularly branched ... 45

44. 44 (43) Thalline margin apically thin (upper part of labia dark grey) ... ... multisulcata-morph (Fig. 12C)

    Thalline margin apically thick (labia more or less invisible) ... ... granulosa-morph (Fig. 11F)

45. 45 (43) Labia pruinose ... albotecta-morph (Fig. 10F)

    Labia non-pruinose ... 46

46. 46 (45) Thalline margin apically thick (labia more or less invisible), lirellae immersed to erumpent ... consanguinea-morph (Fig. 11D)

    Thalline margin apically thin (upper part of labia dark grey), lirellae immersed to prominent ... 47

47. 47 (46) Lirellae immersed to erumpent ... symplecta-morph (Fig. 12E)

    Lirellae prominent ... 48

48. 48 (47) Thalline margin partly flaking off ... lumbricina-morph (Fig. 10E)

    Thalline margin persistent ... acharii-morph (Fig. 12F)

Genera likely to be confused with Graphis

The following key helps to separate genera with carbonized excipulum and well-developed labia from Graphis s.str. We have also mentioned species of genera with hyaline ascospores likely to be confused with Graphis under the corresponding key couplets of the Graphis keys.

1. 1 Ascospores hyaline ... 2

   Ascospores grey-brown ... 6

2. 2 (1) Labia white pruinose, C+ red (lecanoric acid) ... Dyplolabia

   Labia non-pruinose or pruinose but C– ... 3

3. 3 (2) Ascospores I−; labia usually thickly white or pale yellow pruinose; warty peri-physoids above the hymenium usually present ... Carbacanthographis

   Ascospores I+ violet-blue; ascomata non-pruinose or thinly white or brown pruinose; warty periphysoids absent ... 4

4. 4 (3) Lirellae brown pruinose ... Glyphis

   Lirellae non-pruinose or (thinly) white pruinose ... 5

5. 5 (4) Thallus ecorticate, with diverse secondary chemistry (stictic, norstictic, salazinic, and/or protocetraric acid and/or lichexanthone); lirellae with distinctly white-pruinose labia and partially exposed, white-pruinose disc; paraphyses apically anastomosing; ascospores muriform ... Diorygma

   Thallus corticate (if ecorticate then lacking secondary substances or rarely norstictic acid and ascospores transversely septate); lirellae non-pruinose or thinly white-pruinose but not with both labia and exposed disc thickly white-pruinose; paraphyses unbranched or rarely apically branched but not anastomosing; ascospores variable ... Graphis

6. 6 (1) Disc exposed ... Platygramme

   Disc concealed ... 7

7. 7 (6) Thallus white-grey; hymenium clear or rarely inspersed; ascospores I+ violet-blue ... Graphis

   Thallus olive-green; hymenium inspersed; ascospores I+ vine-red ... 8

8. 8 (7) Lirellae sessile, lacking thalline margin; excipulum completely carbonized ... ... Thecographa

   Lirellae erumpent to prominent, with lateral to complete thalline margin; excipulum apically to laterally carbonized ... Platygramme

Group 1: Lirellae with yellow, orange, or red pigment

1. 1 Lirellae with yellow or orange to cinnabar-red pruina covering the labia or in between excipular striae (and then lirellae with complete thalline margin); pigment K+ salmon-yellow slowly turning purple or K+ instantly pink-purple (tetra-hydroxy-anthraquinone-1,3,6,8 and related substances) ... 2

   Lirellae black, lacking pigmented pruina or thalline margin; epithecium and/or hymenium or basal parts of lirellae orange or dark red; pigment K+ purple (tetra-hydroxy-anthraquinone-1,3,6,8 and related substances) or K+ green (isohypocrelline) ... 9

2. 2 (1) Pigment yellow, K+ salmon-yellow turning slowly purple when drying; lirellae prominent to sessile, elongate and irregularly branched, with yellow pruina covering the labia to varying extent; ascospores large ... 3

   Pigment orange-red, K+ instantly pink-purple; lirellae erumpent to sessile, short to elongate and sparsely to irregularly branched, with orange to cinnabar-red pruina covering the labia or disc or in between excipular striae; ascospores small to large ... 5

3. 3 (2) Labia striate; hymenium inspersed; ascospores regularly muriform, 80–100 × 10–17 μm; Neotropics (Central America) [holotype: Costa Rica, Sipman 47811a (B!; isotype:INB!)] ... Graphis flavoaltamirensis Sipman & Lücking

   Labia entire; hymenium clear; ascospores transversely septate or terminally muriform only, size variable ... 4

4. 4 (3) Ascospores transversely septate, 70–100 μm long; Neotropics (Central America) [holotype: Costa Rica, Lücking 15216 (F!)] ... Graphis firferi Lücking

   Ascospores terminally muriform, 100–170 μm long; Neotropics (Central America, Galapagos) [holotype: Costa Rica, Moncada 3009 (INB!)] ... ... Graphis flavominiata Moncada & Lücking

5. 5 (2) Ascospores small (20–35 × 5–8 μm); lirellae erumpent; salazinic acid; Eastern Palaeotropics [lectotype (here selected): Philippines, Host 15028 (TUR-Vainio 27880!)] ... ... Graphis ferruginea Vain.

   Ascospores medium-sized to large (70–190 × 10–25 μm); lirellae prominent to sessile; no additional substances; Neotropics and African Palaeotropics ... 6

6. 6 (5) Excipulum apically carbonized; ascospores transversely septate; Neotropics [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): South America, s.col. (L!)] ... Graphis lutea (Chevall.) Aptroot

   Excipulum completely carbonized; ascospores terminally or regularly muriform, size variable ... 7

7. 7 (6) Lirellae with complete thalline margin; pigment embedded in between excipular striae, not as pruina on labia, only visible when thallus cover is scraped off; ascospores terminally muriform; Neotropics [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Mosén 3145 (S 6493!)] ... Graphis miniata Redinger

   Lirellae with basal to lateral thalline margin; pigment as thick pruina on labia; ascospores terminally or regularly muriform ... 8

8. 8 (7) Ascospores terminally muriform only, 70–120 × 10–15 μm; Neotropics and African Palaeotropics [holotype: Sierra Leone, Deighton M-4307 (FH!)] ... ... Graphis subchrysocarpa Lücking [≡ Phaeographis ochracea C. W. Dodge, non Graphis ochracea Hepp]

   Ascospores regularly muriform, 120–190 × 15–25 μm; Neotropics [holotype: Brazil, s.col. (FI, not seen); neotype: Ecuador, Frisch 96/Eq325 (M, not seen; Staiger Reference Staiger2002)] ... ... Graphis chrysocarpa (Raddi) Spreng.

   Notes. Staiger (Reference Staiger2002) lists three further synonyms under Graphis chrysocarpa: G. ignea Kremp. [holotype: Panama, Wagner s.n. (M, not seen)], G. kermesina Fée [holotype: Brazil, Glaziou 1895 (G, not seen)], G. rubricosa Fée [syntypes: Brazil, Glaziou 2159, 2173 (G, not seen)]. Without the type material, which is possibly lost, it is impossible at this point to ascertain whether these belong to G. chrysocarpa s.str. or any of the other, similar species recognized here (G. subchrysocarpa, G. miniata, G. lutea).

9. 9 (1) Pigment orange, K+ purple (tetra-hydroxy-anthraquinone-1,3,6,8 and related substances); lirellae prominent, lacking thalline margin, with entire labia (hossei-morph) ... 10

   Pigment dark red, K+ green (isohypocrelline); lirellae variable but not as above. 11

10. 10(9) Excipulum completely carbonized; hymenium inspersed; norstictic acid; Neotropics (USA) [holotype: USA. (Florida), Britton & Britton 681 (NY!)] ... ... Graphis chromothecia R. C. Harris

    Excipulum laterally carbonized; hymenium clear; no substances; Neotropics (USA.) [holotype: USA (Florida), Harris 23223 (NY!)] ... Graphis inversa R. C. Harris

11. 11 (9) Hymenium and epithecium inspersed with red pigment; labia entire; excipulum laterally carbonized; lirellae erumpent, with apically thin thalline margins, short and sparsely branched (negrosina-morph); ascospores 20–45 × 6–8 μm; Palaeotropics [holotype: Philippines, Meyen s.n. (B, not seen; Hale Index Cards)] ... ... Graphis persicina G. Mey. & Flot.

    Hymenium clear; red pigment restricted to basal parts of lirellae; labia striate; excipulum laterally (to almost completely) carbonized; lirellae prominent, lacking thalline margin, elongate and irregularly branched (striatula-morph); ascospores 40–50 × 8–13 μm; Neotropics (Central America) [holotype: Costa Rica, Lücking 15297d (F!)] ... Graphis hypocrellinea Lücking & Chaves

Group 2: Labia entire, excipulum apically carbonized, hymenium clear, ascospores transversely septate

1. 1 Ascospores medium-sized to large (60–100 μm long), 15–23-septate; stictic acid ... 2

   Ascospores small to medium-sized (15–60 μm long), 3–15-septate; chemistry variable ... 3

2. 2 (1) Lirellae erumpent with lateral thalline margin, short and sparsely branched (lineola-morph); ascospores 75–100 μm long; Eastern Palaeotropics [holotype: India, Patwardhan & Nagarkar 82.262 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis valparaiensis Adaw. & Makhija

   Lirellae immersed to erumpent, with apically thick complete thalline margin, elongate and irregularly branched (subserpentina-morph); ascospores 65–75 μm long; Eastern Palaeotropics [lectotype (here selected, based on annotation label by Nakanish 1973): Taiwan, Asahina 354 (W!)] ... Graphis epiphloea Zahlbr.

   Notes. On the lectotype of Graphis epiphloea, Nakanishi in 1973 annotated that the species is synonymous with G. awaensis Vain.; both are indeed similar but the type of G. awaensis matches that of G. longiramea in having a laterally carbonized excipulum and a lateral rather than complete thalline margin, which is why we keep G. epiphloea separate at this point.

3. 3 (1) Norstictic, salazinic, or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 4

   No substances (K−) ... 8

4. 4 (3) Stictic acid (K+ yellow) ... 5

   Norstictic or salazinic acid (K+ yellow forming red crystals) ... 6

5. 5 (4) Ascospores medium-sized (50–60 × 10–12 μm), 9–11-septate; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph); Eastern Palaeotropics (China) [holotype: China, Handel-Mazzetti 11222 (W!)] ... ... Graphis elegantula Zahlbr.

   Ascospores small (20–40 × 5–8 μm), 5–9-septate; lirellae immersed, with lateral thalline margin, very long and radiately branched (centrifuga-morph); Eastern Palaeotropics [holotype: Philippines, Merrill 6724 (TUR-Vainio 27810!)] ... ... Graphis arecae Vain.

6. 6 (4) Salazinic acid; lirellae prominent, with apically thick complete thalline margin (illinata-morph); excipulum apically to peripherally carbonized; Eastern Palaeotropics [lectotype (Awashti & Singh Reference Awasthi and Singh1975): India, Thwaites 152 (BM, not seen; Awasthi & Singh Reference Awasthi and Singh1975; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis nematoides Leight.

   Notes. We have not been able to study the type material but this taxon might belong in Carbacanthographis rather than Graphis.

   Norstictic acid; lirellae immersed to erumpent, with lateral to complete thalline margin; excipulum apically carbonized ... 7

7. 7 (6) Ascospores very small (15–25 × 4–5 μm); lirellae immersed, with lateral thalline margin, elongate and irregularly branched, with disc exposed and white-pruinose (scripta-morph); Eastern Palaeotropics [holotype: India, Sethy & Nagarkar 81.578 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis alboglaucescens Adaw. & Makhija

   Ascospores small (30–40 × 4–6 μm); lirellae erumpent, with apically thin complete thalline margin, short to elongate and sparsely to irregularly branched, with disc concealed and labia non-pruinose (negrosina-morph); Eastern Palaeotropics [holotype: India, Patwardhan & Badhe 70.70 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis eburnea Adaw. & Makhija

8. 8 (3) Thallus ecorticate, farinose; ascospores small (25–45 × 6–10 μm) ... 9

   Thallus corticate, smooth to uneven; ascospores variable ... 10

9. 9 (8) Lirellae immersed to erumpent, with lateral thalline margin and pruinose labia, elongate and irregularly branched (glaucescens-morph) (see also Group 13: 16a); pantropical (also saxicolous) ... Graphis glaucescens Fée

   1. A. Labia entire [holotype: Paraguay, Balansa s.n. (G!)] ... Graphis glaucocaesia Müll. Arg.

   2. B. Labia entire [holotype: Philippines, Merrill 7051 (TUR-Vainio 27882!)] ... Graphis bulacana Vain.

   3. C. Labia entire [holotype: Angola, Pechuel-Loesche s.n. (B, not seen; Dodge Reference Dodge1964)] Graphis angolensis C. W. Dodge [≡ Graphis caesia Müll. Arg., nom. illeg., non Spreng.]

   Lirellae immersed to erumpent, with apically thick complete thalline margin and non-pruinose labia, elongate and irregularly branched (subserpentina-morph); Eastern Palaeotropics [holotype: New Caledonia, Thiébaut s.n. (H-Nylander 7681!)] ... ... Graphis malacodes Nyl.

10. 10(8) Disc slightly exposed, white-pruinose; ascospores medium-sized (40–60 × 10–12 μm); lirellae immersed, with thick lateral thalline margin (scripta-morph); Eastern Paleotropics [holotype: Australia, Bailey 778 (G!)] ... ... Graphis epimelaena Müll. Arg.

    Notes. This species was considered a synonym of Graphis glaucescens by Archer (Reference Archer2006) but has larger ascospores, a corticate thallus, and a slightly exposed, pruinose disc.

    Disc concealed; ascospores variable; lirellae variable ... 11

11. 11(10) Labia white-pruinose; lirellae immersed to erumpent, with apically thin thalline margin, elongate and irregularly branched (caesiella-morph) ... 12

    Labia non-pruinose; lirellae variable ... 13

12. 12(11) Ascospores medium-sized (40–55 μm long), 9–15-septate; pantropical ... ... Graphis sayeri Müll. Arg.

    1. A. Ascospores 40–55 × 6–10 μm; lirellae immersed to erumpent [holotype: Australia, Sayer s.n. (G!; isotype: MEL!)] ... Graphis sayeri Müll. Arg.

    2. B. Ascospores 45–55 × 5–7 μm; lirellae erumpent [lectotype (here selected): Paraguay, Malme 1526 Bex (S-6505!)] ... Graphis glaucovirens Redinger

13. Ascospores small (20–40 μm long); 5–9-septate; pantropical ... ... Graphis xanthospora Müll. Arg.

    1. A. Ascospores 30–40 × 8–10 μm; lirellae immersed [holotype: Australia, Sayer s.n. (G!)] ... ... Graphis xanthospora Müll. Arg.

    2. B. Ascospores 25–35 × 6–8 μm; lirellae immersed to erumpent [holotype: Philippines, Merrill 8343 (TUR-Vainio 27885!; isotype: FH!)] ... Graphis mindanaoensis Vain.

    3. C. Ascospores 20–30 × 5–7 μm; lirellae immersed to erumpent [original material: Cuba, Ekman 53 (TUR-Vainio 27372!)] ... Graphis bayatensis Vain. [nom. inval. (not published)]

14. 13(11) Thallus papillose; ascospores very small (15–20 μm long), 3–5-septate; lirellae erumpent, with lateral thalline margin, short and stellately branched (coarctata-morph); Eastern Palaeotropics [lectotype (Awasthi & Singh Reference Awasthi and Singh1975): India, Watt s.n. (BM, not seen; Awasthi & Singh Reference Awasthi and Singh1975)] ... Graphis coarctata Stirt.

    Thallus smooth to uneven; ascospores small (20–45 μm long), 5–9-septate; lirellae erumpent, with lateral thalline margin, unbranched to radiately branched ... 14

15. 14 (13) Lirellae very long and radiately branched (centrifuga-morph); excipulum apically carbonized; thallus smooth to uneven; Neotropics and Eastern Palaeotropics ... ... Graphis intermediella Stirt.

    1. A. Ascospores 20–30 × 6–8 μm [holotype: India, s.col. (BM, not seen; Awasthi Reference Awasthi1991)] ... ... Graphis intermediella Stirt.

    2. B. Ascospores 20–25 × 6–7 μm [holotype: Japan, Yasuda 250 (TUR-Vainio 27814!)] ... ... Graphis batanensis var. rikuzensis Vain.

       Notes. Nakanishi (Reference Nakanishi1966) gives the holotype of Graphis batanensis var. rikuzensis Vain. as number 27489 in hb. Vainio, but the correct number is 27814; number 27489 is Graphis vicarians Vain., which is a synonym of Thecaria quassiicola Fée. Also, Nakanishi (Reference Nakanishi1966) describes the lirellae of G. batanensis var. rikuzensis as having an exposed disc but the disc on the type material is concealed and only slightly exposed in damaged lirellae.

16. Lirellae short and sparsely branched (lineola-morph); excipulum apically to peripherally carbonized; thallus conspicuously rugose; Eastern Palaeotropics [holotype: China, Handel-Mazzetti 2451 (W!; isotype: S-2171!)] ... Graphis oligospora Zahlbr.

Group 3: Labia entire, excipulum apically carbonized, hymenium clear, ascospores muriform

1. 1 Ascospores medium-sized to large (45–160 × 15–35 μm); stictic acid (K+ yellow) or no substances (K−) ... 2

   Ascospores small (15–45 × 7–18 μm); no substances (K−) ... 8

   Notes. If ascospores small and I− and norstictic acid present, cf. Carbacanthographis saxiseda (Zahlbr.) Bungartz.

2. 2(1) Stictic acid (K+ yellow) ... 3

   No substances (K−) ... 5

3. 3(2) Lirellae prominent, with thick lateral thalline margin, very short and unbranched (dussii-morph); Eastern Palaeotropics [holotype: Sri Lanka, s.col. (H-Nylander, not seen; Patwardhan & Nagarkar Reference Patwardhan and Nagarkar1979)] ... Graphis bilabiata Nyl.

   Lirellae immersed to erumpent, with thick lateral to thick complete thalline margin, elongate and irregularly branched (subserpentina-morph) ... 4

4. 4(3) Ascospores 1(–2) per ascus, large (70–100 × 15–30 μm) (see also Group 5: 13a); Neotropics and Eastern Palaeotropics (also saxicolous) ... ... Graphis streblocarpa (Bél.) Nyl.

   1. A. Excipulum apically carbonized; ascospores 70–95 × 20–30 μm [holotype: India, Bélanger s.n. (G!)] ... Opegrapha streblocarpa Bél.

   2. B. Excipulum apically carbonized; ascospores 80–100 × 20–25 μm [holotype: Thailand, Hosseus s.n. (TUR-Vainio 27215!)] ... Graphis streblocarpa var. pauperior Vain.

   3. C. Excipulum apically to peripherally carbonized; ascospores 75–100 × 20–25 μm [holotype: Philippines, Ramos & Edaño s.n. (TUR-Vainio 27227!)] ... Graphis hiascens var. clausior Vain.

   Ascospores (2–)4–8 per ascus, medium-sized (45–80 × 15–25 μm) (see also Group 5: 13b); Eastern Palaeotropics ... ... Graphis japonica (Müll. Arg.) A. W. Archer & Lücking*

   1. A. Excipulum apically to peripherally carbonized; ascospores 2–4 per ascus, 50–80 × 15–21 μm; thallus smooth to uneven [holotype: Indonesia, s.col. (W, not seen; Hale Index Cards)] ... ... Graphina tjibodensis Zahlbr.

   2. B. Excipulum apically to laterally carbonized; ascospores 4–8 per ascus, 45–50 × 15–20 μm; thallus verruculose [lectotype (here selected): China, Chung 600d (W!; isolectotype: FH!)] ... ... Graphina verruculina Zahlbr.

   3. C. Excipulum apically to peripherally carbonized; ascospores 2–4 per ascus, 45–75 × 15–21 μm; thallus smooth to uneven [holotype: Taiwan, Faurie 156 (W!)] ... Graphina filiformis Zahlbr.

   4. D. Excipulum apically to peripherally carbonized; ascospores 2–4 per ascus, 60–80 × 15–25 μm; thallus smooth to uneven [holotype: Taiwan, Faurie 83 (W, not seen; Hale Index Cards] ... ... Graphina petrophila Zahlbr.

5. 5(2) Ascospores 6–8 per ascus; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph) ... 6

   Ascospores 1(–4) per ascus; lirellae variable ... 7

6. 6(5) Ascospores large (70–90 × 20–25 μm); Australia [holotype: Australia, Streimann 56807 (CANB!)] ... Graphis coenensis A. W. Archer [≡ Graphis celata (A. W. Archer) A. W. Archer, nom. illeg., non Stirt.]

   Ascospores (small to) medium-sized (25–75 × 13–20 μm); Europe [holotype: Great Britain, s.col. (BM, not seen; Staiger Reference Staiger2002)] ... Graphis britannica Staiger [≡ Opegrapha pulverulenta Sm. & Sowerby, non Graphis pulverulenta (Pers.) Ach.]

   Notes. We have not yet studied the various synonyms of Graphis britannica, such as Graphis sophistica Nyl. ex Cromb. [non Nyl.] and Graphis inustula Nyl. [non Stirt.].

7. 7(5) Disc exposed, white-pruinose; lirellae prominent, lacking thalline margin, elongate and irregularly branched (scripta-morph); Eastern Palaeotropics [holotype: China, Handel-Mazzetti 6982 (W!)] ... Graphis alpestris (Zahlbr.) Staiger

   Disc concealed, labia non-pruinose; lirellae erumpent, with apically thick complete thalline margin, elongate and irregularly branched (subserpentina-morph); Neotropics ... Graphis subcontorta (Müll. Arg.) Lücking & Chaves

   1. A. Excipulum apically to peripherally carbonized; lirellae with apically thin complete thalline margin; ascospores 80–100 × 25–35 μm [holotype: Paraguay, Balansa 183 (G!)] ... ... Graphina subcontorta Müll. Arg.

   2. B. Excipulum apically carbonized; lirellae with lateral thalline margin; ascospores 60–110 × 15–30 μm [original material: not designated] ... Graphis pseudoserpentina Chaves & Lücking [nom. inval. (not published)]

8. 8(1) Thallus partly ecorticate; lirellae immersed, with apically thick complete thalline margin, elongate and irregularly branched (subserpentina-morph); USA, (Hawaii) [(holotype: Hawaii, Rock 135 (W!)] ... Graphis sulphurella (Zahlbr.) Lücking*

   Thallus corticate; lirellae immersed to erumpent, with lateral thalline margin, short and sparsely or stellately branched (lineola- or coarctata-morph) ... 9

9. 9(8) Ascospores broad (25–45 × 13–20 μm; if ascospores partly longer than 45 μm, cf. G. britannica); African Palaeotropics [lectotype (here selected): South Africa, Van der Byl 912 (W!)] ... Graphis pergracilis (Zahlbr.) Lücking & A. W. Archer*

   Notes. Graphis subnitida Nyl. sensu Archer (Reference Archer2007b) would key out here as potentially earlier name for G. pergracilis; however, the type material of G. subnitida (H-NYL 7449!) represents a Fissurina, which on account of the slightly erumpent, fissurine lirellae and rather large (25– 35 × 12–16 μm), muriform ascospores, in combination with lack of secondary substances, is a synonym of F. rubiginosa (Fée) Staiger). The material reported as G. subnitida by Archer (Reference Archer2007b) is to be named G. pergracilis. The type material of Graphina pergracilis contains two species, one with concealed disc and apically carbonized excipulum (here selected as lectotype) and one with exposed, pruinose disc and laterally carbonized excipulum, representing G. pyrrhocheiloides.

   Ascospores narrow (15–40 × 8–15 μm); Neotropics (also saxicolous) ... 10

10. 10(9) Lirellae stellately branched (coarctata-morph) [holotype: Grand Cayman, Imshaug 24510 (MSC-0024093!)] ... Graphis astrolirellata Lücking

    Lirellae short and sparsely branched (lineola-morph) [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Vainio 322 (TUR-Vainio 27241!)] ... Graphis dimidiata Vain.

Group 4: Labia entire, excipulum laterally carbonized, hymenium clear, ascospores transversely septate

1. 1 Ascospores medium-sized to large (45–135 μm long; mature ascospores usually exceeding 50 μm) ... 2

   Ascospores small (15–45 μm long) ... 16

2. 2(1) Norstictic, salazinic acid, and/or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 3

   No substances (K−) ... 11

3. 3(2) Norstictic and/or salazinic acid, sometimes additionally stictic acid (K+ yellow forming red crystals) ... 4

   Stictic acid, sometimes additionally hypostictic acid (K+ yellow) ... 8

4. 4(3) Lirellae prominent, with thin complete thalline margin that often flakes off to expose black labia, short and sparsely branched (lumbricina-morph); ascospores 15–20 μm broad; norstictic acid (see also Group 17: 6a); Neotropics and Hawaii ... Graphis lumbricina Vain.

   Lirellae erumpent to prominent, with (thick) lateral, persistent thalline margin; ascospores 6–16 μm broad; chemistry variable [if lirellae lacking thalline margin and ascospores medium-sized (35–60 μm long, cf. Graphis elegans] ... 5

5. 5(4) Lirellae prominent to sessile, with thick lateral thalline margin, very short and unbranched (dussii-morph); norstictic acid; Neotropics [holotype: Lesser Antilles (Guadelupe), Duss 515 (TUR-Vainio 27368!)] ... Graphis dussii Vain.

   Lirellae immersed to erumpent, with (thick) lateral thalline margin, short to elongate and sparsely to radiately branched ... 6

6. 6(5) Ascospores medium-sized (45–60 μm long), 7–13-septate; lirellae short and sparsely branched (lineola-morph); norstictic acid; Neotropics and Eastern Palaeotropics ... Graphis erythrocardia Müll. Arg.

   1. A. Lirellae erumpent, with lateral thalline margin; hymenium copper-coloured (artifact?); ascospores 50–60 × 7–10 μm, 9–13-septate [holotype: Tanzania, Holst 3081 (G!)] ... ... Graphis erythrocardia Müll. Arg.

   2. B. Lirellae erumpent, with lateral thalline margin; hymenium colourless; ascospores 40–50 × 8–10 μm, 9–13-septate [lectotype (Archer Reference Archer1999): Australia, Knight 341 (G!)] ... ... Graphis vinosa Müll. Arg.

   Ascospores medium-sized to large (45–135 μm long, mature ascospores exceeding 60 μm), (9–)11–27-septate; lirellae elongate to very long and irregularly to radiately branched (subserpentina-morph); salazinic acid together with norstictic or stictic acids ... 7

7. 7(6) Lirellae very long and radiately branched; thallus verrucose; salazinic, stictic, hypostictic, and hypoconstictic acids; Eastern Palaeotropics [holotype: India, Kulkarni & Prabhu 74.3180 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis salacinilongiramea Adaw. & Makhija

   Lirellae elongate and irregularly branched; thallus smooth to uneven; salazinic and norstictic acids; Eastern Palaeotropics [holotype: India, Awasthi 7973 (LWU, photograph seen; Awasthi & Singh Reference Awasthi and Singh1977; Awasthi Reference Awasthi1991; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis longispora D. D. Awasthi & S. R. Singh

   Notes. Graphis garoana Nagarkar & Patw. would key out here, but according to the description and illustrations, this species might belong in Carbacanthographis.

8. 8(3) Ascospores medium-sized to large (60–100 μm long, mature ascospores usually exceeding 70 μm) ... 9

   Ascospores medium-sized (44–65 μm long) ... 10

9. 9(8) Ascospores 4–8 μm broad; lirellae with thin lateral thalline margin, elongate and irregularly branched, disc concealed (deserpens-morph); Eastern Palaeotropics [holotype: India, Sethy & Patwardhan 86.145 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... Graphis sitapurensis Makhija & Adaw.

   Ascospores 8–16 μm broad; lirellae with thick lateral thalline margin, elongate to very long and irregularly to radiately branched, disc often slightly exposed (longiramea-morph); Eastern Palaeotropics ... Graphis longiramea Müll. Arg.

   1. A. Lirellae radiately branched; ascospores 60–100 × 8–12 μm, 7–23-septate [lectotype (Awasthi & Singh Reference Awasthi and Singh1975): India, Watt 6981 (BM, not seen; paratype: G!)] ... Graphis longiramea Müll. Arg.

   2. B. Lirellae irregularly branched; ascospores 50–70 × 10–12 μm, 11–17-septate [holotype: Japan, Yasuda 251 (TUR-Vainio 27813!)] ... Graphis awaensis Vain.

   3. C. Lirellae radiately branched; ascospores 50–80 × 9–14 μm, 11–17-septate [holotype: China, Handel-Mazzetti 9486 (W!; isotype: S 2172!)] ... Graphis multibrachiata Zahlbr.

   4. D. Lirellae irregularly branched; ascospores 60–85 × 12–16 μm, 13–17-septate [type: China, Handel-Mazzetti 12345 (W!; isotype: US!)] ... Graphis zonatula Zahlbr.

      Notes. The lectotype of Graphis longiramea was given as Watt 6981 by Awasthi & Singh (Reference Awasthi and Singh1975); the specimen in G, labelled Watt 78, according to the illustrations and description by Awasthi & Singh (Reference Awasthi and Singh1975), represents exactly the same morphotype and was also annotated as lectotype by Patwardhan & Kulkarni in 1975 and by Awasthi & Mathur in 1989. Possibly the two stem from the same original collection and there is some confusion with the collection numbers; it is unlikely but not impossible that the same species was recollected by Watt with such disparate numbers within a year.

10. 10(8) Lirellae very narrow, with gently sloping lateral thalline margin, flexuose; excipulum laterally carbonized; Eastern Palaeotropics [holotype: New Caledonia, Deplanche s.n. (H- Nylander 7751!)] ... Graphis leptogramma Nyl.

    Lirellae broader, with thick and abruptly sloping lateral thalline margin, irregularly curved; excipulum laterally to almost completely carbonized; Palaeotropics ... ... Graphis crassilabra Müll. Arg.

    1. A. Ascospores 9–15-septate, 45–65 × 8–12 μm [holotype: Australia, Hartmann 78 (G!; isotype: MEL!)] ... Graphis crassilabra Müll. Arg.

    2. B. Ascospores 9–15-septate, 55–65 × 7–10 μm [holotype: Indonesia (Java), s.col (G!)] ... ... Graphis stenospora Müll. Arg.

11. 11(2) Thallus ecorticate; labia distinctly white-pruinose ... 12

    Thallus corticate; labia non-pruinose or thinly white-pruinose or disc white-pruinose ... 13

12. 12(11) Ascospores 45–60 μm long; lirellae erumpent, with lateral thalline margin (glaucescens-morph); Neotropics [lectotype (Staiger Reference Staiger2002): South America, s.col. (G!)] ... ... Graphis pavoniana Fée

    Ascospores 60–80 μm long; lirellae prominent, with lateral thalline margin (farinulenta-morph); Central America [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Costa Rica, Pittier 5146 (G!)] ... Graphis farinulenta Müll. Arg.

13. 13(11) Lirellae prominent, with basal thalline margin, elongate and irregularly branched (hossei-morph) ... Graphis hossei Vain.

    1. A. Excipulum laterally carbonized; ascospores 2–8 per ascus, 50–110 × 8–14 μm, 11–17-septate [lectotype (based on annotation label by Nakanishi in 1973): Thailand, Hosseus s.n. (TUR- Vainio 27806!)] ... Graphis hossei Vain.

    2. B. Excipulum laterally carbonized; ascospores 8 per ascus, 40–75 × 7–11 μm, 9–15-septate [lectotype (here selected): Thailand, Hosseus s.n. (TUR-Vainio 27803!)] ... Graphis tenuis Vain.

    3. C. Excipulum laterally (to completely) carbonized; ascospores 4 per ascus, 60–110 × 10–16 μm, 11–21-septate [holotype: Philippines, McGregor 8652 (TUR-Vainio 27860!)] ... ... Graphis nematodiza Vain.

    4. D. Excipulum laterally carbonized; ascospores 2–4 per ascus, 70–110 × 10–15 μm, 11–19-septate [holotype: China, Handel-Mazzetti 12348 (W!)] ... Graphis bifera var. cinerea Zahlbr.

    5. E. Excipulum laterally (to almost completely) carbonized; ascospores 8 per ascus, 40–70 × 10–14 μm, 11–17-septate [lectotype (here selected, based on annotation label by Nakanishi 1973): China, Handel-Mazzetti 11429 (W!; isolectotype: S 2186!)] ... Graphis connectens Zahlbr.

    6. F. Excipulum laterally carbonized; ascospores 8 per ascus, 65–75 × 10–14 μm, 15–19-septate [holotype: Indonesia (Java) Groenhart 8744 (BO 7743, not seen; Groenhart Reference Groenhart1954)] ... ... Graphis psidii Groenh.

14. Lirellae erumpent, with (thick) lateral to complete thalline margin ... 14

15. 14(13) Disc (partly) exposed, more or less white-pruinose (see also entry 32a); cosmopolitan ... Graphis scripta (L.) Ach.

    Disc concealed, labia non-pruinose ... 15

16. 15(14) Lirellae stellately branched; ascospores 6–8 μm broad; Neotropics [holotype: Uruguay, Imshaug 42496 (MSC-0009125!)] ... Graphis uruguayensis Lücking

    Lirellae irregularly branched; ascospores 9–15 μm broad; pantropical ... ... Graphis elongata Zenker

    1. A. Excipulum laterally carbonized; lirellae with lateral thalline margin; ascospores 8 per ascus, 70–80 × 11–15 μm, 13–17-septate [lectotype: Brazil, s.col. (not seen; Hale Index Cards; Zenker Reference Zenker, Goebel and Kunze1829)] ... Graphis elongata Zenker

    2. B. Excipulum laterally (to completely) carbonized; lirellae with apically thin thalline margin; ascospores 8 per ascus, 50–75 × 9–12 μm, 11–15-septate [holotype: Australia, Streimann 61366 (CANB!)] ... Graphis stenospora var. deficiens A. W. Archer

       Notes: Graphis elongata Zenker was first considered an invalid name but has been validly published by Zenker (Reference Zenker, Goebel and Kunze1829) and thus has precedence over G. stenospora var. deficiens.

17. 16(1) Lichexanthone (UV+ yellow), together with norstictic acid; ascospores very small (15–20 μm long); lirellae erumpent, with basal thalline margin, elongate and irregularly branched (hossei-morph); Eastern Palaeotropics [holotype: Australia, Streimann 16826 (CANB!; isotypes: H!, US!)] ... Graphis stipitata A. W. Archer

    No lichexanthone (UV−); ascospores small (15–45 μm long, mature ascospores usually exceeding 20 μm); lirellae variable ... 17

18. 17(16) Norstictic, salazinic, stictic, or protocetraric acid, or traces of atranorin (K+ yellow or K+ yellow forming red crystals or P+ red) ... 18

    Notes: If lirellae with thick white pruina containing lecanoric acid (C+ red) and thallus olive green to olive brown, cf. Dyplolabia afzelii (Ach.) A. Massal.; if lirellae with thick white pruina lacking lecanoric acid (C−) but thallus containing psoromic acid (P+ yellow), cf. Carbacanthographis alloafzelii (A. W. Archer) A. W. Archer.

    No substances (K−, P−) ... 35

19. 18(17) Norstictic and/or salazinic acid, sometimes additionally protocetraric or stictic acid (K+ yellow forming red crystals) ... 19

    Stictic and/or protocetraric acid or traces of atranorin (K+ yellow or P+ red) ... 29

20. 19(18) Salazinic acid present, sometimes additionally norstictic, stictic, and/or protocetraric acid ... 20

    Salazinic acid absent, norstictic acid only or rarely additionally stictic acid ... 25

21. 20(19) Salazinic and norstictic acid present, sometimes additionally with stictic or protocetraric acid ... 21

    Salazinic acid only or additionally stictic acid ... 22

22. 21(20) Lirellae stellately branched (coarctata-morph); norstictic and salazinic acids; Eastern Palaeotropics [lectotype (Awasthi & Singh Reference Awasthi and Singh1975): India, Watt s.n. (BM, not seen; Awasthi & Singh Reference Awasthi and Singh1975)] ... Graphis capillacea Stirt.

    Lirellae sparsely to irregularly branched; norstictic and salazinic acids and additionally with either protocetraric or stictic acid ... 23

23. 22(20) Norstictic, salazinic, and protocetraric acids; labia white-pruinose (caesiella-morph); thallus white-grey; ascospores 30–35 μm long; Neotropics (Central America) [holotype: Costa Rica, Lücking 17089b (CR!; isotype: F!)] ... ... Graphis litoralis Lücking, Sipman & Chaves

    Norstictic, salazinic, and stictic acids; labia non-pruinose (deserpens-morph); thallus olive-grey; ascospores 15–30 μm long; Eastern Palaeotropics [holotype: India, Patwardhan & Nagarkar 77.1345 (AMH, photograph seen; Nagarkar & Patwardhan Reference Nagarkar and Patwardhan1982; Awasthi Reference Awasthi1991; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis subasahinae Nagarkar & Patw.

24. 23(21) Salazinic, stictic, and constictic acids; lirellae erumpent, with lateral thalline margin (labia white-pruinose), elongate to very long and irregularly to radiately branched (caesiella- or dendrogramma-morph); Eastern Palaeotropics ... ... Graphis argentia Makhija & Adaw.

    1. A. Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched; labia white pruinose; ascospores 5–7-septate, 20–25 × 4–6 μm [holotype: India, Patwardhan & Nagarkar 85.2226 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis argentia Makhija & Adaw. [as argentius]

    2. B. Lirellae erumpent, with lateral thalline margin, elongate to very long and irregularly to radiately branched; labia white pruinose; ascospores 5–9-septate, 15–35 × 4–6 μm [holotype: India, Sethy & Nagarkar 85.2158 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis insularis Makhija & Adaw.

    Salazinic acid only; lirellae variable ... 24

25. 24(23) Lirellae erumpent, with lateral thalline margin, labia white-pruinose, elongate and irregularly branched (caesiella-morph); Eastern Palaeotropics [lectotype (here selected, based on annotation label by Nakanishi in 1973): Philippines, Baker 547 (TUR-Vainio 27874!)] ... ... Graphis bakeri Vain.

    Notes. If ascospores small and I−, cf. Carbacanthographis saxorum (Egea & Torrente) Bungartz.

    Lirellae prominent, with apically thick complete thalline margin (illinata-morph); Eastern Palaeotropics [lectotype (Awashti & Singh Reference Awasthi and Singh1975): India, Thwaites 152 (BM, not seen; Awasthi & Singh Reference Awasthi and Singh1975; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis nematoides Leight.

26. 25(19) Disc and/or labia white-pruinose ... 26

    Disc concealed, labia non-pruinose ... 27

27. 26(25) Disc exposed, white-pruinose; norstictic acid only; lirellae elongate, irregularly branched (scripta-morph); pantropical [holotype: Thailand, Schmidt s.n. (TUR-Vainio 27556!)] ... Graphis pyrrhocheiloides Zahlbr. [≡ G. pyrrhocheila Vain., nom. illeg., non Mont. & Bosch]

    Disc concealed; labia white-pruinose; norstictic acid only; lirellae elongate and irregularly branched (caesiella-morph), pantropical (also saxicolous) ... ... Graphis caesiella Vain.

    1. A. Ascospores 5–9-septate, 20–40 × 6–8 μm [holotype: Brazil, Vainio s.n. (TUR-Vainio 27730!)] ... Graphis caesiella Vain.

    2. B. Ascospores 7–9-septate, 25–30 × 6–8 μm [holotype: Puerto Rico, Fink 1691 (MICH, not seen; isotype: NY!)] ... Graphis yaucoensis Fink

    3. C. Ascospores 7–11-septate, 25–35 × 6–8 μm [holotype: Australia, Elix 22579 (CANB!)] ... ... Graphis kakaduensis A. W. Archer

28. 27(25) Norstictic and stictic acids; thallus olive-grey; Eastern Palaeotropics [holotype: India, Kushalnagar 74.3281 (AMH, photograph seen; Patwardhan & Kulkarni Reference Patwardhan and Nagarkar1979; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis ajarekarii Patw. & C. R. Kulk.

    Norstictic acid only; thallus white-grey to yellowish ... 28

29. 28(27) Lirellae elongate to very long and irregularly to radiately branched; South-east Asia (India) ... Graphis filiformis Adaw. & Makhija

    1. A. Lirellae very long and radiately branched; ascospores 20–25 × 5–7 μm, 7–9-septate [holotype: India, Mhaskar 71.54 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis filiformis Adaw. & Makhija

    2. B. Lirellae elongate and irregularly branched; ascospores 20–30 × 6–9 μm, 5–9-septate [holotype: Indonesia (Krakatau), Van Leuwen 271 (W!)] ... Graphis tenella var. elongata Zahlbr.

    Lirellae short and sparsely branched; pantropical (also saxicolous) [if excipulum completely carbonized, cf. Graphis assimilis] ... Graphis librata C. Knight

    1. A. Excipulum laterally (to almost completely) carbonized; ascospores 20–30 × 5–8 μm, 5–9 -septate [lectotype (Hayward 1977): New Zealand, Knight 67:23 (WELT; Staiger Reference Staiger2002)] ... ... Graphis librata C. Knight

    2. B. Excipulum laterally carbonized; ascospores 20–25 × 5–8 μm, 5–7-septate [holotype: South Africa, Wilms 11 (G!)] ... Graphis diaphoroides Müll. Arg.

    3. C. Excipulum laterally carbonized; ascospores 25–35 × 5–8 μm [syntypes: USA (Louisiana), Langlois 742, 744 (PH, not seen)] ... Graphis celtidis Müll. Arg.

    4. D. Excipulum laterally carbonized; ascospores 25–35 × 5–8 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): USA (Palmyra Island), Rock 193 (W!; isolectotypes: S 4127!, US!)] ... Graphis palmyrensis Zahlbr.

       Notes. Graphis palmyrensis was considered a separate species lacking lichen substances by Lücking et al. (Reference Lücking and Rivas Plata2008), but re-examination of the type demonstrated norstictic acid, as already annotated by Nakanishi in 1973; the correct name for the taxon lacking norstictic acid is G. pinicola.

30. 29(18) Protocetraric acid or additionally with stictic acid ... 30

    Stictic acid only (with accessory substances) or traces of atranorin ... 31

31. 30(29) Disc exposed, white-pruinose (scripta-morph); protocetraric, stictic, and constictic acids; Eastern Palaeotropics [holotype: India, Patwardhan 85.26 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... Graphis distincta Makhija & Adaw.

    Disc concealed, labia white-pruinose (caesiella-morph); protocetraric acid only; pantropical ... Graphis supracola A. W. Archer

    1. A. Lirellae erumpent, short to elongate and irregularly branched [holotype: Australia, Elix 20222 (CANB!)] ... Graphis supracola A. W. Archer

    2. B. Lirellae immersed (to erumpent), elongate and irregularly to radiately branched [original material: Brazil, Höhnel 158 (W!)] ... Graphis scripta var. candida Zahlbr. [nom. inval. (not published)]

32. 31(29) Lirellae with apically thick complete thalline margin, elongate and irregularly branched (subserpentina-morph); Neotropics [holotype: Puerto Rico, Fink 2167 (MICH, not seen; isotype: FH!) ... Graphis riopiedrensis (Fink) Lücking*

    Lirellae with basal to lateral thalline margin or labia pruinose ... 32

    Notes. If lirellae white-pruinose and labia with warty periphysoids, cf. Carbacanthographis iriomotensis (M. Nakan.) M. Nakan. & Kashiw.

33. 32(31) Ascospores 3-septate, 12–17 μm long; atranorin (trace); lirellae erumpent, with basal thalline margin, short and sparsely branched, non-pruinose (subregularis-morph); African Palaeotropics [holotype: Rwanda, Bock s.n. (GOET, photograph seen; Bock & Hauck Reference Bock and Hauck2005)] ... Graphis tetralocularis C. Bock & Hauck

    Notes. This species is included here with some hesitation; we have not seen the type material, but on account of the small, 3-septate ascospores and the unusual chemistry, its placement in Graphis s.str. is questionable.

    Ascospores 5–11-septate, 20–45 μm long; stictic acid; lirellae variable ... 33

34. 33(32) Labia non-pruinose; lirellae short and sparsely branched (lineola-morph); Palaeotropics ... Graphis immersella Müll. Arg

    1. A. Ascospores 25–40 × 6–8 μm, 7–11-septate [lectotype (Archer Reference Archer1999a): Australia, Shirley 1793 (G!)] ... Graphis immersella Müll. Arg.

    2. B. Ascospores 35–45 × 6–8 μm, 7–9-septate [holotype: China, Handel-Mazzetti 5831 (W!)] ... ... Graphis manhaviensis Zahlbr.

    3. C. Ascospores 30–45 × 6–8 μm, 9–11-septate [holotype: Solomon Islands, Hill 10120 (BM!)] ... ... Graphis leptalocarpa A. W. Archer

    Labia pruinose; lirellae elongate to very long and and irregularly to radiately branched (caesiella-, dendrogramma- or scripta-morph) ... 34

35. 34(33) Disc exposed (scripta-morph); Eastern Palaeotropics [holotype: India, Jagadeesh Ram 828 (CAL, photograph seen; Jagadeesh Ram et al. Reference Jagadeesh Ram, Sinha and Singh2007)] ... ... Graphis sundarbanensis Jagadeesh Ram & G. P. Sinha

    Disc concealed (caesiella- or dendrogramma-morph); pantropical ... ... Graphis dendrogramma Nyl.

    1. A. Lirellae radiately branched; excipulum laterally (to almost completely) carbonized; ascospores 20–30 × 6–8 μm, 7–9-septate [holotype: Malaysia (Admiralty Islands), Moseley 1875 (H-Nylander 7165!)] ... Graphis dendrogramma Nyl.

    2. B. Lirellae irregularly branched; excipulum laterally carbonized; ascospores 20–40 × 5–7 μm, 5–9-septate [holotype: Sri Lanka, Almquist s.n. (H-Nylander 7900!)] ... ... Graphis ceylanica Zahlbr. [≡ Graphis irradians Nyl., nom. illeg., non Fée]

    3. C. Lirellae irregularly branched; excipulum laterally carbonized; ascospores 30–45 × 6–8 μm, 7–11-septate [holotype: Solomon Islands, 5176 (W!)] ... Graphis bougainvillei Zahlbr.

    4. D. Lirellae radiately branched; excipulum laterally carbonized; ascospores 20–30 × 5–7 μm, 7–9-septate [holotype: Philippines, McGregor 10209 (TUR-Vainio 27812!)] ... ... Graphis batanensis Vain.

    5. E. Lirellae irregularly branched; excipulum laterally carbonized; ascospores 20–30 × 5–7 μm, 5–7-septate [holotype: Philippines, Merrill 8529 (TUR-Vainio 27840!)] ... ... Graphis subvirginea var. streblicola Vain.

    6. F. Lirellae irregularly branched; excipulum laterally carbonized; ascospores 25–35 × 5–7 μm, 5–9-septate [holotype: Philippines, McGregor 18421 (TUR-Vainio 27841!)] ... ... Graphis subvirginea var. rhizophorearum Vain.

36. 35(17) Disc at least partly exposed, usually white-pruinose (scripta-morph); ascospores partly longer than 45 μm (see also 35a); cosmopolitan [lectotype: Tab. XVIII and fig. 1 in Dillenius, Hist. Musc.; epitype: Sweden, Malme s.n. (UPS!); Staiger Reference Staiger2002)] ... ... Graphis scripta (L.) Ach.

    Notes. Graphis scripta has an extensive synonymy of about 30 specific and infraspecific epithets (see Index of Fungi) which is not listed here. We have not studied all of these in detail but suspect that this species might represent a collective taxon.

    Disc concealed, labia non-pruinose or rarely thinly white-pruinose; ascospores up to 45 μm long ... 36

37. 36(35) Lirellae prominent to sessile, short to very short ... 37

    Lirellae erumpent, short to elongate (Graphis furcata complex) ... 40

38. 37(36) Lirellae lacking thalline margin or with basal thalline margin only; ascospores very small (15–25 μm long) ... 38

    Lirellae with (thick) lateral thalline margin; ascospores small (30–40 μm long) ... 39

39. 38(37) Lirellae sessile, lacking thalline margin, short and unbranched (nuda-morph); Neotropics and Eastern Palaeotropics [holotype: South America, s.col. (G!)] ... ... Graphis nana (Fée) Spreng.

    Lirellae prominent, with basal thalline margin, elongate and partly branched (hossei-morph); Neotropics (Guatemala) and Eastern Palaeotropics [lectotype (here selected): Philippines, Elmer 14634 (TUR-Vainio 27876!)] ... Graphis urandrae Vain.

40. 39(37) Lirellae prominent, with lateral thalline margin, elongate and irregularly branched (marginata-morph); ascospores 9–11 μm broad; Eastern Palaeotropics [holotype: Japan, Yasuda 182 (TUR-Vainio 27802, not seen; Nakanishi et al. Reference Nakanishi, Kashiwadani and Moon2003b)] ... ... Graphis prunicola Vain.

    Lirellae prominent to sessile, with thick lateral thalline margin, short and unbranched (dussii-morph); ascospores 4–6 μm broad; Neotropics [holotype: South America, s.col. (H-Acharius 658!)] ... Graphis enteroleuca (Ach.) Lücking*

41. 40(36) Thallus partly ecorticate; lirellae thin (0·1–0·2 mm), flexuose, with gently sloping thalline margins; labia often thinly white-pruinose (caesiella-morph); pantropical (also foliicolous) [holotype: South America, s.col. (G, not seen; Staiger Reference Staiger2002)] ... ... Graphis furcata Fée

    1. A. Lirellae with lateral thalline margin; thallus partly ecorticate; ascospores 25–35 × 7–10 μm, 5–9-septate [holotype: South America, s.col. (G, not seen; Staiger Reference Staiger2002)] ... Graphis furcata Fée

    2. B. Lirellae with lateral thalline margin; thallus not well-developed (foliicolous); ascospores 30–40 × 6–8 μm, 7–9-septate [holotype: Africa, Welwitsch 393 (TUR-Vainio 27615!)] ... ... Graphis foliicola Vain.

    3. C. Lirellae with lateral thalline margin; thallus corticate; ascospores 35–40 × 8–10 μm, 9–11-septate [holotype: China, Handel-Mazzetti 3075 (W, not seen; Zahlbruckner Reference Zahlbruckner and Handel-Mazetti1930; Hale Index Cards) ... Graphis setchwanensis Zahlbr.

    Thallus corticate; lirellae thicker (0·15–0·3 mm), straight or curved, with abruptly sloping thalline margin (deserpens- or subserpentina-morph); labia always non- pruinose ... 41

42. 41(40) Lirellae elongate to very long and irregularly to radiately branched, with thick lateral thalline margin (subserpentina-morph); Eastern Palaeotropics ... ... Graphis albissima Müll. Arg.

    1. A. Ascospores 30–40 × 7–8 μm, 9–11-septate [holotype: Australia, Knight 147 (G!)] ... ... Graphis albissima Müll. Arg.

    2. B. Ascospores 35–45 × 5–7 μm, 9–13-septate [holotype: India, Sethy & Patwardhan 87.348 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... Graphis leptocarpoides Makhija & Adaw.

43. Lirellae short to elongate and sparsely to irregularly branched (lineola- or deserpens-morph); pantropical (also foliicolous) ... Graphis pinicola Zahlbr.

    1. A. Lirellae with lateral thalline margin; ascospores 25–35 × 10–12 μm, 7-septate [holotype: China, Handel-Mazzetti 2829 (W!; isotype: US!)] ... Graphis pinicola Zahlbr.

    2. B. Lirellae with lateral thalline margin; ascospores 20–30 × 5–8 μm, 5–7-septate [holotype: China, Handel-Mazzetti 11462 (W, not seen; Zahlbruckner Reference Zahlbruckner and Handel-Mazetti1930; Hale Index Cards)] ... ... Graphis castanopsidis Zahlbr.

    3. C. Lirellae with apically thin complete thalline margin; ascospores 30–45 × 5–8 μm, 7–11-septate [holotype: India, Singh 71.429 (LWU; not seen; Awasthi & Singh Reference Awasthi and Singh1972)] ... ... Graphis foliicola var. major Awasthi & Singh

    4. D. Lirellae with apically thin complete thalline margin; ascospores 25–30 × 7–9 μm, 9-septate [holotype: Sierra Leone, Deighton M-4345 (FH!)] ... Graphis guineensis C. W. Dodge

       Notes. This taxon was named Graphis palmyrensis by Lücking et al. (Reference Lücking and Rivas Plata2008), but re-examination of the type demonstrated norstictic acid, as already annotated by Nakanishi in 1973; the correct name for the taxon lacking norstictic acid is thus G. pinicola.

Group 5: Labia entire, excipulum laterally carbonized, hymenium clear, ascospores muriform

1. 1 Norstictic, salazinic, or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   Notes. If lirellae with thick white pruina containing lecanoric acid (C+ red) and thallus olive-green to olive-brown, cf. Dyplolabia oryzoides (Leight.) Kalb & Staiger.

   No substances or in two species with unknown substances (K−) ... 14

2. 2 (1) Norstictic acid, sometimes additionally with salazinic acid (K+ yellow forming red crystals) ... 3

   Stictic acid (K+ yellow) ... 12

   Notes. If salazinic acid major and ascospores I−, 12–17 × 5–7 μm, cf. Carbacanthographis marcescens (Fée) Staiger & Kalb.

3. 3 (2) Ascospores 1(–2) per ascus, medium-sized to large [(40–)50–150 μm long] ... 4

   Ascospores (2–)4–8 per ascus, small (20–50 μm long) ... 8

4. 4 (3) Ascospores medium-sized [(40–)50–70 μm long]; lirellae immersed to erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph); Eastern Palaeotropics [holotype: Philippines, Copeland 196 (TUR-Vainio 27257!)] ... ... Graphis copelandii Vain.

   Ascospores large (70–150 μm long); lirellae erumpent or prominent, with (thick) lateral thalline margin ... 5

5. 5 (4) Lirellae erumpent, with (thick) lateral thalline margin, elongate and irregularly branched (subserpentina-morph); Palaeotropics ... Graphis subserpentina Nyl.

   1. A. Ascospores 80–130 × 20–35 μm [holotype: Sri Lanka, Hooker 8012 (G!; isotype: US!)] ... ... Graphis subserpentina Nyl.

   2. B. Ascospores 70–150 × 20–25 μm [holotype: Malaysia (Malacca), Maingay s.n. (H-NYL 8018!)] ... Graphis adtenuans Nyl.

   3. C. Ascospores 70–120 × 15–25 μm [holotype: Singapore, s.col. (W, not seen; Hale Index Cards)] ... Phaeographina alutacea Zahlbr.

   Lirellae erumpent to prominent, with thick lateral to complete thalline margin, short and unbranched ... 6

6. 6 (5) Lirellae with apically thick complete thalline margin (illinata-morph); ascospores up to 170 × 45 μm; Eastern Palaeotropics (Thailand) [holotype: Thailand, Moon & Nakanishi 2295 (TNS, photograph seen; isotype: RAMK!) ... ... Graphis rongklaensis Sutjaritturakan [≡ Graphina vestita M. Nakan., Kashiw. & K. H. Moon, non Graphis vestita Fr. nec Graphis vestita Fée]

   Lirellae with lateral thalline margin (dussii-morph); ascospores up to 120 ×35 μm ... 7

7. 7 (6) Norstictic and salazinic acid; Eastern Palaeotropical [holotype: India, Patwardhan & Nagarkar 77.1417 (AMH, photograph seen; Patwardhan & Nagarkar Reference Patwardhan and Nagarkar1979)] ... ... Graphis awasthii (Patw. & Nagarkar) Lücking*

   Norstictic acid only; pantropical [holotype: South America, s.col. (G!)] ... ... Graphis hiascens (Fée) Nyl.

   Notes. This taxon requires further study; except for the type, all collections available have very short lirellae of the dussii-morph, whereas in the type they are longer and less prominent; potentially this represents a collective species.

8. 8 (3) Lirellae prominent, with basal thalline margin (hossei-morph); Palaeotropics ... ... Graphis analoga Nyl.

   1. A. Lirellae short to elongate; excipulum laterally (to almost completely) carbonized; ascospores 6–8 per ascus, 20–30 × 8–12 μm [holotype: Tahiti, Viellard & Planchet G13:8 (H-Nylander 7432!)] ... Graphis analoga Nyl.

   2. B. Lirellae short; excipulum laterally (to almost completely) carbonized; ascospores 8 per ascus, 25–35 × 8–12 μm [holotype: Philippines, Merrill 9056 (TUR-Vainio 27251!)] ... ... Graphis analoga var. nana Vain.

   3. C. Lirellae elongate; excipulum laterally (to almost completely) carbonized; ascospores 8 per ascus, 30–40 × 10–14 μm [holotype: Thailand, Schmidt s.n. (TUR-Vainio 27198!)] ... ... Graphis simplex Vain.

   Lirellae erumpent to prominent, with lateral thalline margin ... 9

9. 9 (8) Lirellae prominent, with apically thick complete lateral thalline margin, very short and unbranched (cleistomma-morph); ascospores (small to) medium-sized (40–50 × 13–16 μm); Eastern Palaeotropics [holotype: Papua New Guinea, Aptroot 38161 (B!)] ... Graphis nadurina Aptroot*

   Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph) ... 10

10. 10 (9) Ascospores small (20–35 × 8–14 μm); pantropical ... ... Graphis renschiana (Müll. Arg.) Stizenb.

    1. A. Lirellae with lateral thalline margin; excipulum laterally carbonized; ascospores 8 per ascus, 25–35 × 10–14 μm [holotype: unknown locality, s.col. (UPS!)] Opegrapha gracilis Fr. [non Fée]

    2. B. Lirellae with lateral thalline margin; excipulum laterally carbonized; ascospores 8 per ascus, 25–35 × 9–12 μm [holotype: Madagascar, Rensch 969 (G!)] ... Graphina renschiana Müll. Arg.

    3. C. Lirellae with thick lateral thalline margin; excipulum laterally carbonized; ascospores 4–8 per ascus, 20–30 × 8–12 μm [holotype: Philippines, Rosenbluth & Tamesis 13753 (TUR-Vainio 27185!)] ... Graphis antillarum var. manilensis Vain.

    4. D. Lirellae with lateral thalline margin; excipulum laterally (to almost completely) carbonized; ascospores 4–8 per ascus, 25–35 × 9–13 μm [holotype: China, Handel-Mazzetti 11459 (W!; isotype: S 3963!)] ... Graphina symplocorum Zahlbr.

       Notes. There is some confusion as to the application of the name Graphis gracilis: Zahlbruckner (Reference Zahlbruckner1923) cited G. gracilis (Fr.) Leight. as the combination; however, Leighton (Reference Leighton1869) did not actually recombine Opegrapha gracilis Fr. into Graphis, as assumed by Zahlbruckner (Reference Zahlbruckner1923) and Lücking et al. (Reference Lücking and Rivas Plata2008), but instead based his G. gracilis (Eschw.) (sic!) on Ustalia gracilis Eschw. and cited a Fée illustration that refers to a taxon now included in Arthoniales. Thus, the earliest available epithet, gracilis (from Opegrapha gracilis Fr.), cannot be used for the taxon delimited here, since the combination Graphis gracilis is blocked by Leighton's original combination, and hence the next available epithet renschiana has to be taken up.

    Ascospores (small to) medium-sized (35–50 × 10–25 μm) ... 11

11. 11(10) Ascospores 17–25 μm broad; Eastern Palaeotropics (Australia) [holotype: Australia, Streimann 56574 (CANB!)] ... Graphis borealis (A. W. Archer) A. W. Archer

    Ascospores 10–16 μm broad; Palaeotropics [holotype: Philippines, Baker 1253 (TUR-Vainio 27260!)] ... Graphis norstictica A. W. Archer & Lücking*

12. 12(2) Ascospores small (20–35 × 8–12 μm); lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph); Eastern Palaeotropics [holotype: Philippines, McGregor 8544 (TUR-Vainio 27234!)] ... Graphis deserpens Vain.

    Ascospores medium-sized to large (45–100 × 15–25 μm); lirellae erumpent, with thick lateral to complete thalline margin (subserpentina-morph) ... 13

    Notes. If lirellae prominent and with distinct white pruina, cf. Carbacanthographis cleitops (Fée) Lücking* and C. triphoroides (M. Wirth & Hale) Lücking*.

13. 13(12) Ascospores single (see also Group 3: 4a); Neotropics and Eastern Palaeotropics ... Graphis streblocarpa (Bél.) Nyl.

    1. A. Excipulum laterally carbonized; ascospores 60–100 × 15–25 μm [lectotype (Archer Reference Archer2006): Sri Lanka, Thwaites CL 75 (BM!)] ... Graphis fissofurcata Leight.

       Notes. According to the description and illustrations in the protologue, Graphina indica D. D. Awasthi & S. R. Singh (Awasthi & Singh Reference Awasthi and Singh1977) would key out here. The described K-reaction suggests stictic acid, but we have not studied the type and TLC is necessary to establish the exact compounds present.

       If lirellae prominent and with distinct white pruina, cf. Carbacanthographis cleitops (Fée) Lücking*.

14. Ascospores 2–8 per ascus (see also Group 3: 4); Eastern Palaeotropics ... ... Graphis japonica (Müll. Arg.) A. W. Archer & Lücking*

    1. A. Lirellae with thick complete thalline margin; ascospores 2–4/ascus, 50–70 × 18–25 μm [holotype: Japan, Miyoshi 23 (G!)] ... Graphina japonica Müll. Arg.

    2. B. Lirellae with thick lateral thalline margin; ascospores 2/ascus, 60–80 × 20–25 μm [lectotype (here selected): Taiwan, Asahina 338 (W!)] ... Graphina japonica var. major Zahlbr.

       Notes. If lirellae prominent and with distinct white pruina, cf. Carbacanthographis triphoroides (M. Wirth & Hale) Lücking*.

15. 14 (1) Ascospores medium-sized to large (50–140 × 15–40 μm) ... 15

    Ascospores small (20–50 × 7–18 μm) ... 22

    Notes. Graphina pauciloculata Coppins & P. James (Coppins & James Reference Coppins and James1978) would key out here; we have not seen the type material but judging from the excellent description and photographs, due to its aggregate-stromatic lirellae and very small ascospores this species does not appear to belong in Graphis s.str.; its generic home is unclear.

16. 15(14) Ascospores large (100–140 × 30–40 μm); thallus verrucose; lirellae prominent, with apically thin complete thalline margin (rhizocola-morph); Eastern Palaeotropics [holotype: Australia, Hafellner 16977 (CANB!)] ... ... Graphis atrocelata (A. W. Archer) A. W. Archer

    Ascospores medium-sized to large (50–100 × 15–30 μm); thallus smooth to uneven; lirellae variable ... 16

17. 16(15) Lirellae very short, almost round to ellipsoid, prominent, with apically thick complete thalline cover (globosa-morph); Neotropics [holotype: Colombia, Lindig 2652 (H- Nylander 7588!)] ... Graphis triphora Nyl.

    Lirellae distinctly elongate, erumpent to prominent, with lateral to apically thick complete thalline margin ... 17

18. 17(16) Ascospores single ... 18

    Ascospores 2–8 per ascus ... 20

19. 18(17) Ascospores 50–60 × 15–20 μm; lirellae short and stellately branched (coarctata- morph); Eastern Paleotropics [holotype: Japan, Nakanishi 1902 (HIRO, not seen; Nakanishi Reference Nakanishi1966)] ... Graphis kousyuensis (Horik. & M. Nakan.) Lücking*

    Notes. This taxon was placed as a synonym of Graphis alpestris by Kurokawa (Reference Kurokawa2003) but according to the description given by Nakanishi (Reference Nakanishi1966, Reference Nakanishi1967) differs in the laterally carbonized excipulum and the stellate lirellae.

    Ascospores 55–100 × 15–30 μm; lirellae elongate and irregularly branched ... 19

20. 19(18) Lirellae prominent, with basal thalline margin (hossei-morph); Eastern Palaeotropics ... Graphis daintriensis (A. W. Archer) A. W. Archer

    1. A. Excipulum laterally (to completely) carbonized; ascospores 55–95 × 15–30 μm [holotype: Australia, Streimann 45823 (CANB!)] ... Graphina daintriensis A. W. Archer

    2. B. Excipulum laterally carbonized; ascospore size unknown [original material: not designated] ... ... Graphina chartarna F. Wilson [nom. inval. (not published)]

21. Lirellae erumpent, with lateral to apically thick complete thalline margin (subserpentina-morph); pantropical [holotype: Tanzania, Holst 696 (G!)] ... ... Graphis subhiascens (Müll. Arg.) Lücking

22. 20(17) Lirellae prominent, with basal thalline margin, elongate and irregularly branched (hossei-morph); Eastern Palaeotropics [holotype: Papua New Guinea, Aptroot 37428 (B!)] ... Graphis myolensis Aptroot*

    Lirellae erumpent, with lateral to apically thick complete thalline margin (subserpentina-morph) ... 21

23. 21(20) Ascospores 1–2 per ascus, 70–90 × 20–30 μm; lirellae with lateral thalline margin; Neotropics [holotype: USA (Florida), Harris 23900 (NY!)] ... ... Graphis xylophaga (R. C. Harris) Lendemer [non (R. C. Harris) Lücking]

    Notes. This species was simultaneously recombined in Graphis by Lendemer & Knudsen (Reference Lendemer and Knudsen2008) and Lücking et al. (Reference Lücking and Rivas Plata2008), and the first publication predates the second one.

    Ascospores 2–4/ascus, 50–70 × 15–22 μm; lirellae with almost complete thalline margin; Neotropics and Eastern Palaeotropics [holotype: Brazil, Glaziou 3303 (G!)] ... Graphis lapidicola Fée

    Notes. If lirellae with exposed, brown-pruinose disc and apically anastomosing paraphyses, cf. Glyphis atrofusca (Müll. Arg.) Lücking*.

24. 22(14) Lirellae erumpent; disc exposed (scripta-morph) ... 23

    Lirellae erumpent to prominent; disc concealed ... 24

25. 23(22) Ascospores 20–30 μm long; pruina on disc thin; Eastern Palaeotropics [holotype: Australia, Gwyther s.n. (BRI!)] ... Graphis tenuirima (Shirley) A. W. Archer

    Ascospores 15–20 μm long; pruina on disc thick; Eastern North America [holotype: USA., Calkins s.n. (H-Nylander!)] ... Graphis sophisticascens Nyl.

    Notes. We have kept the above two taxa separate since they differ slightly in ascospore size and also in distribution, but further data are needed to test whether this separation can be maintained. A collection from French Guiana (LeGallo 4532, FH!) is somewhat intermediate and would support merging the two taxa.

26. 24(22) Lirellae with apically thin complete thalline margin, stellately branched (evirescens- morph); thallus green; unknown substances; Neotropics [holotype: Brazil, Malme 3527 (S 5540!)] ... Graphis evirescens (Redinger) Lücking*

    Lirellae with basal to lateral thalline margin, sparsely branched (hossei- or lineola- morph); thallus white-grey; substances absent ... 25

27. 25(24) Lirellae with basal thalline margin (hossei-morph); ascospores 30–40 μm long; Eastern Paleotropics [holotype: Thailand, Schmidt s.n. (TUR-Vainio 27256!)] ... ... Graphis consimilis Vain.

    Lirellae with lateral thalline margin (lineola-morph); ascospores 30–50 μm long; pantropical [lectotype (here selected): Philippines, Copeland 1382 (TUR-Vainio 27261!)] ... ... Graphis nanodes Vain.

Group 6: Labia entire, excipulum laterally carbonized, hymenium inspersed, ascospores transversely septate

1. 1 Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   No substances (K−) ... 9

2. 2 (1) Stictic acid (K+ yellow) ... 3

   Norstictic acid (K+ yellow forming red crystals) ... 4

3. 3 (2) Ascospores small (20–40 × 6–8 μm); lirellae erumpent, with lateral thalline margin, short and sparsely branched (lineola-morph); pantropical [lectotype (Wirth & Hale Reference Wirth and Hale1978): South America, s.col. (G!)] ... Graphis leptocarpa Fée

   Ascospores medium-sized (40–60 × 10–14 μm); lirellae erumpent, with lateral to complete thalline margin, short to elongate and sparsely to irregularly branched (subserpentina-morph); Eastern Palaeotropics [holotype: Taiwan, Faurie 30 (W!)] ... ... Graphis kelungana Zahlbr.

4. 4 (2) Ascospores (small to) medium-sized (40–70 μm long); lirellae erumpent, with lateral thalline margin, elongate and irregularly branched; disc exposed, white-pruinose (scripta-morph); Australia [holotype: Australia, Streimann 48626 (CANB!)] ... ... Graphis streimannii A. W. Archer

   Ascospores small to very small (15–45 × 5–9 μm); lirellae variable ... 5

5. 5 (4) Disc exposed; lirellae erumpent, with lateral thalline margin, short to elongate and sparsely to irregularly branched (handelii- or scripta-morph) ... 6

   Notes. This is a difficult complex including four described names whose types differ chiefly in the pruinosity of the disc and (slightly) in ascospore size. We have separated two taxa, one with pruinose disc and one with non-pruinose disc. The slight differences in ascospore size between the type material of Graphis crebra and Graphis apertella are not considered taxonomically important, since both fall within the typical range of small-sized ascospores.

   Disc concealed; lirellae variable ... 7

6. 6 (5) Disc white-pruinose (scripta-morph); pantropical ... Graphis crebra Vain.

   1. A. Ascospores 5–9-septate, 20–30 × 5–8 μm [holotype: Guadeloupe, Duss 541 (TUR-Vainio 27617!)] ... Graphis crebra Vain.

   2. B. Ascospores 7–11-septate, 25–45 × 6–9 μm [holotype: Australia, Streimann 9367 (CANB!)] ... ... Graphis apertella A. W. Archer

   Disc non-pruinose (handelii-morph); pantropical ... Graphis handelii Zahlbr.

   1. A. Excipulum laterally (to almost completely) carbonized; ascospores 7–9-septate, 30–45 × 6–9 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Paraguay, Malme 1900 (S- 6496!)] ... ... Graphis lineola var. chondroplaca Redinger

   2. B. Excipulum laterally carbonized; ascospores 7–11-septate, 25–40 × 7–10 μm [holotype: India,Watt s.n. (BM, not seen; Staiger Reference Staiger2002)] ... ... Schistostoma dehiscens Stirt. [non Graphis dehiscens Vain.]

   3. C. Excipulum laterally carbonized, ascospores 5–11-septate, 20–40 × 6–9 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): China, Handel-Mazzetti 12788 (W!; isolectotype: S 2181!)] ... Graphis handelii Zahlbr.

      Notes. Initially we took up the epithet chondroplaca for this taxon (Lücking et al. Reference Lücking and Rivas Plata2008), but revision of the lectotype material of G. handelii, first considered a synonym of G. lineola, showed more or less exposed discs and norstictic acid and hence handelii becomes the valid epithet for the species.

7. 7 (5) Labia white-pruinose; lirellae immersed, with lateral thalline margin (caesiella- morph); Neotropics [holotype: Guadeloupe, Duss 1189 (TUR-Vainio 27732!)] ... ... Graphis plumierae Vain.

   Labia non-pruinose; lirellae variable ... 8

8. 8 (7) Lirellae short and broad (up to 0·6 mm broad), immersed, with apically thick complete thalline margin (subserpentina-morph); ascospores very small (15–20 μm long); Oceania [holotype: Vanuatu, Kashiwadani 43246 (TNS, not seen; Nakanishi et al. Reference Nakanishi, Kashiwadani and Moon2002)] ... Graphis brevicarpa M. Nakan., Kashiw. & K. H. Moon

   Lirellae elongate and slender (up to 0·25 mm broad), erumpent, with basal to (thick) lateral thalline margin (lineola-morph); ascospores small (25–45 μm long); pantropical ... Graphis cincta (Pers.) Aptroot*

   1. A. Lirellae with lateral thalline margin; ascospores 7–9-septate, 25–35 × 7–8 μm [holotype: Dominican Republic, s.col. (L!; isotypes: L!)] ... Opegrapha cincta Pers.

   2. B. Lirellae with basal to lateral thalline margin; ascospores 7–9-septate, 35–45 × 7–9 μm [holotype: Dominica, Raunkiaer 490 (TUR-Vainio 27701!; isotype: FH!)] ... Graphis tenellula Vain.

   3. C. Lirellae with lateral thalline margin; ascospores 7–11-septate, 25–45 × 7–9 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): Philippines, Merrill 6716 (TUR- Vainio 27811!)] ... Graphis guimarana Vain.

   4. D. Lirellae with basal thalline margin; ascospores 7–9-septate, 25–35 × 6–8 μm [holotype: Philippines, Merrill 6344 (TUR-Vainio 27722!)] ... Graphis tenella var. jatrophae Vain.

   5. E. Lirellae erumpent, with lateral thalline margin; ascospores 30–40 × 6–8 μm, 5–9-septate [holotype: Taiwan, Faurie 291 (W!)] ... Graphis latibasa Zahlbr.

   6. F. Lirellae with lateral thalline margin; ascospores 5–9-septate, 25–30 × 5–7 μm [lectoype (here selected): Paraguay, Malme 1663Bc (S 6500!)] ... Graphis inspersa Redinger

   7. G. Lirellae with thick lateral thalline margin; ascospores 7–9-septate, 35–45 × 6–8 μm [original material: Philippines, Merrill 6736 (TUR-Vainio 27706!)] ... ... Graphis tenellula var. eutypodes Vain. [nom. inval. (not published)]

      Notes. Opegrapha cincta was found to be the oldest epithet available for this taxon usually known as G. tenellula.

9. 9 (1) Ascospores (small to) medium-sized (40–80 × 8–12 μm), 11–17-septate ... 10

   Ascospores small (20–40 × 6–9 μm), 5–11-septate ... 12

10. 10 (9) Lirellae erumpent, with basal thalline margin (labia black), short and sparsely branched (subregularis-morph); Australia [holotype: Australia, Elix 17392 (CANB!)] ... ... Graphis subregularis A. W. Archer

    Lirellae distinctly elongate, partly to radiately branched, with (thick) lateral thalline margin ... 11

11. 11(10) Lirellae with lateral thalline margin, very long and radiately branched (centrifuga- morph); ascospores 50–80 μm long; Neotropics [holotype: Jamaica, Imshaug 14632 (MSC-0029034!)] ... Graphis inspersoradians Lücking

    Lirellae with thick lateral thalline margin, short and sparsely branched (subserpen- tina-morph); ascospores 40–55 μm long; Eastern Palaeotropics [holotype: Thailand, Hosseus s.n. (TUR-Vainio 27214!)] ... Graphis intermedians Vain.

12. 12(9) Disc exposed, white-pruinose; lirellae erumpent, with lateral thalline margin (scripta-morph); pantropical [holotype: Indonesia (Java), s.col. (not seen)] ... ... Graphis submarginata Lücking [≡ Graphis marginata G. Mey. & Flot., nom. illeg., non Raddi]

    Disc concealed, labia non-pruinose; lirellae erumpent, with lateral thalline margin (lineola-morph); pantropical ... Graphis lineola Ach.

    1. A. Lirellae with lateral thalline margin; ascospores 7–9-septate, 20–40 × 6–8 μm [holotype: Lesser Antilles, Swartz s.n. (H-Acharius 584!)] ... Graphis lineola Ach.

    2. B. Lirellae with lateral thalline margin; ascospores 7–9-septate, 25–35 × 5–8 μm [holotype: Sierra Leone, Deighton 4307E (FH!)] ... Graphis deightonii C. W. Dodge

Group 7: Labia entire, excipulum laterally carbonized, hymenium inspersed, ascospores muriform

1. 1 Norstictic, stictic, or protocetraric acid (K+ yellow or K+ yellow forming red crystals or P+ red) ... 2

   No substances (K−, P−) ... 6

2. 2(1) Norstictic acid (K+ yellow forming red crystals) ... 3

   Stictic or protocetraric acid (K+ yellow or P+ red) ... 4

3. 3(2) Ascospores small (25–40 × 8–14 μm), 8 per ascus; lirellae erumpent, with apically thin thalline margin, short and sparsely branched (negrosina-morph); thallus verrucose; Eastern Palaeotropics including Australia [holotype: Malaysia (Borneo), Beccari 165 (M!; isotype: W!)] ... Graphis pertricosa (Kremp.) A. W. Archer

   Ascospores large (50–110 × 15–30 μm), 1–2 per ascus; lirellae immersed to erumpent, with thick lateral to complete thalline margin, elongate and irregularly branched (subserpentina-morph); pantropical ... ... Graphis insulana (Müll. Arg.) Lücking

   1. A. Lirellae erumpent, with thick lateral thalline margin; ascospores 1–2/ascus, 60–90 × 20–30 μm [holotype: Fiji, s.col. (G!)] ... Graphina insulana Müll. Arg.

   2. B. Lirellae immersed, with lateral thalline margin; ascospores single, 80–100 × 18–28 μm [holotype: Australia, Sayer s.n. (G!)] ... Graphina subtartarea Müll. Arg.

   3. C. Lirellae erumpent, with thick lateral thalline margin; ascospores single, 50–100 × 20–30 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): São Tomé & Principé, Fr. Quintas s.n. (H-Nylander 7414!)] ... Graphina rudescens Nyl.

   4. D. Lirellae erumpent, with lateral thalline margin; ascospores single, 80–100 × 20–30 μm [holotype: Costa Rica, Pittier 5238 (G!)] ... Graphina robusta Müll. Arg.

   5. E. Lirellae erumpent, with thick lateral thalline margin; ascospores 1–2/ascus, 55–110 × 15–30 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Philippines, McGregor 18445 (TUR-Vainio 27230!)] ... ... Graphis orientalis Vain.

   6. F. Lirellae immersed, with thick lateral thalline margin; ascospores single, 50–110 × 15–25 μm [holotype: Mariana Islands (Guam), McGregor 607 (TUR-Vainio 27242!)] ... ... Graphis macgregorii Vain.

   7. G. Lirellae erumpent, with thick lateral thalline margin; ascospores single, 70–100 × 20–30 μm [holotype: Sierra Leone, MacDonald s.n. (FH!)] ... Graphina sierraleonensis C. W. Dodge

   8. H. Lirellae erumpent, with thick lateral thalline margin; ascospores single, 80–100 × 20–30 μm [holotype: India, Patwardhan & Nagarkar 74.3053 (AMH!)] ... ... Graphina nylanderi Patw. & C. R. Kulk.

4. 4(2) Protocetraric acid (P+ red); lirellae prominent, with thick lateral thalline margin, very short and unbranched (dussii-morph); ascospores large (90–105 × 20–30 μm); Eastern Palaeotropics [holotype: Philippines, Elmer 14924 (TUR-Vainio 27262!; isotype: FH!)] ... Graphis sorsogona Vain.

   Stictic acid (K+ yellow); lirellae variable; ascospores small to large (35–150 × 11–25 μm) ... 5

5. 5(4) Ascospores 8 per ascus, small to medium-sized (35–50 × 11–13 μm); lirellae erumpent, with lateral thalline margin, short and sparsely branched (lineola-morph); thallus yellow-green; Australia [holotype: Australia, Fawcett s.n. (M!)] ... ... Graphis polyclades Kremp.

   Ascospores 1–2 per ascus, medium-sized to large (50–130 × 15–25 μm); lirellae erumpent, with thick complete thalline margin, short to elongate and sparsely to irregularly branched (subserpentina-morph); thallus white-grey with yellow tinge; Palaeotropics [holotype: Philippines, McGregor 8605 (TUR-Vainio 27239!)] ... ... Graphis diplocheila Vain.

6. 6(1) Ascospores small to medium-sized (35–50 × 8–15 μm), lirellae erumpent, with lateral thalline margin, short to elongate and sparsely to irregularly branched (lineola- or deserpens-morphs); Palaeotropics (also saxicolous) ... ... Graphis subvelata Stirt.

   1. A. Lirellae erumpent, with lateral thalline margin; ascospores 35–45 × 8–12 μm [lectotype (Rogers 1982): Australia, Bailey s.n. (BM!)] ... Graphis subvelata Stirt.

   2. B. Lirellae erumpent, with thick lateral thalline margin; ascospores 40–50 × 10–15 μm [holotype: Angola, Welwitsch 392 (TUR-Vainio 27197!)] ... Graphis ambrizensis Vain.

   Ascospores very large (75–250 × 25–30 μm); lirellae prominent, with thin complete thalline margin, short and sparsely branched (negrosina-morph); Neotropics [holotype: Brasil, Cáceres & Lücking 01-0514 (F!; isotype: B!)] ... ... Graphis pilarensis Cáceres & Lücking

Group 8: Labia entire, excipulum completely carbonized, hymenium clear, ascospores transversely septate

1. 1 Ascospores medium-sized to large (45–120 μm long, mature ascospores exceeding 50 μm) ... 2

   Ascospores small (15–50 μm long, mature ascospores rarely exceeding 45 μm) ... 29

2. 2 (1) Lichexanthone (UV+ yellow); lirellae prominent, with basal thalline margin or thalline margin lacking (hossei-morph); ascospores 8 per ascus, 7–15-septate, 45–60 ×10–14 μm; Neotropics [holotype: Brazil, Spruce 331 (BM, not seen; Staiger Reference Staiger2002)] ... Graphis sauroidea Leight.

   Notes. If additionally protocetraric acid and ascospores I− and up to 100 μm long, cf. Carbacanthographis candidata (Nyl.) Staiger & Kalb; if additionally protocetraric acid and ascospores I− and 20–25 × 6–8 μm, cf. Carbacanthographis chionophora (Redinger) Staiger & Kalb.

   Lichexanthone absent (UV−); lirellae and ascospores variable ... 3

3. 3 (2) Norstictic, stictic, and/or salazinic acids (K+ yellow or K+ yellow forming red crystals) ... 4

   No substances (K−) ... 12

4. 4 (3) Stictic acid, norstictic and salazinic acids absent (K+ yellow) ... 5

   Norstictic and/or salazinic acid present, sometimes additionally stictic acid (K+ yellow forming red crystals) ... 7

   Notes. If protocetraric acid and ascospores I−, cf. Carbacanthographis candidata (Nyl.) Staiger & Kalb.

5. 5 (4) Lirellae prominent, with thick lateral thalline margin, elongate and irregularly branched (marginata-morph); pantropical ... Graphis rustica Kremp.

   1. A. Ascospores 8 per ascus, 11–17-septate, 65–85 × 8–11 μm [lectotype (Archer Reference Archer2006 based on annotation label by Nakanishi in 1973): Singapore, Beccari 258 (M!)] ... Graphis rustica Kremp.

   2. B. Ascospores 6–8 per ascus, 9–17-septate, 60–100 × 10–14 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Mauritius, s.col. (BM!)] ... Graphis turgidula Müll. Arg.

   3. C. Ascospores 4–8 per ascus, 15–19-septate, 65–100 × 9–12 μm [holotype: Philippines, Merrill 7958 (TUR-Vainio 27867!)] ... Graphis marginifera Vain.

   4. D. Ascospores 4–8 per ascus, 9–17-septate, 40–80 × 9–12 μm [holotype: Philippines, Merrill 7985 (TUR-Vainio 27863!)] ... Graphis tonglonensis Vain.

   5. E. Ascospores 4–8 per ascus, 13–19-septate, 80–120 × 11–15 μm [holotype: Indonesia (Celebes), Kjellberg 3L (S-2178!)] ... Graphis kjellbergii Redinger

      Notes. There is a probably undescribed species from Taiwan with similar lirellae anatomy and chemistry but with very long and radiately branched lirellae.

   Lirellae immersed to erumpent, with lateral or apically thick complete thallin margin (caesiella- or subserpentina-morph) ... 6

6. 6 (5) Ascospores medium-sized (50–70 × 7–11 μm); lirellae with lateral thalline margin, labia thinly white-pruinose (caesiella-morph); Eastern Palaeotropics ... ... Graphis subassimilis Müll. Arg.

   1. A. Ascospores 9–17-septate, 50–70 × 7–10 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): Indonesia (Java), L. B. 445 (L!; isolectotype: G!)] ... ... Graphis subassimilis Müll. Arg.

   2. B. Ascospores 13–17-septate, 55–70 × 9–11 μm [type: Indonesia (Java), Overeem s.n. (W, not seen; Hale Index Cards)] ... Graphis overeemii Zahlbr.

   3. C. Ascospores 11–15-septate, 45–65 × 7–11 μm [holotype: Taiwan, Faurie 243 (W!)] ... ... Graphis formosana Zahlbr.

   Ascospores large (70–90 × 10–14 μm); lirellae with apically thick complete thalline margin (subserpentina-morph); Palaeotropics [holotype: Tanzania, Schröder 160 (W!)] ... ... Graphis schroederi Zahlbr.

   Notes. According to the original description and the information provided by Redinger (Reference Redinger1935), Graphis balansana Müll. Arg. might represent an earlier name for G. schroederi, but we have not seen type material of the former.

7. 7 (4) Lirellae sessile, lacking or with basal thalline margin, very short and unbranched (nuda-morph); ascospores 100–200 μm long; Neotropics [holotype: Ecuador (Galápagos), Bungartz 4701 (CDS!)] ... Graphis pedunculata Bungartz & Aptroot

   Lirellae erumpent to prominent, with basal to complete thalline margin (hossei-, marginata-, or subserpentina-morphs); ascospores 45–120 μm long ... 8

8. 8 (7) Ascospores 60–120 μm long; lirellae prominent, with thick lateral thalline margin, elongate and irregularly branched (marginata-morph); norstictic acid; pantropical ... Graphis marginata Raddi

   1. A. Lirellae with thick lateral thalline margin; ascospores 4–8 per ascus, 11–25-septate, 60–120 × 9–18 μm [holotype: Brazil, s.col. (not seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... Graphis marginata Raddi

   2. B. Lirellae with thick lateral thalline margin; ascospores 8 per ascus, 15–29-septate, 60–120 × 10– 17 μm [lectotype (Archer Reference Archer2006): Indonesia (Borneo), Beccari 134 (M!)] ... Graphis leucoparypha Kremp.

   3. C. Lirellae with thick lateral thalline margin; ascospores 8 per ascus, 15–19-septate, 60–85 × 10–12 μm [holotype: Australia, Streimann 56877 (CANB!)] Graphis turgidula var. norstictica A. W. Archer

      Notes. If lirellae with thick white pruina and ascospores I–, cf. Carbacanthographis induta (Müll. Arg.) Lücking*.

   Ascospores 45–70 μm long; lirellae erumpent, if prominent then with basal thalline margin (hossei-morph); norstictic or salazinic acid ... 9

9. 9 (8) Disc exposed, white-pruinose (scripta-morph); lirellae otherwise with complete thalline margin; norstictic acid; Eastern Palaeotropics [holotype: India, Patwardhan & Nagarkar 73.2254 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... ... Graphis albidofarinacea Adaw. & Makhija

   Disc concealed, labia non-pruinose ... 10

10. 10(9) Lirellae prominent, with basal thalline margin, short and sparsely branched (hossei- morph); norstictic acid; Eastern Palaeotropics [holotype: India, Nagarkar & Patwardhan 85.1526 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... ... Graphis kollaimalaiensis Adaw. & Makhija

    Lirellae erumpent, with thick lateral to complete thalline margin (subserpentina- morph) ... 11

11. 11(10) Lirellae with thick lateral thalline margin; norstictic acid; Eastern Palaeotropics [holotype: India, Nagarkar & Gole 76.623 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... Graphis nigrocarpa Adaw. & Makhija

    Lirellae with complete thalline margin; salazinic and stictic acids; Eastern Palaeotropics [holotype: India, Patwardhan & Nagarkar 77.1334 (AMH, photograph seen; Nagarkar & Patwardhan Reference Nagarkar and Patwardhan1982; Awasthi Reference Awasthi1991)] ... Graphis assamensis Nagarkar & Patw.

12. 12(3) Ascospores large (70–300 μm) ... 13

    Ascospores medium-sized (40–70 μm) ... 19

13. 13(12) Lirellae sessile, lacking thalline margin, very short and unbranched (nuda-morph) ... 14

    Lirellae prominent to sessile, with basal to (thick) lateral or complete thalline margin (rarely thalline margin flaking off to expose black labia), short to elongate and sparsely to irregularly branched ... 15

14. 14(13) Ascospores 15–21-septate, 80–100 μm long; Neotropics and Eastern Palaeotropics [holotype: China, Handel-Mazzetti 12262 (W!; isotype: US!)] ... Graphis bifera Zahlbr.

    Notes. According to Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b), Graphis subdura M. Nakan. would key out in our treatment as either a synonym of G. bifera or G. adpressa, depending on the length of the ascospores; we have not seen the original publication or type, but the ascospores of G. subdura are given by Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b) as either less than 60 μm or more than 65 μm.

    Ascospores 25–31-septate, 100–140 μm long; Eastern Palaeotropics [holotype: Indonesia (Java), s.col. (W, not seen; Hale Index Cards)] ... Graphis curtiuscula Zahlbr.

15. 15(13) Lirellae prominent, with basal thalline margin, short and sparsely branched (Opegrapha-morph); Eastern Palaeotropics [holotype: Japan, Yatabe 256 (G!)] ... ... Graphis parallela Müll. Arg.

    Lirellae prominent to sessile, with complete thalline margin, elongate and irregularly branched ... 16

16. 16(15) Ascospores very large (120–300 μm long); Neotropics ... ... Graphis tumidula (Fée) Spreng.

    1. A. Thallus smooth to uneven [holotype: Peru, s.col. (G!)] ... Opegrapha tumidula Fée

    2. B. Thallus uneven to rugulose [holotype: South America, s.col. (G, not seen; Fée Reference Fée1824, Tab. VII, Fig. 1, 1a; isotype: L!)] ... Opegrapha rugulosa Fée

    Ascospores large (70–130 μm long) ... 17

17. 17(16) Lirellae sessile, very short to short and unbranched to rarely sparsely branched; thalline margin sometimes flaking off to expose black labia (nudaeformis-morph); Neotropics (Central America) [holotype: Costa Rica, Lücking 18025 (INB!)] ... ... Graphis nudaeformis Lücking

    Lirellae prominent, short to elongate and irregularly to stellately branched; thalline margin persistent ... 18

18. 18(17) Lirellae short and stellately branched (evirescens-morph); Neotropics [holotype: Peru, s.col. (G, not seen; Fée Reference Fée1824, Tab. XIII, Fig. 1, 1a; isotype: L!)] ... ... Graphis conglomerata Spreng.[≡ Opegrapha conglomerata Fée, nom. illeg., non Pers.]

    Lirellae elongate and irregularly branched (rhizocola-morph); Neotropics ... ... Graphis rhizocola (Fée) Lücking & Chaves

    1. A. Ascospores not seen [holotype: South America, s.col. (G, not seen; Fée Reference Fée1824, Tab. XIII, Fig. 2, 2a)] ... Opegrapha rhizocola Fée

    2. B. Ascospores 11–19-septate, 70–100 × 10–20 μm [holotype: Peru, s.col. (not seen; Zenker Reference Zenker, Goebel and Kunze1829; Müller Argoviensis Reference Müller Argoviensis1887)] ... Graphis cooperta Zenker

    3. C. Ascospores 19–25-septate, 70–130 × 11–20 μm [holotype: Caribbean (St. Vincent), s.col. (FH!)] ... Graphis anguilliformis Taylor

    4. D. Ascospores 15–25-septate, 70–130 × 11–20 μm [holotype: Brazil, Glaziou 6279 (G!)] ... ... Graphis serpens Fée

    5. E. Ascospores 13–19-septate, 70–120 × 10–20 μm [original material: Trinidad and Tobago (Trinidad), Thaxter 92 (TUR-Vainio 27529!)] ... ... Graphis anguillaeformis var. infecunda Vain. [nom. inval. (not published)]

    6. F. Ascospores 15–19-septate, 70–80 × 10–13 μm [holotype: Brazil, Malme 634 (S-6501!)] ... ... Graphis illota var. leopoldensis Redinger

19. 19(12) Lirellae prominent to sessile, lacking thalline margin or very rarely with basal thalline margin ... 20

20. Lirellae immersed to prominent, with basal to (thick) lateral thalline margin ... 23

21. 20(19) Lirellae short and sparsely branched (hossei-morph); Eastern Palaeotropics [lectotype (here selected): Japan, Yatabe 260 (G, not seen; isolectotype: US!)] ... ... Graphis cognata Müll. Arg.

    Lirellae very short and unbranched (nuda-morph) ... 21

22. 21(20) Ascospores grey-brown; Neotropics (Central America) [holotype: Costa Rica, Lücking 15247g (INB!; isotype: F!)] ... ... Graphis pittieri Lücking, Umaña, Sipman & Chaves

    Ascospores hyaline ... 22

23. 22(21) Ascospores with terminal gelatinous caps, 40–60 × 8–13 μm; Neotropics ... ... Graphis ovata (Fée) A. Massal.

    1. A. Ascospores 9–11-septate, 40–60 × 10–12 μm [holotype: Peru, s.col. (G!)] ... Opegrapha ovata Fée

    2. B. Ascospores 7–11-septate, 40–50 × 8–13 μm [original material: Brazil, Kalb & Plöbst 31277 (hb. Kalb!)] ... Graphis nuda Staiger [nom. inval. (not validly published)]

    Ascospores lacking gelatinous caps, 50–70 × 12–15 μm; Neotropics [holotype: Brazil, Vainio s.n. (TUR-Vainio 27851!)] ... Graphis adpressa Vain.

    Notes. According to Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b), Graphis subdura M. Nakan. would key out in our treatment as either a synonym of G. bifera or G. adpressa, depending on the length of the ascospores; we have not seen the original publication or type, but the ascospores of G. subdura are given by Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b) as either less than 60 μm or more than 65 μm.

24. 23(19) Lirellae very short and unbranched (subregularis- or dussii-morph) ... 24

    Lirellae elongate and irregularly branched ... 25

25. 24(23) Lirellae erumpent with basal thalline margin (subregularis-morph); Neotropics and Eastern Paleotropics [lectotype (here selected): South America, s.col. (G; Fée Reference Fée1824, Tab. VI, Fig. 3 and 3a); Müller Argoviensis Reference Müller Argoviensis1887)] ... Graphis subimmersa (Fée) A. Massal.

    Lirellae prominent with thick lateral thalline margin (dussii-morph); Eastern Palaeotropics [holotype: Indonesia (Java), s.col. (G!)] ... Graphis regularis Müll. Arg.

26. 25(23) Lirellae immersed to erumpent, with lateral thalline margin, elongate and irregularly branched (deserpens-morph); Eastern Palaeotropics ... ... Graphis subdisserpens Nyl.

    1. A. Ascospores 9–17-septate, 40–70 × 6–10 μm [holotype: India, Kurz 37 (H-Nylander 7156!)] ... ... Graphis subdisserpens Nyl.

    2. B. Ascospores 13–15-septate, 45–65 × 8–12 μm [holotype: Australia, Hartmann 77 (G!; isotype: MEL!)] ... Graphis propinqua Müll. Arg.

    3. C. Ascospores 11–15-septate, 55–70 × 6–8 μm [holotype: India, Patwardhan & Adawadkar 87.314 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis dispersa Makhija & Adaw. [nom. illeg., non Redinger]

    Notes. Graphis propinqua was treated as a separate species by Archer (Reference Archer2006) but is identical with G. subimmersa.

    Lirellae prominent, with (thick) lateral to complete thalline margin ... 26

27. 26(25) Labia white pruinose along slit (farinulenta-morph); thallus usually matt, ecorticate; pantropical ... Graphis seminuda Müll. Arg.

    1. A. Lirellae prominent; ascospores 11–13-septate, 40–65 × 8–11 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Costa Rica, Pittier 5241 (G!)] ... Graphis seminuda Müll. Arg.

    2. B. Lirellae (erumpent to) prominent; ascospores 11–15-septate, 45–65 × 8–11 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Costa Rica, Pittier s.n. (G!)] ... ... Graphis seminuda var. sublaevis Müll. Arg.

    3. C. Lirellae (erumpent to) prominent; ascospores 11–15-septate, 45–70 × 8–11 μm [holotype: Australia, Streimann 28307 (CANB!)] ... Graphis catherinae A. W. Archer

    Labia non-pruinose; thallus slightly nitidous ... 27

28. 27(26) Lirellae prominent, with thick lateral thalline margin, elongate and irregularly branched (marginata-morph); pantropical ... Graphis flavens Müll. Arg.

    1. A. Ascospores 11–15-septate, 50–70 × 9–11 μm [holotype: Indonesia (Java), Junghuhn 446 (L!; isotype: G!)] ... Graphis flavens Müll. Arg.

    2. B. Ascospores 13–19-septate, 60–80 × 10–14 μm [holotype: Guadeloupe, Duss 530 (TUR-Vainio 27845!)] ... Graphis angustata var. denudata Vain.

    3. C. Ascospores 13–19-septate, 60–80 × 7–11 μm [holotype: Indonesia (Java), Kjellberg 101L (S-2166!)] ... Graphis subintegra Redinger

    4. D. Ascospores 9–15-septate, 40–60 × 7–12 μm [original material: not designated] ... ... Graphis atrolabiata Chaves & Lücking [nom. inval. (not published)]

    Lirellae erumpent to prominent, with complete thalline margin ... 28

29. 28(27) Lirellae with apically thin complete thalline margin, very long and radiately branched (anguilliradians-morph); Neotropics [holotype: Trinidad and Tobago (Trinidad), Imshaug & Imshaug 31858 (MSC-0024122!)] ... Graphis anguilliradians Lücking

    Lirellae with apically thick complete thalline margin, short and sparsely branched (illinata-morph); Eastern Paleotropics [holotype: Australia, Archer 747 (NSW!)] ... ... Graphis subcelata A. W. Archer

30. 29(1) Norstictic, salazinic, and/or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 30

    Notes. If protocetraric acid (P+ red), cf. Carbacanthographis hertelii Kalb & Staiger.

    No substances (K−) ... 43

31. 30(29) Stictic acid (K+ yellow) ... 31

    Norstictic and/or salazinic acid, sometimes additionally stictic acid (K+ yellow forming red crystals) ... 36

32. 31(30) Disc exposed, white-pruinose (scripta-morph); Palaeotropics [type: Solomon Islands, s.col. (W!)] ... Graphis modesta Zahlbr.

    Disc concealed ... 32

33. 32(31) Lirellae with thick complete thalline margin (labia with conspicuous white cover); Eastern Palaeotropics [holotype: Solomon Islands, Hill 8400 (BM!)] ... ... Graphis discarpa A. W. Archer

    Lirellae with (thick) lateral thalline margin; labia black or white-pruinose ... 33

    Notes. If ascospores I− and lirellae with thick pruinose cover, cf. Carbacanthographis stictica Staiger.

34. 33(32) Labia white-pruinose (caesiella- or glaucescens-morph) ... 34

    Labia non-pruinose ... 35

35. 34(33) Thallus ecorticate (glaucescens-morph); Neotropics [holotype: Mexico, Pringle 24 (W!)] ... ... Graphis subamylacea Zahlbr.

    Thallus corticate; lirellae with split between labia and thalline margin; Neotropics [holotype: Antilles (Dominica), Imshaug 32724B (MSC!)] ... ... Graphis imshaugii M. Wirth & Hale

36. 35(33) Lirellae very long and radiately branched, lateral thalline margin thin (centrifuga- morph); Neotropics and Eastern Palaeotropics [holotype: India, Nagarkar & Sethy 85.2264 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis flavovirens Makhija & Adaw.

    Lirellae elongate and irregularly branched, lateral thalline margin thick (subserpentina-morph); Neotropics (Antilles) and Eastern Palaeotropics (Australia) [holotype: Australia, Knight 339 (G!)] ... Graphis descissa Müll. Arg.

    Notes. Graphis ocellata Zahlbr. may belong here but we have not been able to track down the type material.

37. 36(30) Lirellae prominent, lacking thalline margin, very short and unbranched (nuda-morph); norstictic acid; pantropical [holotype: Australia, Bailey 217 (G!)] ... ... Graphis emersa Müll. Arg.

    1. A. Ascospores 7–11-septate, 30–40 × 7–9 μm [holotype: Australia, Bailey 217 (G!)] ... ... Graphis emersa Müll. Arg.

    2. B. Ascospores 7–9-septate, 25–35 × 7–8 μm [holotype: Japan (HIRO, not seen; Nakanishi Reference Nakanishi1966, Reference Nakanishi1967; Nakanishi et al. Reference Nakanishi, Kashiwadani and Moon2003b)] ... Graphis meridionalis M. Nakan.

    Lirellae immersed to erumpent, with lateral to apically thin complete thalline margin or rarely lacking thalline margin ... 37

38. 37(36) Norstictic acid, additionally with salazinic and/or stictic acid ... 38

    Norstictic or salazinic acid only ... 40

39. 38(37) Lirellae erumpent, lacking thalline margin, short and sparsely branched (subregularis- morph); norstictic and salazinic acids; Eastern Palaeotropics [holotype: India, Patwardhan & Prabhu 75.184 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... ... Graphis cinnamomea Adaw. & Makhija

    Lirellae immersed to erumpent, with lateral thalline margin; norstictic and stictic acids ... 39

40. 39(38) Lirellae erumpent, with thick lateral thalline margin, elongate and irregularly branched (subserpentina-morph); ascospores 25–45 μm long; Eastern Palaeotropics [holotype: India, Patwardhan & Prabhu 75.220 (AMH, photograph seen; Awasthi Reference Awasthi1991)] ... Graphis asahinae Patw. & C. R. Kulk.

    Lirellae immersed to erumpent, with lateral thalline margin, short and sparsely branched (lineola-morph); ascospores 15–25 μm long; Eastern Palaeotropics (often saxicolous) [holotype: Japan, Yatabe 266 (G, not seen; Staiger Reference Staiger2002)] ... ... Graphis cervina Müll. Arg.

41. 40(37) Salazinic acid; disc partly exposed (handelii-morph); Eastern Palaeotropics [holotype: Philippines, Baker 545 (TUR-Vainio 27868!; isotype: FH!)] ... Graphis ficicola Vain.

    Norstictic acid; disc concealed ... 41

42. 41(40) Labia white-pruinose (caesiella-morph); Neotropics and Eastern Palaeotropics (Papua New Guinea) [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Malme 3602 (S 6506!)] ... ... Graphis caesiocarpa Redinger

    Labia non-pruinose ... 42

43. 42(41) Lirellae short, sparsely branched (lineola-morph); ascospores 30–45 μm long; Eastern Palaeotropics ... Graphis assimilis Nyl.

    1. A. Lirellae erumpent, with lateral thalline margin; excipulum (laterally to) completely carbonized; ascospores 7–11-septate, 30–45 × 7–9 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): New Caledonia, Viellard s.n. (H-Nylander 7125!)] ... Graphis assimilis Nyl.

    2. B. Lirellae prominent, with thick lateral thalline margin; excipulum completely carbonized; ascospores 7–11-septate, 35–45 × 7–11 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): Indonesia (Java), Van Overeem s.n. (W!)] ... Graphis inamoena Zahlbr.

    3. C. Lirellae erumpent, with lateral thalline margin; excipulum completely carbonized; ascospores 7–9-septate, 30–40 × 6–8 μm [holotype: China, Chung 596f (W!)] ... ... Graphis spodoplaca Zahlbr.

    4. D. Lirellae erumpent, with thick lateral thalline margin; excipulum completely carbonized; ascospores 5–11-septate, 35–50 × 4–6 μm [holotype: India, Patwardhan & Nagarkar 85.1756 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... Graphis verruciformis Adaw. & Makhija

       Notes. Graphis inamoena was treated as separate species by Archer (Reference Archer2006), but G. assimilis is an older name for this taxon. Lücking et al. (Reference Lücking and Rivas Plata2008) listed G. spodoplaca as synonym of G. tenellula (= G. cincta), but the holotype has a completely carbonized excipulum and clear hymenium.

    Lirellae elongate to very long and irregularly to radiately branched (deserpens- or centrifuga-morph); ascospores 15–30 μm long; pantropical ... ... Graphis intricata Fée

    1. A. Lirellae very long and radiately branched; ascospores 5–7-septate, 15–30 × 6–8 μm [lectotype: South America, s.col. (G!; isolectotype: H-Nylander 7142!)] ... Graphis intricata Fée

    2. B. Lirellae elongate and irregularly branched; ascospores 7-septate, 20-30 × 6–8 μm [holotype: New Zealand, s.col. (not seen; Hale Index Cards)] ... Graphis confinis C. Knight & Mitt.

    3. C. Lirellae elongate and irregularly branched; ascospores 5–7-septate, 20-30 × 6–8 μm [holotype: U.S.A. (Hawaii), Rock 99 (W!)] ... Graphis tapetica Zahlbr.

    4. D. Lirellae very long and radiately branched [holotype (saxicolous): China, Handel-Mazzetti 12233 (W, not seen; Zahlbruckner Reference Zahlbruckner and Handel-Mazetti1930)] ... Graphis hunana Zahlbr.

    5. E. Lirellae elongate and irregularly branched; ascospores 5–7-septate, 25–30 × 4–6 μm [holotype: India, Patwardhan 76.701 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... ... Graphis colliculoides Adaw. & Makhija

44. 43(29) Lirellae immersed in pseudostromata, stellately branched (hyphosa-morph); labia white-pruinose; Neotropics ... Graphis hyphosa Staiger

    1. A. Ascospores 5–9-septate, 20–40 × 6–9 μm [holotype: Brazil, Kalb s.n. (hb. Kalb 28859; Staiger Reference Staiger2002)] ... Graphis hyphosa Staiger

    2. B. Ascospores 7–11-septate, 30–40 × 7–9 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Malme 2357 (S 6499!)] ... Graphis intricata f. meizospora Redinger

    Lirellae erumpent to prominent, unbranched to irregularly branched, if stellately branched then not forming pseudostromata and labia non-pruinose ... 44

45. 44(43) Lirellae sessile, lacking thalline margin, very short to short and unbranched to sparsely branched (nuda- or hossei-morph); pantropical ... ... Graphis conferta Zenker

    1. A. Ascospores 7–11-septate, 35–45 × 10–13 μm [holotype: Unknown locality, s.col. (not seen; Zenker Reference Zenker, Goebel and Kunze1829)] ... Graphis conferta Zenker

    2. B. Ascospores 7–11-septate, 25–45 × 7–10 μm [holotype: Brazil, Puiggari 337 (G!)] ... ... Graphis virescens Müll. Arg.

       Notes. Lücking et al. (Reference Lücking and Rivas Plata2008) used the name Graphis virescens for this taxon, but G. conferta represents an earlier epithet.

    Lirellae erumpent to prominent, with basal to lateral or complete thalline margin, short to elongate and sparsely to radiately or stellately branched ... 45

46. 45(44) Lirellae with basal to lateral thalline margin ... 46

    Lirellae with apically thin complete thalline margin or labia white-pruinose ... 50

47. 46(45) Disc exposed; lirellae elongate and irregularly branched (handelii-morph); Eastern Palaeotropics [holotype: India, Sethy & Patwardhan 86.820 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis palmicola Makhija & Adaw. [non Graphina palmicola Müll. Arg.]

    Disc concealed; labia variable ... 47

48. 47(46) Lirellae with basal to lateral thalline margin, short and stellately branched (dracaenae- or stellata-morph) ... 48

    Lirellae with (thick) lateral thalline margin, short to elongate and sparsely to irregularly branched ... 49

49. 48(47) Thallus verrucose, olive-green, with scattered isidia; lirellae with lateral thalline margin (stellata-morph); Neotropics and Eastern Palaeotropics (Thailand) [holotype: Brazil, Cáceres & Lücking s.n. (F!; isotype: URM!)] ... ... Graphis stellata Cáceres & Lücking

    Thallus smooth, white-grey, lacking isidia; lirellae with basal thalline margin (geraensis-morph); pantropical [holotype: Africa, Welwitsch 397 (TUR-Vainio 27561!)] ... ... Graphis dracaenae Vain.

    1. A. Ascospores 5–7-septate, 20–30 × 5–7 μm [holotype: Africa, Welwitsch 397 (TUR-Vainio 27561!) ... Graphis dracaenae Vain.

    2. B. Ascospores 5–9-septate, 20–40 × 5–7 μm [holotype: Brazil, Malme 320 (S 6503!)] ... ... Graphis geraensis Redinger

       Notes. The concept of Graphis geraensis provided by Lücking et al. (Reference Lücking and Rivas Plata2008) turned out to be incorrect; the material identified as such from Costa Rica is conspecific with G. conferta, whereas the type of G. geraensis is conspecific with G. dracaenae.

50. 49(47) Lirellae prominent, with thick lateral thalline margin, elongate and irregularly branched (marginata-morph); pantropical ... Graphis oxyclada Müll. Arg.

    1. A. Ascospores 9–11-septate, 30–45 × 5–7 μm [holotype: Kenya, Hildebrandt 2540 (G!)] ... ... Graphis oxyclada Müll. Arg.

    2. B. Ascospores 7–9-septate, 30–35 × 5–7 μm [holotype: Sierra Leone, Deighton M-5539 (FH!)] ... ... Graphis njalensis C. W. Dodge

    Lirellae erumpent, with lateral thalline margin, short to elongate and sparsely to irregularly branched (lineola- or deserpens-morph); pantropical ... ... Graphis immersicans A. W. Archer

    1. A. Ascospores 5–7-septate, 20–30 × 5–8 μm μm [holotype: Philippines, Elmer 14926 (TUR-Vainio 27678!)] ... Graphis leptocarpa var. invita Vain.

    2. B. Ascospores 5–7-septate, 20–30 × 6–8 μm [holotype: Australia, Streimann 31317 (CANB!; isotypes: B!, H!, US!)] ... Graphis immersicans A. W. Archer

       Notes. Graphis immersicans and G. dracaenae are kept separate here due to the stellately branched lirellae with basal thalline margin observed in the latter. More material is necessary to confirm these differences.

51. 50(45) Labia white-pruinose; lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (farinulenta-morph); thallus matt; ascospores 3– 5-septate; Neotropics and African Palaeotropics ... ... Graphis sitiana Vain.

    1. A. Ascospores 3–5-septate, 20–30 × 7–9 μm [holotype: Brazil, Vainio 533 (TUR-Vainio 27854!)] ... Graphis sitiana Vain.

    2. B. Ascospores 3–7-septate, 15–30 × 7–10 μm [holotype: Puerto Rico, Fink 1777 (MICH, not seen, Fink Reference Fink1927)] ... Graphis albida Fink

    Labia non-pruinose; lirellae variable; thallus slightly nitidous ... 51

52. 51(50) Labia very long and radiately branched (anguilliradians-morph); Neotropics [holotype: Brazil, Cáceres & Lücking 00-0655 (F!; isotype: URM!))] ... ... Graphis pernambucoradians Cáceres & Lücking

    Labia short and sparsely branched ... 52

53. 52(51) Lirellae with apically thick complete thalline margin (subserpentina-morph); Neotropics [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Spruce 341 (H-Nylander 7138!)] ... ... Graphis subvirginea Nyl.

    Lirellae with apically thin complete thalline margin (negrosina-morph); Eastern Palaeotropics [holotype: Philippines, Merrill 6790 (TUR-Vainio 27630!)] ... ... Graphis negrosina (Vain.) Lücking*

Group 9: Labia entire, excipulum completely carbonized, hymenium clear, ascospores muriform

1. 1 Ascospores large (80–200 μm long) ... 2

   Ascospores small to medium-sized (15–80 μm long) ... 20

2. 2(1) Norstictic, stictic, salazinic, and/or protocetraric acids (K+ yellow or K+ yellow forming red crystals or P+ red) ... 3

   Notes. If psoromic and subpsoromic acids (P+ yellow) and ascospores I−, cf. Carbacanthographis hillii (A. W. Archer) A. W. Archer; if protocetraric acid (P+ red) and lichexanthone (UV+ yellow) and ascospores I−, cf. Carbacanthographis subalbotecta Staiger.

   No substances (K−, P−) ... 9

3. 3(2) Stictic acid (K+ yellow) ... 4

   Norstictic and/or protocetraric and/or salazinic acids (K+ yellow forming red crystals or P+ red) ... 5

4. 4(3) Ascospores terminally muriform; lirellae prominent, with apically thin complete thalline margin (negrosina-morph); ascospores 11–15 μm broad; Neotropics (Central America) [holotype: Costa Rica, Sipman 48390a (INB!; isotype: B!)] ... ... Graphis subturgidula Lücking & Sipman

   Ascospores regularly muriform; lirellae prominent to sessile, with apically thick complete thalline margin (illinata-morph); ascospores 15–20 μm broad; Eastern Palaeotropics [holotype: Indonesia (Borneo), Hale 30435 (US!)] ... ... Graphis sarawakensis Hale ex Lücking*

   Notes. If ascospores I− and 120–180 × 25–30 μm, thallus ecorticate and lirellae with pruinose cover, cf. Carbacanthographis crassa (Müll. Arg.) Staiger & Kalb; if lichexanthone present (UV+yellow), thallus ecorticate, and lirellae thickly white-pruinose with partially exposed disc, cf. Diorygma confluens (Fée) Kalb, Staiger & Elix and D. epiglaucum (Müll. Arg.) Kalb, Staiger & Elix.

5. 5(3) Protocetraric acid, sometimes additionally norstictic and salazinic acids ... 6

   Norstictic acid only ... 7

6. 6(5) Protocetraric acid only; lirellae prominent, with thick lateral thalline margin, labia black, sharply delimited, short and sparsely branched (marginata-morph); ascospores 20–35 μm broad; Eastern Palaeotropics [holotype: Philippines, Elmer 14583 (TUR-Vainio 27233!)] ... Graphis elmeri Vain.

   Protocetraric, norstictic, and salazinic acids; lirellae sessile, with thick lateral thalline margin, very short and unbranched (dussii-morph); ascospores 15–20 μm broad; Australia [holotype: Australia, Lumbsch 5409c (CANB!)] ... ... Graphis lumbschii (A. W. Archer) A. W. Archer

   Notes. If protocetraric acid (P+ red) and lichexanthone (UV+ yellow), ascospores I−, and lirellae with distinct pruina, cf. Carbacanthographis subalbotecta Staiger; if thallus ecorticate and lirellae thickly white-pruinose with partially exposed disc, cf. Diorygma reniforme (Fée) Kalb, Staiger & Elix.

7. 7(5) Ascospores terminally muriform; lirellae prominent, with apically thick complete thalline margin, elongate and irregularly branched (illinata-morph); Eastern Palaeotropics [holotype: Australia, Streimann 56768 (CANB!)] ... ... Graphis aquilonia (A. W. Archer) Staiger

   Notes. The reports of Graphis aquilonia from Costa Rica (Lücking et al. Reference Lücking and Rivas Plata2008) do not belong to that taxon but represent G. norvestitoides (see Group 18: 3b).

   Ascospores regularly muriform; lirellae erumpent with thick complete thalline margin or sessile with thick lateral thalline margin and then short ... 8

8. 8(7) Lirellae elongate and irregularly branched, erumpent, with thick complete thalline margin (subserpentina-morph); ascospores single, 25–35 μm broad; Eastern Palaeotropics (Australia) [holotype: Australia, Elix 37306 (CANB!)] ... ... Graphis cycasicola A. W. Archer & Elix

   Lirellae very short and unbranched, sessile, with thick lateral thalline margin (dussii-morph); ascospores 2–4 per ascus, 15–25 μm broad; pantropical (also saxicolous) ... Graphis cleistoblephara Nyl.

   1. A. Excipulum completely carbonized; ascospores 80–130 × 15–25 μm [holotype: China (Hong Kong), s.col. (H-Nylander 7589!)] ... Graphis cleistoblephara Nyl.

   2. B. Excipulum (laterally to) completely carbonized; ascospores 75–120 × 15–22 μm [lectotype (here selected): Taiwan, Asahina 300 (W!)] ... Graphina subpulicaris Zahlbr.

9. 9(2) Lirellae round, prominent to sessile, with complete thalline margin (globosa-morph); ascospores single ... 10

   Lirellae at least shortly elongate, otherwise variable; ascospores variable ... 12

10. 10(9) Thallus with abundant, delicate isidia; Neotropics [holotype: Venezuela, Hale 42425 (US!)] ... Graphis isidiata (Hale) Lücking*

    Thallus lacking isidia ... 11

11. 11(10) Ascospores 150–270 × 25–50 μm; Neotropics ... ... Graphis mexicana (Hale) Lücking, Lumbsch & Kalb

    1. A. Ascospores 150–200 × 35–40 μm [holotype: Mexico, Nakanishi 82 (US!)] ... ... Thelotrema mexicanum Hale

    2. B. Ascospores 150–270 × 25–50 μm [holotype: Ecuador, Kalb s.n. (hb. Kalb!)] ... ... Graphina muscicola Kalb

    3. C. Ascospores 160–260 × 30–40 μm [holotype: Brazil, Aptroot 41709 (ABL!)] ... ... Topeliopsis globosa Aptroot

    Ascospores 80–140 × 17–35 μm; Neotropics ... Graphis globosa (Fée) Spreng.

    1. A. Thallus white-grey; ascospores 80–140 × 17–35 μm [lectotype (here selected, based on annotation by Singh in 1983): South America, s.col. (G!)] ... Opegrapha globosa Fée

    2. B. Thallus brown; ascospores 80–130 × 20–30 μm [holotype: Brazil, Vainio s.n. (TUR-Vainio 27168!)] ... Graphis hemisphaerica Vain.

12. 12(9) Lirellae very short and unbranched, prominent to sessile ... 13

    Lirellae distinctly elongate, immersed to prominent ... 15

13. 13(12) Lirellae lacking thalline margin (nuda-morph); Neotropics [holotype: Brazil, Aptroot 41545 (B!)] ... Graphis lourdesina Aptroot*

    Lirellae with thick lateral thalline margin or with thick white cover differing from lateral thalline margin ... 14

14. 14(13) Lirellae with apically thick thalline margin (cleistomma-morph); ascospores single, 120–180 × 30–40 μm; Neotropics (Central America) [holotype: Costa Rica, Lücking 16579 (F!)] ... Graphis oryzaecarpa Lücking

    Lirellae with thick lateral thalline margin but lacking apical cover (dussii-morph); ascospores 1–2 per ascus; 80–120 × 10–20 μm; pantropical ... ... Graphis plagiocarpa Fée

    1. A. Ascospores 80–120 × 10–18 μm [holotype: Caribbean, s.col. (G!)] ... Graphis plagiocarpa Fée

    2. B. Ascospores 95–120 × 15–20 μm [holotype: Australia, Streimann 45427 (CANB!)] ... ... Graphina lumbschii var. deficiens A. W. Archer

15. 15(12) Lirellae immersed, with thick lateral thalline margin (subserpentina-morph); ascospores 2–4/ascus, (40–)80–90 × (16–)20–28 μm (see also 30b); Neotropics and Eastern Palaeotropics (Australia) ... Graphis dolichographa Nyl.

    Lirellae prominent, with basal or complete thalline margin; ascospores variable ... ... 16

16. 16(15) Lirellae with basal thalline margin (hossei-morph); ascospores 80–90 μm long; Neotropics [holotype: America, s.col. (G, not seen; Hale Index Cards)] ... ... Graphis subvernicosa Lücking* [≡ Opegrapha vernicosa Fée, non Graphis vernicosa Nyl.]

    Notes. If ascospores grey-brown and hymenium distinctly inspersed, cf. Thecographa prosiliens (Mont. & Bosch) A. Massal.

    Lirellae with complete thalline margin; ascospores at least partly longer than 90 μm ... 17

17. 17(16) Lirellae very short, unbranched, with apically thin complete thalline margin (cleistomma-morph); Neotropics [lectotype (here selected): Colombia, Lindig 2626 (H-Nylander 6054!)] ... Graphis cleistomma Nyl.

    Notes. There are three syntypes in hb. Nylander, all with the same collection number (Lindig 2626) but with different accession numbers (6054, 7533, 7534). The first two represent parts of the same collection and the well-developed 6054 is here selected as lectotype, although 7533 was annotated by Nakanishi in 1973 (but apparently not published).

    Lirellae distinctly elongate, partly branched, with apically thin or thick thalline margin (rhizocola-or illinata-morph) ... 18

18. 18(17) Ascospores single; lirellae with apically thick thalline margin (illinata-morph); Neotropics [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Martius s.n. (M!)] ... ... Graphis illinata Eschw.

    Ascospores 2–4 per ascus ... 19

19. 19(18) Lirellae with apically thin complete thalline margin (rhizocola-morph); large clusters of calcium oxalate crystals between lateral excipulum and covering thalline margin (see also Group 18: 8a); Neotropics and Eastern Palaeotropics ... ... Graphis acharii Fée

    1. A. Ascospores 80–170 × 15–30 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Glaziou 6286 (BM!; isolectotype: M!)] ... Graphis inturgescens Kremp.

    2. B. Ascospores 100–150 × 20–30 μm [syntypes: Mexico, Pringle 5, 25 (MICH, W, not seen; Wirth & Hale Reference Wirth and Hale1963)] ... Graphina acharii var. subintegra Zahlbr.

    Notes. We first considered Graphis inturgescens as a distinct species, but the type material agrees completely with non-striate, first-generation lirellae of G. acharii.

    Lirellae with apically thick thalline margin (illinata-morph); no crystals between lateral excipulum and covering thalline layer; Neotropics [holotype: Brazil, Vainio s.n. (TUR-Vainio 27171!)] ... Graphis carassensis Vain.

20. 20(1) Norstictic, stictic, and/or protocetraric acids (K+ yellow or K+ yellow forming red crystals or P+ red) ... 21

    Notes. If ascospores I− and salazinic acid present, cf. Carbacanthographis salazinica (A. W. Archer) A. W. Archer.

21. No substances (K−, P−) ... 24

    Notes. If ascospores very small (6–12 μm long), submuriform, I−, cf. Carbacanthographis coccospora (Aptroot) Aptroot & Lücking*.

22. 21(20) Protocetraric acid (P+ red); ascospores very small (15–20 μm long); lirellae prominent, with apically thin complete thalline margin (negrosina-morph); Eastern Palaeotropics [holotype: Solomon Islands, Hill 9367 (BM!)] ... ... Graphis subelmeri (A. W. Archer) A. W. Archer

    Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals); ascospores small to medium-sized (20–50 μm long); lirellae variable ... 22

23. 22(21) Stictic acid (K+ yellow); lirellae prominent, lacking or with basal thalline margin (hossei-morph); Eastern Palaeotropics (China) [holotype: China, Li Jing FJI182 (LHS, not seen; Jia & Wei Reference Jia and Wei2008)] ... Graphis fujianensis Z. F. Jia & J. C. Wei

    Norstictic acid (K+ yellow forming red crystals; lirellae erumpent to prominent, with lateral thalline margin ... 23

    Notes. If salazinic acid, thallus ecorticate, lirellae prominent and white pruinose, and ascospores I−, cf. Carbacanthographis amicta (Nyl.) Staiger & Kalb.

24. 23(22) Lirellae prominent, with apically thick complete thalline margin, very short and unbranched (cleistomma-morph); Eastern Palaeotropics [holotype: Papua New Guinea, Aptroot 38161 (B!)] ... Graphis nadurina Aptroot*

    Lirellae immersed, with lateral thalline margin; pantropical (also saxicolous) [holotype: U.S.A. (Hawaii), Faurie 720 (UPS, not seen; Magnusson Reference Magnusson1955)] ... ... Graphis cremicolor (H. Magn.) Lücking & Archer

25. 24(20) Lirellae prominent to sessile, very short, usually unbranched ... 25

    Lirellae immersed to erumpent, short to elongate, at least partly branched ... 28

    Notes. If disc exposed and brown pruinose, cf. Glyphis atrofusca (Müll. Arg.) Lücking*.

26. 25(24) Lirellae with thick lateral thalline margin (dussii-morph); ascospores 2/ascus, 50–70 × 25–30 μm; Neotropics and Eastern Palaeotropics [holotype: South America, s.col. (H-Acharius 656!)] ... Graphis scaphella Ach.

    Lirellae lacking thalline margin (nuda-morph); ascospores variable ... 26

27. 26(25) Ascospores 55–80 × 20–28 μm, 2–4 per ascus; Neotropics (Central America) [holotype: Costa Rica, Lücking 17750gb (F!)] ... ... Graphis subruiziana Sipman, Chaves & Lücking

    Ascospores 25–65 × 10–20 μm, 4–8 per ascus ... 27

    Notes. If ascospores 20–25 × 8–11 μm, cf. Graphina hartmanniana Müll. Arg., which is not a Graphidaceae; see Ertz & Diederich Reference Ertz and Diederich2007.

28. 27(26) Ascospores 25–40 μm long, 8 per ascus; Neotropics and Eastern Palaeotropics (Hawaii) [holotype: U.S.A. (Hawaii), Faurie 1025b (UPS!)] ... ... Graphis nuda (Magn.) Staiger & Lücking

    Ascospores 35–65 μm long, 4–8 per ascus; cosmopolitan [holotype: South America, s.col. (G!)] ... Graphis ruiziana (Fée) A. Massal.

29. 28(24) Ascospores single, (30–)50–70 × 15–30 μm; lirellae erumpent, with basal thalline margin, elongate and irregularly branched (hossei-morph); Eastern Palaeotropics [holotype: Philippines, Robinson 9083 (TUR-Vainio 27263!)] ... Graphis polillensis Vain.

    Ascospores 2–8 per ascus; lirellae variable ... 29

30. 29(28) Ascospores medium-sized (35–80 μm long) and broad (15–30 μm broad) ... 30

    Ascospores small (20–40 μm long) and narrow (10–12 μm broad) ... 31

31. 30(29) Lirellae erumpent, with lateral thalline margin, very long and radiately branched (centrifuga-morph); ascospores 35–50 × 15–20 μm; Neotropics (Central America) [holotype: Costa Rica, Will-Wolf 12740a (CR!; isotypes: F!, INB!, USJ!, WIS!)] ... ... Graphis gomezii Lücking, Will-Wolf & Umaña

    Lirellae immersed, with thick lateral thalline margin and indistinctly delimited labia (subserpentina-morph); irregularly branched and shorter (up to 5 mm); ascospores 40–80(–90) × 16–28 μm (see also 15a); Neotropics and Eastern Palaeotropics (Australia) [holotype: Colombia, Lindig 866 (H-Nylander 6052!)] ... ... Graphis dolichographa Nyl.

32. 31(29) Lirellae erumpent, with apically thin complete thalline margin (upper part of labia dark grey), elongate and irregularly branched (negrosina-morph); Neotropics [holotype: Colombia, Lindig 2725 (H-Nylander 7408b!)] ... Graphis subtecta (Nyl.) Lücking*

    Lirellae with basal to lateral thalline margin or thalline margin lacking, upper part of labia black ... 32

33. 32(31) Lirellae very long (up to 10 mm), radiately branched (centrifuga-morph); Neotropics [holotype: Brazil, Puiggari s.n. (G!)] ... Graphis jeanmuelleri Lücking*[≡ Graphina elegantula Müll. Arg., non Graphis elegantula Zahlbr.]

    Lirellae rather short (1–3 mm), unbranched or sparsely branched ... 33

34. 33(32) Ascospores 20–30 μm long; lirellae erumpent, with basal thalline margin, short and sparsely branched (subregularis-morph); African Palaeotropics [lectotype (here selected, lower middle piece extracted for TLC by Staiger in 1998): Guinea, s.col. (H-Acharius 628!)] ... Graphis comma (Ach.) Spreng. [non Eschw.]

    Notes. The species was reported from America by Fée (Reference Fée1824) but we could not verify that record.

    Ascospores 30–40 μm long; lirellae with lateral thalline margin (lineola-morph); Eastern Palaeotropics [holotype: Solomon Islands, Hill 9821 (BM!)] ... ... Graphis maritima (A. W. Archer) A. W. Archer

Group 10: Labia entire, excipulum completely carbonized, hymenium inspersed, ascospores transversely septate

1. 1 Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   No substances ... 9

2. 2(1) Stictic acid (K+ yellow); lirellae erumpent to prominent, with apically thin or thick complete thalline margin ... 3

   Norstictic acid (K+ yellow forming red crystals); lirellae erumpent, with (thick) lateral thalline margin or thalline margin absent ... 5

3. 3(2) Ascospores 5–7-septate, 25–35 × 6–8 μm; lirellae erumpent, with apically thin complete thalline margin (negrosina-morph); thallus verrucose; Eastern Palaeotropics [type: Solomon Islands (BM!)] ... Graphis luluensis A. W. Archer

   Ascospores 9–19-septate, 50–90 × 9–16 μm; lirellae variable ... 4

4. 4(3) Lirellae prominent, short and sparsely branched, with apically thick complete thalline margin (illinata-morph); thallus uneven; Eastern Palaeotropics (Hawaii) [holotype: U.S.A. (Hawaii), Rock 131 (W!; isotype: FH!)] ... ... Graphis apoda (Zahlbr.) Lücking*

   Lirellae erumpent to prominent, elongate and irregularly branched, with thick lateral thalline margin (marginata-morph); thallus smooth; Eastern Palaeotropics (Australia) [holotype: Australia, Rogers 7613 (BRI!)] ... ... Graphis gloriosensis A. W. Archer & Elix

5. 5(2) Disc exposed, white-pruinose, lirellae erumpent, with lateral thalline margin, short and sparsely branched (scripta-morph); ascospores 20–35 μm long; Palaeotropics ... Graphis aperiens Müll. Arg.

   1. A. Ascospores 5–9-septate, 20–30 × 6–9 μm [holotype: Japan, Miyoshi s.n. (G!)] ... ... Graphis aperiens Müll. Arg.

   2. B. Ascospores 5–7-septate, 20–35 × 6–9 μm [holotype: Australia, Bailey 488 (G!)] ... ... Graphis semiaperta Müll. Arg.

      Notes. Archer (Reference Archer2006) treated Graphis semiaperta a separate species but G. aperiens is an older name available for this taxon.

   Disc concealed, labia non-pruinose; ascospores variable ... 6

6. 6(5) Lirellae erumpent, lacking thalline margin, short to elongate and irregularly branched (hossei-morph); ascospores small to medium-sized (25–50 μm long); Neotropics and Eastern Palaeotropics [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Glaziou 5082 (M!)] ... Graphis desquamescens (Fée) Zahlbr.[≡ Graphis compulsa Kremp. nom. illeg.]

   Notes. Krempelhuber (Reference Krempelhuber1876) deliberately replaced the epithet desquamescens with compulsa, which is illegitimate, but the name change was accepted by Redinger (Reference Redinger1935).

   Lirellae erumpent, with lateral thalline margin; ascospores (very) small (15–30 μm long) ... 7

7. 7(6) Lirellae very long and radiately branched (centrifuga-morph); ascospores very small (15–25 μm long); Eastern Palaeotropics [holotype: Australia, Wilson s.n. (H-Räsänen!)] ... ... Graphis centrifuga Räsänen

   Lirellae short and stellately branched or short to elongate and sparsely to irregularly or branched; ascospores small (20–30 μm long) ... 8

8. 8(7) Lirellae short, stellately branched (coarctata-morph); Eastern Palaeotropics [holotype: Taiwan, Faurie 236 (W!)] ... Graphis cervinonigra Zahlbr.

   Lirellae short to elongate, sparsely to irregularly branched (lineola- or deserpens- morph); Palaeotropics ... Graphis gonimica Zahlbr.

   1. A. Lirellae short, sparsely branched [lectotype (here selected, based on annotation label by Nakanishi in 1973): China, Chung 591a (W!)] ... Graphis gonimica Zahlbr.

   2. B. Lirellae elongate, irregularly branched [lectotype (here selected, based on annotation label by Nakanishi in 1973): China, Handel-Mazzetti 12819 (W!; isolectotype: S 2169!)] ... ... Graphis sapii Zahlbr.

   3. C. Lirellae short, sparsely branched [holotype: Australia, Stevens s.n. (BRI!)] ... ... Graphis eimeoensis A. W. Archer & Elix

      Notes. The type material of Graphis sapii also contains G. pinicola and G. tsunodae.

9. 9(1) Ascospores large (70–135 × 9–30 μm) ... 10

   Ascospores small to medium-sized (25–70 × 6–9 μm) ... 12

10. 10(9) Lirellae erumpent, with complete thalline margin (subserpentina-morph); hymenial inspersion of type A; African Palaeotropics [holotype: Tanzania, Holst s.n. (G!)] ... ... Graphis superans Müll. Arg.

    Lirellae prominent, with apically thin complete thalline margin (negrosina-morph); hymenial inspersion of type B ... 11

11. 11(10) Ascospores 11–15 μm broad; Neotropics (Central America) [holotype: Costa Rica, Lücking 15449a (INB!)] ... Graphis bettinae Lücking, Umana, Chaves & Sipman

    Ascospores 20–30 μm broad; South America [holotype: Brazil, Kalb 31118 (hb. Kalb; Staiger Reference Staiger2002)] ... Graphis subserpens Staiger*

12. 12(9) Ascospores medium-sized (60–70 μm long), 13–17-septate; lirellae prominent, with basal thalline margin (hossei-morph); Neotropics [holotype: Cuba, Ekman 19 (TUR- Vainio 27371!)] ... Graphis cupei Vain. ex Lücking*

    Ascospores small (25–45 μm long), 7–11-septate; lirellae variable ... 13

13. 13(12) Lirellae prominent, lacking thalline margin, short and sparsely branched (hossei-morph); possibly pantropical [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Martius s. n. (M!; isolectotype: G!)] ... Graphis anfractuosa (Eschw.) Eschw.

    Lirellae erumpent, with lateral thalline margin, very long and radiately branched (centrifuga-morph); Palaeotropics [holotype: Thailand, Schmidt s. n. (TUR-Vainio 27563!)] ... Graphis arbusculaeformis (Vain.) Lücking*

Group 11: Labia entire, excipulum completely carbonized, hymenium inspersed, ascospores muriform

1. 1 Ascospores terminally muriform; lirellae prominent, with apically thin complete thalline margin (upper part of labia dark grey), elongate and irregularly branched (negrosina-morph) (see also Group 20: 1a); Neotropics (Central America) ... ... Graphis subflexibilis Lücking & Chaves

   Ascospores regularly muriform ... 2

2. 2(1) Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 3

   Notes. If protocetraric acid and ascospores I−, 14–18 × 5–7 μm, cf. Carbacanthographis inspersa Staiger.

   Hirtifructic acid or no substances (K−) ... 6

3. 3(2) Stictic acid (K+ yellow); lirellae prominent, with apically thin complete thalline margin (negrosina-morph); Neotropics (Central America) [holotype: Costa Rica, Sipman 48189 (INB!; isotype: B!)] ... Graphis inspersostictica Sipman & Lücking

   Norstictic acid (K+ yellow forming red crystals); lirellae erumpent to prominent, with thick lateral thalline margin ... 4

4. 4(3) Ascospores single, 100–150 × 25–40 μm; lirellae erumpent, with thick lateral thalline margin, elongate and irregularly branched (subserpentina-morph); pantropical [holotype: Australia, Sayer s.n. (G!)] ... ... Graphis novopalmicola A. W. Archer & Lücking* [≡ Graphina palmicola Müll. Arg., non Graphis palmicola Makhija & Adaw.]

   Ascospores 1–4 per ascus, 40–110 × 15–30 μm ... 5

5. 5(4) Ascospores 1–2 per ascus, 60–110 μm long; lirellae prominent, with thick lateral thalline margin, very short and unbranched (dussii-morph); pantropical [lectotype (Staiger Reference Staiger2002): Sri Lanka, Almquist 1879 (H-Nylander 7398!)] ... Graphis leprographa Nyl.

   Ascospores (2–)4 per ascus, 40–55 μm long; lirellae erumpent, with thick lateral thalline margin, short and sparsely branched (subserpentina-morph); Eastern Palaeotropics (Australia) [holotype: Australia, Hartmann s.n. (G!)] ... ... Graphis saxicola (Müll. Arg.) A. W. Archer

6. 6(2) Ascospores single; lirellae with apically thick complete thalline margin (illinata-morph); hirtifructic acid; Australia [holotype: Australia, Elix 16321 (CANB!)] ... ... Graphis elixiana A. W. Archer [≡ Phaeographis elixii A. W. Archer, non Graphis elixii A. W. Archer]

   Ascospores (1–)2–8 per ascus; lirellae with apically thin complete thalline margin (negrosina-morph); no substances ... 7

7. 7(6) Ascospores 15–25 μm broad, 4–8 per ascus (see also Group 20: 5) ... ... Graphis argentata Lücking & Umaña

   Ascospores 25–35 μm broad, 1–4 per ascus (see also Group 20: 5); Neotropics [holotype: Brazil, Vainio s.n. (TUR-Vainio 27166!)] ... Graphis phaeospora Vain.

Group 12: Labia striate, excipulum apically and basally carbonized, interrupted by lateral non-carbonized parts, hymenium clear, ascospores transversely septate to muriform

1. 1 Ascospores distinctly muriform, single, 150–180 × 30–45 μm (often grey-brown when old but I+ violet); lirellae prominent, with apically thin complete thalline margin (upper part of labia dark grey); elongate and irregularly branched (acharii- morph); Neotropics (Central America) [holotype: Costa Rica, Sipman 48208 (INB!; isotype: B!)] ... Graphis mirabilis Lücking, Sipman, Umaña & Chaves

   Ascospores transversely septate or with occasional longitudinal septa in some of the segments, 2–8 per ascus, 45–110 × 10–14 μm; lirellae immersed to erumpent, with apically thin complete thalline margin (symplecta-morph) ... 2

2. 2(1) Ascospores transversely septate, 45–80 μm long; lirellae erumpent; Neotropics (Central America) and Asian Palaeotropics [holotype: Costa Rica, Sipman 51764 (INB!; isotype: B!)] ... Graphis gregmuelleri Sipman & Lücking

   Ascospores with a few longitudinal septa, 60–110 μm long; lirellae immersed; Neotropics (South America) [holotype: Paraguay, Balansa s.n. (G!)] ... ... Graphis immersoides Lücking [≡ Graphina immersa Müll. Arg., non Graphis immersa Fink]

Group 13: Labia striate, excipulum apically (to peripherally) carbonized, hymenium clear, ascospores transversely septate

1. 1 Norstictic and/or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   No substances (K−) ... 10

2. 2(1) Norstictic acid, sometimes additionally stictic acid (K+ yellow forming red crystals) ... 3

   Stictic acid only (K+ yellow) ... 7

3. 3(2) Ascospores large (75–100 μm long) ... 4

   Ascospores small to medium-sized (20–60 μm long) ... 5

4. 4(3) Norstictic acid only; lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Eastern Palaeotropics [holotype: Thailand, Hosseus s.n. (TUR-Vainio 27801!)] ... Graphis trichospora Vain.

   Notes. Graphis trichospora, G. verminosa, and G. leptospora share the same lirellae morphology, ascospore type and chemistry, but differ in the degree of excipulum carbonization (apical versus lateral versus complete). As in other cases, and contrary to other authors, we consider these to represent distinct species, but the available material is sparse and the variation in excipulum carbonization cannot be properly assessed at this point.

   Norstictic and stictic acids; lirellae erumpent, with apically thin complete thalline margin (upper part of labia dark grey), elongate and irregularly branched (symplecta-morph); Eastern Palaeotropics [holotype: India, Patwardhan & Nagarkar 77.1115 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis meghalayaensis Adaw. & Makhija

5. 5(3) Lirellae immersed, with lateral thalline margin; labia white-pruinose (glaucescens-morph); norstictic acid only; Palaeotropics ... Graphis exalbata Nyl.

   1. A. Excipulum apically (to laterally) carbonized; ascospores 7–11-septate, 25–45 × 6–9 μm [lectotype (based on annotation label by Awasthi & Mathur 1989): São Tomé and Principe, Fr. Quintas s. n. (H-Nylander 7661!)] ... Graphis exalbata Nyl.

   2. B. Excipulum apically carbonized; ascospores 7–11-septate, 20–50 × 6 μm [holotype: India (AMH, not seen; Makhija et al. Reference Makhija, Dube, Adawadkar and Chitale2006; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis nerurensis Makhija, A. Dube, Adaw. & Chitale

6. Lirellae erumpent to prominent, with basal thalline margin or lacking thalline margin; labia non-pruinose; norstictic, stictic, and constictic acids ... 6

7. 6(5) Lirellae erumpent, lacking thalline margin, elongate and irregularly branched (striatula-morph); ascospores 30–60 μm long; Eastern Palaeotropics [lectotype (Awashti & Singh Reference Awasthi and Singh1975): India, Thwaites 82 (BM, not seen; Awasthi & Singh Reference Awasthi and Singh1975)] ... ... Graphis nigroglauca Leight.

   Notes. This species has also been reported as having an inspersed hymenium, but we have not been able to study the type and confirm this feature; inspersed hymenia are extremely rare in species with striatula-morph lirellae.

   Lirellae prominent, with basal thalline margin, short and sparsely branched (striatula-morph); ascospores 25–35 μm long; Eastern Palaeotropics [holotype: India, Nagarkar & Sethy 81.508 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis subvittata Adaw. & Makhija

8. 7(2) Ascospores medium-sized to large (60–160 × 12–16 μm), 2–4 per ascus; labia pruinose; lirellae erumpent, lacking thalline margin, elongate and irregularly branched (albotecta-morph); Eastern Palaeotropics ... Graphis sp. 1 in Awasthi (Reference Awasthi1991)

   Ascospores small to medium-sized (20–60 × 6–10 μm), 8/ascus; labia non-pruinose; lirellae variable ... 8

9. 8(7) Thallus isidiate; lirellae erumpent, with complete thalline margin; ascospores 20–35 μm long; Eastern Palaeotropics [India, Makhija 01.106 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2004)] ... Graphis isidiza Adaw. & Makhija

   Thallis lacking isidia; lirellae variable; ascospores variable ... 9

10. 9(8) Lirellae erumpent, with lateral thalline margin; ascospores 30–50 μm long; pantropical ... Graphis vittata Müll. Arg.

    1. A. Ascospores 7–11-septate, 30–50 × 7–8 μm [holotype: Indonesia, Junghuhn 111 (Java) (G!)] ... ... Graphis vittata Müll. Arg.

    2. B. Ascospores 7–11-septate, 30–40 × 7–10 μm [holotype: Indonesia (Java), Van Overeem 93 (W!)] ... Graphis treubii Zahlbr.

    3. C. Ascospores 7–11-septate, 30–40 × 7–10 μm [holotype: China, Handel-Mazzetti 11217 (W!;isotype: US!)] ... Graphis theae Zahlbr.

    4. D. Ascospores 7–9-septate, 30–35 × 6–8 μm [holotype: Taiwan, Asahina 368 (W!)] ... ... Graphis flabellans Zahlbr.

       Notes. Archer (Reference Archer2006) maintained Graphis treubii separate from G. vittata but reexamination of the types confirmed conspecificity of the two taxa.

    Lirellae erumpent to prominent, lacking thalline margin, short and sparsely branched (striatula-morph); ascospores 20–40 μm long; pantropical [lectotype (Archer Reference Archer2006): Philippines, Fénix 12786 (TUR-Vainio 27887!)] ... Graphis stenotera Vain.

11. 10(1) Ascospores medium-sized to large (50–125 μm long) ... 11

    Ascospores small (20–50 μm long) ... 15

12. 11(10) Thallus isidiate; lirellae erumpent, with lateral thalline margin, very long and radiately branched (dichotoma-morph); ascospores 9–19-septate, 60–110 × 8–12 μm; Eastern Palaeotropics [holotype: India, Prabhu & Nagarkar 74.3534 (AMH, photograph seen; isotype: US!)] ... Graphis patwardhanii C. R. Kulk.

    Thallus lacking isidia; lirellae variable; ascospores variable ... 12

13. 12(11) Ascospores large (80–125 μm long) ... 13

    Ascospores medium-sized (50–70 μm long) ... 14

14. 13(12) Thallus white-grey; lirellae erumpent, with lateral thalline margin (tenella-morph) and distinct white lines inbetween striae, formed by clusters of calcium-oxalate crystals; Neotropics [holotype: Guadeloupe, Duss 517 (TUR-Vainio 27847!)] ... ... Graphis subalbostriata Lücking* [≡ G. angustata var. albostriata Vain., non G. albostriata Vain.]

    Thallus dark olive-grey; lirellae erumpent, with apically thin complete thalline margin, elongate and irregularly branched (symplecta-morph); pantropical [holotype: Brazil, Malme 2267B (S-2170!)] ... Graphis olivacea Redinger

15. 14(12) Thallus yellow-green; labia non-pruinose; lirellae prominent, lacking thalline margin, elongate and irregularly branched (striatula-morph); Neotropics (Central America, Caribbean) [holotype: Dominica, Imshaug & Imshaug 32777-A2 (MSC!)] ... ... Graphis caribica Lücking

    Thallus white-grey; labia white-pruinose; lirellae immersed, with lateral thalline margin, elongate and irregularly branched (glaucescens-morph); Neotropics [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Costa Rica, Tonduz s.n. (G!)] ... ... Graphis supertecta Müll. Arg.

16. 15(10) Labia white-pruinose; thallus ecorticate or corticate ... 16

17. Labia non-pruinose; thallus corticate ... 17

18. 16(15) Thallus ecorticate (glaucescens-morph); ascospores 25–35 μm long (see also Group 2: 9); pantropical ... Graphis glaucescens Fée

    1. A. Labia striate; ascospores 7–9-septate, 25–35 × 5–7 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): South America, s.col. (G!; isolectotype: S 2184!)] ... Graphis glaucescens Fée

    2. B. Labia striate; ascospores 5–9-septate, 25–35 × 6–8 μm [holotype: Paraguay, Malme 1526 (S-3947!; isotype: S-3948!)] ... Graphis caesioglauca Redinger

19. Thallus corticate (chloroalba-morph); ascospores 20–25 μm long; Eastern Palaeotropics [holotype: India, Nagarkar & Patwardhan 86.529 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... Graphis chloroalba Makhija & Adaw.

20. 17(15) Lirellae prominent to sessile, lacking thalline margin or very rarely with basal thalline margin ... 18

    Lirellae erumpent, with basal to lateral thalline margin ... 20

21. 18(17) Lirellae very short, usually unbranched (granulocarpa-morph); Neotropics [lectotype (here selected): Brazil, Malme 3680 (S-6502!)] ... Graphis granulocarpa Redinger

    Lirellae short to elongate, sparsely to irregularly branched (striatula-morph) ... 19

22. 19(18) Lirellae sessile, resembling Glyphis substriatula; thallus yellow-olive; ascospores 20–30 μm long; pantropical ... Graphis endoxantha Nyl.

    1. A. Ascospores 5–7-septate, 20–30 × 6–8 μm [holotype: New Caledonia, Pancher s.n. (H-Nylander 7782!)] ... Graphis endoxantha Nyl.

    2. B. Ascospores 5–7-septate, 20–30 × 6–10 μm [holotype: Mexico, Pringle 162 (H-Nylander 7776!)] ... Graphis subelegans Nyl.

23. Lirellae prominent; thallus white-grey; ascospores 20–45 μm long; pantropical ... Graphis tsunodae Zahlbr.

    1. A. Ascospores 7–11-septate, 20–40 × 6–9 μm [holotype: Japan, Tsunoda 182 (W!)] ... ... Graphis tsunodae Zahlbr.

    2. B. Ascospores 5–13-septate, 20–45 × 7–9 μm [holotype: China, s.col. (W!; isotype: S 3950!)] ... ... Graphis rockii Redinger

       Notes. Lücking et al. (Reference Lücking and Rivas Plata2008) treated this taxon as G. rockii but G. tsunodae is an earlier epithet. The latter was listed as synonym of G. proserpens by Nakanishi (Reference Nakanishi1966) and Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b), but differs morphologically from that species in the shorter, much less branched, thicker and prominent lirellae, thus agreeing with G. rockii.

24. 20(17) Lirellae with lateral thalline margin, short to elongate and sparsely to irregularly branched (tenella-morph); pantropical ... Graphis chlorotica A. Massal.

    1. A. Lirellae erumpent, with lateral thalline margin, short and sparsely branched; ascospores 9–13-septate, 30–45 × 5–8 μm [holotype: Indonesia (Java), s.col. (W, not seen; Hale Index Cards)] ... Graphis chlorotica A. Massal.

    2. B. Lirellae erumpent, with lateral thalline margin, elongate and almost radiately branched; ascospores 7–9-septate, 20–35 × 5–8 μm long [holotype: Australia, Sayer s.n. (G!)] ... ... Graphis subtenella Müll. Arg.

    3. C. Lirellae erumpent (to prominent), with lateral (to basal) thalline margin, short to elongate and sparsely branched; ascospores 9–13-septate, 30–50 × 7–10 μm [holotype: Philippines (TUR-Vainio 27628!)] ... Graphis duplicata var. australasiatica Vain.

25. Lirellae lacking or with basal thalline margin, elongate and irregularly branched (striatula-morph) ... 21

26. 21(20) Thallus verrucose; Neotropics [holotype: Jamaica, Imshaug 14579 (MSC-0029035!)] ... ... Graphis verrucoserpens Lücking

    Thallus smooth to uneven; possibly pantropical ... Graphis proserpens Vain.

    1. A. Ascospores 7–11-septate, 20–40 × 6–9 μm [holotype: Brazil, Vainio s. n. (TUR-Vainio 27557A!)] ... Graphis proserpens Vain. [≡ Graphis disserpens Vain., nom. illeg., non Nyl.]

    2. B. Ascospores 7–9-septate, 25–45 × 6–8 μm [holotype: India, Patwardhan & Nagarkar 77.1806 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis sikkimensis Nagarkar & Patw.

       Notes. According to Nakanishi (Reference Nakanishi1966) and Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003b), Graphis endoxantha Asahina [nom. illeg., non Nyl.] is also a synonym of G. proserpens, but since their concept of the latter species is rather broad and we have not seen type material, the current status of the former could not be ascertained.

Group 14: Labia striate, excipulum apically (to peripherally) carbonized, hymenium clear, ascospores muriform

1. 1 Norstictic and/or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   No substances (K−) ... 8

2. 2(1) Norstictic acid (K+ yellow forming red crystals); ascospores single or 2–8 per ascus ... 3

   Stictic acid (K+ yellow) ... 6

3. 3(2) Ascospores single, medium-sized to large (60–125 μm long) ... 4

   Ascospores 2–8 per ascus, small to medium-sized (20–60 μm long) ... 5

4. 4(3) Lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Eastern Palaeotropics [isotype: India, Watt 83 (G!)] ... Graphis semirigida (Müll. Arg.) Lücking*

   Lirellae erumpent, with apically thick complete thalline margin, elongate and irregularly branched (consanguinea-morph); Neotropics ... ... Graphis bipartita (Müll. Arg.) Lücking

   1. A. Ascospores 60–100 × 25–40 μm [lectotype (Wirth & Hale Reference Wirth and Hale1963): Paraguay, Balansa 1878 (G!)] ... Graphina bipartita Müll. Arg.

   2. B. Ascospores 60–110 × 20–30 μm [holotype: Mexico, Pringle 222 (H!; isotype: FH!)] ... ... Graphis peralbida Nyl.

5. 5(3) Ascospores 30–60 × 17–25 μm; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); pantropical ... ... Graphis antillarum Vain.

   1. A. Lirellae with lateral thalline margin; ascospores 30–60 × 17–25 μm [holotype: Guadeloupe, Duss 540 (TUR-Vainio 27182!; isotype: FH!)] ... Graphis antillarum Vain.

   2. B. Lirellae with lateral thalline margin; ascospores 35–55 × 18–25 μm [holotype: Guadeloupe,Duss 1198 (TUR-Vainio 27178!)] ... Graphis platycarpoides Vain.

   Ascospores 20–45 × 10–17 μm; lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); pantropical ... ... Graphis perstriatula Nyl.

   1. A. Lirellae with basal thalline margin; ascospores 20–40 × 11–15 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): India, Kurz s.n. (H-Nylander 7779!)] ... Graphis perstriatula Nyl.

   2. B. Lirellae with lateral thalline margin; ascospores 35–45 × 13–17 μm [holotype: Lesser Antilles, Raunkiaer 437 (TUR-Vainio 27243!)] ... Graphis acuminata Vain.

   3. C. Lirellae with basal thalline margin; ascospores 20–40 × 11–15 μm [holotype: Mariana Islands (Guam), McGregor 604 (TUR-Vainio 27238!)] ... Graphis gracilescens Vain.

   4. D. Lirellae with lateral thalline margin; ascospores 20–35 × 10–17 μm [holotype: Puerto Rico, Fink 659 (MICH, not seen; isotype: NY!)] ... Graphina sulcata Fink

   5. E. Lirellae with basal thalline margin; ascospores 20–35 × 8–13 μm [original material: Cuba, Ekman 5 (TUR-Vainio 27169!)] ... Graphis polyschizans Vain. [nom. inval. (not validly published)]

6. 6(2) Ascospore medium-sized (45–70 × 16–22 μm); lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); Neotropics ... Graphis parilis Kremp.

   1. A. Ascospores 45–70 × 16–20 μm [holotype: Brazil, Glaziou 3394 (G!)] ... Graphis parilis Kremp.

   2. B. Ascospores 50–70 × 18–20 μm [holotype: Brazil, Glaziou 3388 (M, not seen; Krempelhuber Reference Krempelhuber1876)] ... Graphis annulata Kremp.

   3. C. Ascospores 45–55 × 18–22 μm [holotype: Costa Rica, Tonduz s.n. (G!)] ... ... Graphina acromelaena Müll. Arg.

   4. D. Ascospores 45–65 × 16–20 μm [holotype: U.S.A. (Louisiana), Eckfeldt 751 (G!)] ... ... Graphina acrophaea Müll. Arg.

7. Ascospores small (20–45 × 9–16 μm); lirellae variable ... 7

8. 7(6) Lirellae erumpent, with apically thin complete thalline margin, elongate and irregularly branched (symplecta-morph); Eastern Palaeotropics [holotype: Philippines, Merrill 8576 (TUR-Vainio 27237!)] ... Graphis subducta Vain.

   Lirellae erumpent to prominent, with lateral thalline margin, short and sparsely branched (lineola-morph); Eastern Palaeotropics [holotype: China, Handel-Mazzetti 5912 (W, not seen; Zahlbruckner Reference Zahlbruckner and Handel-Mazetti1930; Hale Index Cards)] ... ... Graphis galactoderma (Zahlbr.) Lücking*

9. 8(1) Ascospores medium-sized to large (50–100 μm long) ... 9

   Ascospores small (25–45 μm long) ... 10

10. 9(8) Ascospores large (70–100 × 20–30 μm), 1–2 per ascus; lirellae erumpent, with lateral thalline margin (tenella-morph); Neotropics (Central America) [holotype: Costa Rica, Lücking 17265aa (USJ!; isotype: F!)] ... Graphis tenoriensis Lücking & Chaves

    Ascospores medium-sized (50–70 × 14–20 μm), 8 per ascus; lirellae erumpent, with apically thin complete thalline margin (symplecta-morph); Neotropics ... ... Graphis platycarpa Eschw.

    1. A. Ascospores 55–70 × 15–20 μm [holotype: Brazil, Martius s.n. (M!)] Graphis platycarpa Eschw.

    2. B. Ascospores 50–70 × 14–20 μm [lectotype (Archer Reference Archer2001d): Colombia, Lindig 2726 (H-Nylander 7379!)] ... Graphis sophistica Nyl.

11. 10(8) Ascospores 15–25 μm broad ... 11

    Ascospores 9–15 μm broad ... 12

12. 11(10) Lirellae immersed to erumpent, with apically thin complete thalline margin, short to elongate and sparsely to irregularly branched (symplecta-morph); ascospores 6–8 per ascus; Neotropics [holotype: Colombia, Lindig s.n. (H-Nylander 7404!)] ... ... Graphis symplecta Nyl.

    Lirellae erumpent, with lateral thalline margin, very long and radiately branched (dichotoma-morph); ascospores 1–2 per ascus; Neotropics [holotype: Colombia, Lindig 93 (H-Nylander 7415!)] ... Graphis disserpens Nyl.

13. 12(10) Lirellae erumpent, with apically thin complete thalline margin, elongate and irregularly branched (symplecta-morph); Neotropics (Central America) [holotype: Costa Rica, Lücking 15667 (USJ!; isotype: F!)] ... Graphis paraserpens Lizano & Lücking

    Lirellae erumpent, lacking or with basal thalline margin, very long and radiately branched (sorediosa-morph); Neotropics (Central America) [holotype: Costa Rica, Sipman 47989f (INB!; isotype: B!)] ... Graphis paradisserpens Sipman & Lücking

Group 15: Labia striate, excipulum laterally carbonized, hymenium clear, ascospores transversely septate

1. 1 Norstictic or stictic acid or lichexanthone (K+ yellow or K+ yellow forming red crystals or UV+ yellow) ... 2

   No substances (K−) ... 8

2. 2(1) Lichexanthone (UV+ yellow, K−); lirellae prominent, lacking thalline margin, short to elongate and sparsely to irregularly branched (striatula-morph) ... 3

   Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals, UV–); lirellae variable ... 4

3. 3(2) Ascospores medium-sized (50–85 × 7–12 μm), 9–19-septate; Neotropics ... ... Graphis haleana R. C. Harris

   1. A. Ascospores 9–15-septate, 50–75 × 9–12 μm [holotype: U.S.A. (Florida), Harris 25248 (NY!)] ... ... Graphis haleana R. C. Harris

   2. B. Ascospores 11–19-septate, 60–85 × 7–12 μm [original material: Brazil, Kalb 28244 (hb. Kalb, not seen; Staiger Reference Staiger2002)] ... Graphis cuiabensis Staiger [nom. inval. (not validly published)]

4. Ascospores small (20–40 × 7–10 μm), 5–9-septate; Neotropics ... ... Graphis lucifica R. C. Harris

   1. A. Ascospores 5–7-septate, 20–30 × 7–9 μm [holotype: USA (Florida), Harris 25174 (NY!)] ... ... Graphis lucifica R. C. Harris,

   2. B. Ascospores 7–9-septate, 30–40 × 8–10 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Malme 3605 (S 6490!)] ... Graphis rimulosa var. tetraspora Redinger

5. 4(2) Norstictic acid (K+ yellow forming red crystals); ascospores (small to) medium-sized to large (35–150 μm long) ... 5

   Stictic acid (K+ yellow); ascospores small (20–35 μm long) ... 7

6. 5(4) Ascospores (small to) medium-sized (35–60 μm long), 7–13-septate, conspicuously thick-walled and often somewhat greyish; lirellae erumpent to prominent, lacking thalline margin, short and sparsely branched (striatula-morph); subcosmopolitan ... Graphis elegans (Sm.) Ach.

   1. A. Ascospores 9–13-septate, 35–60 × 9–13 μm [holotype: Great Britain (England), Borrer s.n. (BM; Staiger Reference Staiger2002)] ... Graphis elegans (Sm.) Ach.

   2. B. Ascospores 7–9-septate, 35–60 × 11–13 μm [holotype: America, s.col. (G, not seen; Hale Index Cards)] ... Opegrapha rhabdotis Fée

      Notes. Graphis ramificans Nyl. keys out here but we have been unable to locate type material and confirm the described characters of this taxon.

7. Ascospores medium-sized to large (60–150 μm long), 15–35-septate, not conspicuously thick-walled and always hyaline; lirellae erumpent to prominent, with lateral to apically thin complete thalline margin, elongate and irregularly branched ... 6

8. 6(5) Lirellae prominent, with lateral thalline margin, elongate and irregularly branched (celata-morph); Eastern Palaeotropics [holotype: India, s.col. (BM, not seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... Graphis celata Stirt.

   Lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Eastern Palaeotropics [lectotype (Awasthi & Singh Reference Awasthi and Singh1975): India, Watt s.n. (BM, not seen; paratype: Watt 76, G!)] ... ... Graphis verminosa Müll. Arg.

   Notes. Graphis trichospora, G. verminosa, and G. leptospora share the same lirella morphology, ascospore type, and chemistry, but differ in the degree of excipulum carbonization (apical versus lateral versus complete). As in other cases, and contrary to other authors, we consider these to represent distinct species, but the available material is sparse and the variation in excipulum carbonization cannot be properly assessed at this point.

9. 7(4) Thallus sorediate; lirellae erumpent, lacking thalline margin, very long and radiately branched (sorediosa-morph); Eastern Palaeotropics [holotype: India, Nagarkar 78.364 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2007)] ... ... Graphis sorediosa Nagarkar & Patw.

   Thallus lacking soralia; lirellae erumpent, with basal thalline margin, elongate and irregularly branched (striatula-morph); Neotropics (SE USA) and Eastern Palaeotropics [holotype: Papua New Guinea, Aptroot 31685 (B!)] ... ... Graphis brahmanensis Aptroot*

   Notes. This taxon appears to be conspecific with Graphis sp. 25050 in Harris (Reference Harris1995).

10. 8(1) Ascospores medium-sized to large (70–190 μm long); lirellae lacking thalline margin (striatula-morph) ... 9

    Ascospores small to medium-sized (15–70 μm long); lirellae variable ... 10

11. 9(8) Lirellae strongly sessile (see also Group 17: 12); Eastern Palaeotropics [lectotype (Awashti & Singh Reference Awasthi and Singh1975): India, Watt 6399 (G, not seen; Awasthi & Singh Reference Awasthi and Singh1975; Hale Index Cards)] ... Graphis contortuplicata Müll. Arg.

    Lirellae erumpent; Eastern Palaeotropics [holotype: India (AMH; photograph seen; Makhija et al. Reference Makhija, Dube, Adawadkar and Chitale2006)] ... Graphis polystriata Makhija & Dube

12. 10(8) Lirellae erumpent to prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Neotropics [holotype: Guadeloupe, Duss 1190 (TUR-Vainio 27848!)] ... Graphis ingarum (Vain.) Lücking*

    Lirellae erumpent to prominent, lacking thalline margin or with basal to lateral thalline margin ... 11

13. 11(10) Ascospores small to medium-sized (30–65 × 7–12 μm; large ascospores exceeding 50 μm ... 12

    Ascospores small (15–45 × 6–9 μm) ... 13

14. 12(11) Lirellae prominent, lacking thalline margin or with basal thalline margin, elongate and irregularly branched (striatula-morph); pantropical ... ... Graphis striatula (Ach.) Spreng.

    1. A. Excipulum laterally (to almost completely) carbonized; lirellae lacking thalline margin, irregularly branched; ascospores 9–15-septate, 40–60 × 8–12 μm [holotype: Guinea, s.col. (H-Acharius 629!)] ... Opegrapha striatula Ach.

    2. B. Excipulum laterally (to almost completely) carbonized; lirellae with basal thalline margin; ascospores 11–15-septate, 50–65 × 7–9 μm [holotype: Brazil, Malme 640 (S-6508!)] ... ... Graphis bicrenatula Redinger

    3. C. Excipulum laterally carbonized; lirellae with basal thalline margin, stellately branched; ascospores 9–15-septate, 40–60 × 8–12 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Brazil, Malme 3635 (S-6488!)] ... Graphis striatula var. substellaris Redinger

    Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); pantropical ... Graphis leptoclada Müll. Arg.

    1. A. Excipulum laterally carbonized; ascospores 9–13-septate, 30–50 × 7–10 μm [lectotype (Archer Reference Archer2001e): Indonesia (Java), Junghuhn s.n. (G!)] ... Graphis leptoclada Müll. Arg.

    2. B. Excipulum laterally (to completely) carbonized; ascospores 9–13-septate, 40–65 × 9–12 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Costa Rica, Pittier 5291 (G!)] ... Graphis rigidula Müll. Arg.

    3. C. Excipulum laterally carbonized; ascospores 7–13-septate, 30–50 × 7–10 μm [holotype: Puerto Rico, Fink 1873 (MICH, not seen; isotype: NY!)] ... Graphis rimulosa subsp. lignicola Fink

15. 13(11) Lirellae erumpent to prominent, lacking thalline margin or with basal thalline margin, elongate and irregularly branched (striatula-morph); pantropical ... ... Graphis duplicata Ach.

    1. A. Excipulum laterally carbonized; lirellae lacking thalline margin; ascospores 7–11-septate, 25–45 × 6–8 μm [lectotype (Staiger Reference Staiger2002): South America, s.col. (H-Acharius 586!)] ... ... Graphis duplicata Ach.

    2. B. Excipulum laterally (to almost completely) carbonized; lirellae with basal thalline margin; ascospores 7–13-septate, 25–45 × 7–9 μm [syntypes: Indonesia (Java), s.col. (G!)] ... ... Graphis schizograpta Müll. Arg.

    3. C. Excipulum laterally carbonized; lirellae lacking thalline margin; ascospores 7–9-septate, 25–35 × 6–8 μm [holotype: China, Handel-Mazzetti 10213 (W!)] ... Graphis lopingensis Zahlbr.

    4. D. Excipulum laterally carbonized; lirellae lacking thalline margin; ascospores 9–11-septate, 25–30 × 7–8 μm [holotype: China, Handel-Mazzetti 9103 (W!; isotype: S 2173!)] ... ... Graphis lussuensis Zahlbr.

    Lirellae erumpent, with lateral thalline margin, short and sparsely branched (tenella- morph); pantropical ... Graphis tenella Ach.

    1. A. Ascospores 5–9-septate, 20–30 × 6–8 μm [holotype: Guinea, s.col. (H-Acharius 585!)] ... ... Graphis tenella Ach.

    2. B. Ascospores 5–7-septate, 20–30 × 6–8 μm [holotype: Jamaica, Hart 101 (G!)] ... ... Phaeographis sulcata Müll. Arg.

    3. C. Ascospores 5–7-septate, 15–30 × 7–8 μm [holotype: India, Patwardhan & Nagarkar 77.1495 (AMH, photograph seen; Awasthi Reference Awasthi1991; Nagarkar & Patwardhan Reference Nagarkar and Patwardhan1982)] ... ... Graphis subglauconigra Nagarkar & Patw.

Group 16: Labia striate, excipulum laterally carbonized, hymenium clear, ascospores muriform

1. 1 Norstictic and stictic acid (K+ yellow forming red crystals); lirellae immersed, with lateral thalline margin, very long and radiately branched (dichotoma-morph); Neotropics [holotype: Brazil, Vainio s.n. (TUR-Vainio 27202!; G!)] ... ... Graphis neoelongata Lücking* [≡ Graphis elongata Vain., nom. illeg., non Zenker]

2. No substances (K−); lirellae variable ... 2

3. 2(1) Ascospores 40–75 × 20–30 μm, 2 per ascus; lirellae erumpent (to prominent), with basal thalline margin, short and sparsely branched (striatula-morph); Eastern Palaeotropics [holotype: Malaysia (Borneo: Sabah), Hale 29240 (US!)] ... ... Graphis subintermedians Hale ex Lücking*

   Ascospores 20–60 × 8–16 μm, (2–)4–8 per ascus; lirellae variable ... 3

4. 3(2) Lirellae erumpent, with lateral thallin margin, very long and radiately branched (dichotoma-morph); ascospores 20–35 μm long; Neotropics ... ... Graphis dichotoma (Müll. Arg.) Lücking

   1. A. Ascospores 8/ascus, 20–25 × 7–10 μm [holotype: Brazil, Puiggari 508 (G!)] ... ... Graphina dichotoma Müll. Arg.

   2. B. Ascospores (2–)4–8 per ascus, 25–35 × 10–13 μm [holotype: Puerto Rico, Fink 2049 (MICH, not seen; Wirth & Hale Reference Wirth and Hale1978)] ... Graphis elongatoradians Fink

5. Lirellae erumpent, with lateral to apically thin complete thalline margin, short and sparsely branched (tenella- or symplecta-morph); ascospores 25–60 μm long ... 4

6. 4(3) Ascospores 35–60 μm long; lirellae with lateral thalline margin (tenella-morph); Neotropics (Central America) [holotype: Costa Rica, Chaves 466 (INB!; isotype: CR!)] ... ... Graphis pseudoserpens Chaves, Lücking & Umaña

   Ascospores 25–40 μm long; lirellae with apically thin complete thalline margin (symplecta-morph); Neotropics ... Graphis puiggarii (Müll. Arg.) Lücking

   1. A. Excipulum laterally carbonized; ascospores 25–35 × 8–12 μm [holotype: Brazil, Puiggari 506 (G!)] ... Graphina puiggarii Müll. Arg.

   2. B. Excipulum laterally (to completely) carbonized; ascospores 30–40 × 12–16 μm [holotype: Paraguay, Balansa s.n. (G!)] ... Graphina endoschiza Müll. Arg.

Group 17: Labia striate, excipulum completely carbonized, hymenium clear, ascospores transversely septate

1. 1 Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 2

   No substances (K−) ... 9

2. 2(1) Norstictic acid (K+ yellow forming red crystals) ... 3

   Stictic acid (K+ yellow) ... 7

3. 3(2) Ascospores small (20–40 μm long) ... 4

   Ascospores medium-sized to large (50–120 μm long) ... 5

4. 4(3) Lirellae erumpent, with lateral thalline margin, short and sparsely branched (tenella-morph); Eastern Palaeotropics [holotype: Indonesia, Schiffner 3138 (W!; isotype: FH!)] ... ... Graphis schiffneri Zahlbr.

   Notes. The material identified with that name from Costa Rica (Lücking et al. Reference Lücking and Rivas Plata2008) does not belong to that species but represents an aberrant form of Graphis caesiella in which the labia pruina appears in thin lines making the labia appear striate.

   Lirellae prominent, with basal thalline margin, short and sparsely branched (striatula-morph); pantropical [holotype: India, Watt s.n. (G!)] ... ... Graphis paralleloides Cáceres & Lücking* [≡ Graphis rimulosa var. parallela Müll. Arg., non Graphis parallela Müll. Arg.]

5. 5(3) Lirellae prominent, with thick lateral thalline margin, elongate and irregularly branched (marginata-morph); ascospores medium-sized (50–60 μm long); Neotropics [holotype: Colombia, Lindig 858 (H-Nylander 7604!)] ... Graphis subtracta Nyl.

   Lirellae erumpent to prominent, with apically thin complete thalline margin or thin thalline margin that flakes off to expose black labia; ascospores large (80–120 μm long) ... 6

6. 6(5) Lirellae prominent, with thin thalline margin that often flakes off to expose black labia, short and sparsely branched, with very distinct and coarse striation (lumbricina-morph); ascospores 15–21 μm broad (see also Group 4: 4); Neotropics and Hawaii [lectotype (Wirth & Hale Reference Wirth and Hale1978): Guadeloupe, Duss 1036 (TUR-Vainio 27524!; isolectotype: FH!)] ... Graphis lumbricina Vain.

   Lirellae erumpent to prominent, with persistent, apically thin thalline margin, elongate and irregularly branched, with fine striation; ascospores 6–10 μm broad; Eastern Palaeotropics [holotype: Thailand, Hosseus s.n. (TUR-Vainio 27807!)] ... ... Graphis leptospora Vain.

   Notes. Graphis trichospora, G. verminosa, and G. leptospora share the same lirella morphology, ascospore type, and chemistry, but differ in the degree of excipulum carbonization (apical versus lateral versus complete). As in other cases, and contrary to other authors, we consider these to represent distinct species, but the available material is sparse and the variation in excipulum carbonization cannot be properly assessed at this point.

7. 7(2) Ascospores large (75–90 μm long); lirellae erumpent, with lateral thalline margin, elongate to very long and irregularly branched, labia thinly white-pruinose (albotecta-morph); Eastern Palaeotropics [holotype: India, Sethy & Patwardhan 85.2715 (AMH, photograph seen; Makhija & Adawadkar Reference Adawadkar and Makhija2005)] ... ... Graphis longissima Makhija & Adaw.

   Ascospores small (25–45 μm long); lirellae variable ... 8

8. 8(7) Lirellae erumpent, with apically thin complete thalline margin, elongate and irregularly branched (symplecta-morph); ascospores 5–7-septate, 25–35 μm long; Neotropics (Antilles) and Eastern Palaeotropics [holotype: Vietnam, Balansa s.n. (G!)] ... ... Graphis fumosa Müll. Arg.

   Lirellae with basal thalline margin, elongate and irregularly branched; ascospores 9–13-septate, 35–45 μm long; Eastern Palaeotropics [holotype: India, Patwardhan 78.109 (AMH, photograph seen; Adawadkar & Makhija Reference Adawadkar and Makhija2006)] ... ... Graphis nilgiriensis Adaw. & Makhija

9. 9(1) Ascospores medium-sized to large (50–160 μm long) ... 10

   Ascospores (very) small (15–50 μm long) ... 17

10. 10(9) Ascospores large (most mature ascospores exceeding 70 × 12 μm) ... 11

    Ascospores medium-sized (mature ascospores not exceeding 70 × 12 μm) ... 15

11. 11(10) Lirellae prominent to sessile, lacking or with basal thalline margin (striatula-morph) ... 12

    Lirellae immersed to prominent, with lateral to complete thalline margin ... 13

12. 12(11) Lirellae short and sparsely branched; ascospores 1–2 per ascus; Neotropics [holotype: Brazil, Glaziou s.n.; G, not seen; Awasthi Reference Awasthi1991)] ... Graphis granulata Fée

    Lirellae elongate and irregularly branched; ascospores 6–8 per ascus (see also Group 15:9); Eastern Palaeotropics [lectotype (Awashti & Singh Reference Awasthi and Singh1975): India, Watt 6399 (G, not seen; Awasthi & Singh Reference Awasthi and Singh1975; Hale Index Cards)] ... Graphis contortuplicata Müll. Arg.

13. 13(11) Ascospores 120–160 × 17–25 μm; lirellae prominent, short and sparsely branched (acharii-morph); pantropical [holotype: unknown locality (UPS, not seen; Müller Argoviensis Reference Müller Argoviensis1887)] ... Graphis vestita Fr. [non Fée]

    Ascospores 70–120 × 10–17 μm; lirellae immersed to prominent ... 14

14. 14(13) Lirellae prominent (acharii-morph); pantropical ... Graphis angustata Eschw.

    1. A. Ascospores 11–15-septate, 70–100 × 12–18 μm [holotype: Brazil, Martius s.n. (M, not seen; Hale Index Cards)] ... Graphis angustata Eschw.

    2. B. Ascospores 15–21-septate, 90–120 × 12–15 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Glaziou 5106 (M, not seen; Staiger Reference Staiger2002)] ... Graphis flexibilis Kremp.

    3. Ascospores 15–21-septate, 70–100 × 11–13 μm [lectotype (Lücking et al. Reference Lücking and Rivas Plata2008): Philippines, Elmer 14443 (TUR-Vainio 27866!)] ... Graphis subangustata Vain.

       Notes. Graphis subcurva Zenker might represent an earlier name for this species but we have been unable to locate authentic material.

    Lirellae immersed to erumpent (symplecta-morph); pantropical (also saxicolous) ... Graphis calcea (Fée) A. Massal.

    1. A. Lirellae erumpent; ascospores 15–25-septate, 80–190 × 12–16 μm [holotype: Antilles or South America, s.col. (G, not seen; Wirth & Hale Reference Wirth and Hale1978)] ... Opegrapha calcea Fée

    2. B. Lirellae erumpent; ascospores 15–19-septate, 80–110 × 14–17 μm [holotype: New Caledonia, Saves s.n. (G!)] ... Graphis noumeana Müll. Arg.

    3. C. Lirellae immersed; ascospores 15–19-septate, 70–100 × 14–17 μm [holotype: Brazil, Ule s.n. (G!)] ... Graphis virens Müll. Arg.

    4. D. Lirellae erumpent; ascospores 11–19-septate, 70–120 × 10–15 μm [holotype: Brazil, Ule 273 (G!; isotype: MICH!)] ... Graphis illota Müll. Arg.

    5. E. Lirellae erumpent; ascospores 9–13-septate, 70–100 × 13–16 μm [holotype: Paraguay, Malme 1438 (S 6482!)] ... Graphis virens var. saxicola Redinger

    6. F. Lirellae immersed; ascospores 15–19-septate, 70–90 × 11–14 μm [lectotype (here selected): Brazil, Malme 1293 (S-6484!)] ... Graphis tenuescens var. subimmersa Redinger

15. 15(10) Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); pantropical ... Graphis longula Kremp.

    1. A. Lirellae erumpent, with lateral thalline margin; ascospores 11–17-septate, 50–70 × 10–13 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Glaziou 5497 (M!)] ... Graphis longula Kremp.

    2. B. Lirellae erumpent (to prominent), with (basal to) lateral thalline margin; ascospores 9–17-septate, 60–70 × 9–11 μm [holotype: Brazil, Glaziou 5498 (G!)] ... Graphis flavicans Müll. Arg.

    3. C. Lirellae erumpent, with basal thalline margin; ascospores 11–15-septate, 40–65 × 7–10 μm [lectotype (here selected): Philippines, Copeland 1362 (TUR-Vainio 27862!)] ... ... Graphis supernata Vain.

    Lirellae erumpent to prominent, with apically thin complete thalline margin (upper part of labia dark grey) ... 16

16. 16(15) Lirellae short to elongate, stellately branched (asterizans-morph); thallus verrucose; pantropical [holotype: China (Hong Kong), s.col. (H-Nylander 7781!)] ... ... Graphis asterizans Nyl.

    Lirellae short to elongate, sparsely to irregularly branched (acharii-morph); thallus smooth to uneven; Palaeotropics ... Graphis glauconigra Vain.

    1. A. Ascospores 9–13-septate, 40–80 × 8–12 μm [lectotype (here selected, based on annotation labels by Nakanishi in 1973 and Awasthi & Mathur in 1989): Philippines, Merrill 6791 (TUR-Vainio 27884!)] ... Graphis glauconigra Vain.

    2. B. Ascospores 11–13-septate, 55–65 × 10–12 μm [holotype: Puerto Rico, Fink 1737 (MICH, not seen; isotype: NY!)] ... Graphis tumidulella Fink

    3. C. Ascospores 11–13-septate, 60–70 × 9–11 μm [holotype: China, Chung 393a (W!; isotype: FH!)] ... ... Graphis chungii Zahlbr.

    4. D. Ascospores 11–15-septate, 65–75 × 9–11 μm [holotype: China, Chung 388 (W!; isotype: FH!)] ... Graphis chungii var. oligospora Zahlbr.

17. 17(9) Lirellae broad (0·5–1), erumpent, with basal to lateral thalline margin, short and sparsely branched (tenella-morph); Neotropics [holotype: Brazil, Aptroot 41306 (SP!; isotype: ABL!)] ... Graphis funilina Aptroot*

    Lirellae narrow (0.2–0.4 mm), erumpent to sessile, lacking or with basal to lateral thalline margin ... 18

18. 18(17) Lirellae erumpent to prominent, with lateral thalline margin, short and sparsely branched (tenella-morph); ascospores 15–30 μm long; pantropical ... ... Graphis aurita Eschw.

    1. A. Lirellae prominent, with lateral thalline margin; ascospores 7-septate, 15–25 × 6–8 μm [holotype: Brazil, s.col. (M, not seen; Hale Index Cards)] ... Graphis aurita Eschw.

    2. B. Lirellae erumpent, with lateral thalline margin; ascospores 7–11-septate, 25–30 × 5–7 μm [holotype: India, s.col. (BM, not seen; Awasthi Reference Awasthi1991)] ... Graphis persulcata Stirt.

    3. C. Lirellae erumpent, with lateral thalline margin; ascospores 7–13-septate, 20–45 × 7–8 μm [holotype: Puerto Rico, Fink 1613 (MICH, not seen; Fink Reference Fink1927)] ... Graphis immersa Fink

       Notes. Unfortunately no type material of the above names was available for study and the concept of this taxon must be considered provisional at this point.

    Lirellae erumpent to sessile, lacking thalline margin or with basal thalline margin; ascospores 20–50 μm long ... 19

19. 19(18) Lirellae sessile, very long and radiately branched (slendrae-morph); Eastern Palaeotropics [holotype: Malaysia (Borneo: Sabah), Hale 28009 (US!)] ... ... Graphis slendrae Hale ex Lücking*

    Lirellae erumpent to prominent, short and sparsely branched (striatula-morph) ... 20

20. 20(19) Ascospores 30–50 × 7–13 μm; pantropical ... Graphis rimulosa (Mont.) Trevis.

    1. A. Lirellae erumpent, lacking thalline margin; ascospores 7–11-septate, 30–50 × 7–13 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Guyana, Leprieur 200 (PC, not seen; Hale Index Cards)] ... ... Opegrapha rimulosa Mont.

    2. B. Lirellae prominent, with basal thalline margin; ascospores 7–11-septate, 35–50 × 7–10 μm [holotype: Australia, Shirley 1821 (G!)] ... Graphis rimulosa var. brachycarpa Müll. Arg.

       Notes. Graphis rimulosa var. brachycarpa was listed as synonym of G. striatula by Archer (Reference Archer2006), but the type has a completely carbonized excipulum and slightly smaller ascospores and thus agrees with G. rimulosa.

    Ascospores 20–40 × 5–9 μm; pantropical ... Graphis dupaxana Vain.

    1. A. Lirellae prominent, lacking thalline margin; ascospores 7–9-septate, 25–40 × 7–9 μm [lectotype (here selected, based on annotation label by Nakanishi in 1973): Philippines, McGregor 14313 (TUR-Vainio 27869!)] ... Graphis dupaxana Vain.

    2. B. Lirellae prominent, with basal thalline margin; ascospores 5–11-septate, 20–35 × 5–8 μm [holotype: Indonesia (Borneo), Moulton s.n. (TUR-Vainio 27859!)] ... Graphis moultonii Vain.

    3. C. Lirellae prominent, with basal thalline margin; ascospores 3–7-septate, 15–20 × 7–8 μm [holotype: Brazil, Puiggari 136 (G, not seen; Müller Argoviensis Reference Müller Argoviensis1880; Redinger Reference Redinger1935)] ... ... Graphis leioplaca Müll. Arg.

Group 18: Labia striate, excipulum completely carbonized, hymenium clear, ascospores muriform

1. 1 Ascospores terminally muriform, with longitudinal septa in terminal segments only ... 2

   Ascospores regularly muriform or submuriform with at least a few longitudinal septa in middle segments ... 4

2. 2(1) No substances (K−); ascospores large (80–150 μm long); lirellae prominent, with apically thin complete margin; elongate and irregularly branched (acharii-morph); Neotropics and Eastern Palaeotropics ... Graphis vestitoides (Fink) Staiger

   1. A. Ascospores 80–140 × 12–16 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Puerto Rico, Fink 1986 (MICH, not seen; Staiger Reference Staiger2002)] ... Graphina vestitoides Fink

   2. B. Ascospores 90–140 × 11–16 μm [lectotype (here selected): Philippines, Elmer 14443 (TUR-Vainio 27224!; isolectotype: FH!)] ... Graphis fruticicola Vain.

   3. C. Ascospores 80–150 × 12–17 μm [holotype: Brazil, Malme 3531 (S!)] ... ... Phaeographis cerviculata Redinger

   Norstictic acid (K+ yellow forming red crystals); ascospores and lirellae variable ... 3

3. 3(2) Ascospores medium-sized (50–80 μm long); lirellae prominent, with lateral thalline margin, short and stellately branched (stellata-morph); Neotropics and Eastern Palaeotropics [holotype: Brazil, s.col. (G, not seen; Wirth & Hale Reference Wirth and Hale1978; Hale Index Cards)] ... ... Graphis congesta (Fée) Müll. Arg.

   Notes. This taxon is usually understood as having transversely septate ascospores (Wirth & Hale Reference Wirth and Hale1978), which would make it a synonym of G. marginata, but the ascospores contain (easily overlooked) terminal longitudinal septa.

   Ascospores large (80–150 μm long); lirellae prominent, with apically thin complete margin; elongate and irregularly branched (acharii-morph); Neotropics and Eastern Palaeotropics [holotype: Thailand, Sutjaritturakan 12736 (RAMK!)] ... ... Graphis norvestitoides Sutjaritturakan

   Notes. This taxon was erroneously identified with Graphis aquilonia by Lücking et al. (Reference Lücking and Rivas Plata2008) but the latter differs in the entire labia and lirellae with apically thick complete thalline margin.

4. 4(1) Norstictic or stictic acid (K+ yellow or K+ yellow forming red crystals) ... 5

   Notes. If protocetraric acid (P+ red) and ascospores I− and very small (15–25 μm long), cf. Graphis pauaiensis Vain which probably belongs in Carbacanthographis.

   No substances or hypostictic acid (K−) ... 7

5. 5(4) Ascospores single, medium-sized to large (60–140 × 20–35 μm); lirellae erumpent, with apically thin complete margin, elongate and irregularly branched (symplecta-morph); norstictic acid; Neotropics [holotype: Colombia, Lindig s.n. (H-Nylander 7530!)] ... Graphis componens Nyl.

   Ascospores 4–8 per ascus, small (15–50 × 7–12 μm); lirellae and chemistry variable ... 6

6. 6(5) Ascospores 4 per ascus, 30–50 μm long; norstictic acid; lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Neotropics [holotype: Venezuela, Neuwirth 8517 (W!)] ... ... Graphis breussii Neuwirth & Lücking

   Ascospores 8 per ascus, 15–25 μm long; stictic acid; lirellae erumpent to prominent, lacking or with basal thalline margin, elongate and irregularly branched (striatula-morph); Eastern Palaeotropics [holotype: China, Handel-Mazzetti 11507 (W!)] ... ... Graphis hunanensis (Zahlbr.) M. Nakan. & Kashiw.

7. 7(4) Ascospores (medium-sized to) large [(70–)80–170 × 15–45 μm, most mature ascospores exceeding 80 μm] ... 8

   Ascospores small to medium-sized (20–80 × 10–25 μm) ... 13

8. 8(7) Ascospores 2–6 per ascus; lirellae prominent to sessile, with apically thin complete thalline margin, short to elongate and sparsely to irregularly branched (acharii-morph); Neotropics and Eastern Palaeotropics (also saxicolous) ... ... Graphis acharii Fée

   1. A. Thallus smooth to uneven; lirellae prominent, elongate and irregularly branched; ascospores 80–170 × 15–30 μm [holotype: South America, s.col. (G; not seen; isotype: L!)] ... ... Graphis acharii Fée

   2. B. Thallus smooth to uneven; lirellae prominent, short and sparsely branched; ascospores 90–150 × 18–30 μm [holotype: Brazil, Glaziou 5486 (G!)] ... Graphis curta Fée

   3. C. Thallus verrucose; lirellae prominent, elongate and irregularly branched; ascospores 100–120 × 20–30 μm [holotype: Brazil, Glaziou 11777 (G!)] ... Graphina tuberculifera Müll. Arg.

   4. D. Thallus smooth to uneven; lirellae sessile, short and sparsely branched; ascospores 100–150 × 20–30 μm [holotype: Brazil, Glaziou 16671 (G!)] ... Graphina pachypleura Müll. Arg.

   Ascospores single or 1–2 per ascus; lirellae variable ... 9

9. 9(8) Thallus and lirellae verrucose; hypostictic acid ... 10

   Thallus and lirellae smooth to uneven; no substances ... 11

10. 10(9) Lirellae sessile, with distinctly verrucose, apically thin complete thalline margin, very short and unbranched (granulosa-morph); Neotropics [holotype: Jamaica, Hart s.n. (G!)] ... Graphis granulosa (Müll. Arg.) Lücking

    Lirellae immersed to erumpent, with slightly verrucose, apically thick complete thalline margin, elongate and irregularly branched (consanguinea-morph); Neotropics and Eastern Palaeotropics ... Graphis dealbata Nyl.

    1. A. Ascospores 80–110 × 20–35 μm [holotype: Brazil, Glaziou 1899 (H-Nylander 7531!)] ... ... Graphis dealbata Nyl.

    2. B. Ascospores 80–100 × 20–30 μm μm [holotype: Brazil, Vainio s.n. (TUR-Vainio 27177!)] ... ... Graphis albostriata Vain.

11. 11(9) Lirellae erumpent, with lateral thalline margin (tenella-morph); Neotropics and Eastern Palaeotropics (Thailand) ... Graphis myrtacea (Müll. Arg.) Lücking

    1. A. Ascospores single, 100–150 × 25–40 μm [holotype: Brazil, Ule 127 (G!)] ... ... Graphina myrtacea Müll. Arg.

    2. B. Ascospores 1–2 per ascus, 75–115 × 30–45 μm [lectotype (Staiger Reference Staiger2002): Brazil, Malme 2311 (S!)] ... Graphina crystallifera Redinger

    Lirellae immersed to prominent, with apically thin to thick complete thalline margin (acharii- or consanguinea-morph) ... 12

12. 12(11) Lirellae prominent, with apically thin complete thalline margin (acharii-morph); Neotropics ... Graphis macella Kremp.

    1. A. Ascospores single, 70–170 × 20–40 μm [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Glaziou 6289b (M!; Staiger Reference Staiger2002)] ... Graphis macella Kremp.

    2. B. Ascospores single, 80–150 × 20–30 μm [holotype: Brazil, Vainio s.n. (TUR-Vainio 27186!)] ... Graphis subvestita Vain.

       Ascospores 1–2 per ascus, 85–135 × 17–25 μm [holotype: Brazil, Mosén 2143 (S-2203!)] ... ... Graphina carbocarpa Redinger

    Lirellae immersed to erumpent, with apically thick complete thalline margin (consanguinea-morph); Neotropics ... ... Graphis consanguinea (Müll. Arg.) Lücking

    1. A. Lirellae immersed; ascospores single, 110–125 × 30–35 μm [holotype: Brazil, Glaziou s.n. (G!)] ... ... Graphina consanguinea Müll. Arg.

    2. B. Lirellae erumpent; ascospores single, 70–170 × 25–40 μm [holotype: Brazil, Vainio s.n. (TUR-Vainio 27092!; isotype: M)] ... Graphina includens Vain.

    3. C. Lirellae erumpent; ascospores single, 140–170 × 30–40 μm [lectotype (Staiger Reference Staiger2002): Brazil, Vainio s.n. (M; Staiger Reference Staiger2002; isolectotype: TUR-Vainio 27244!)] ... ... Graphina pseudosophistica Vain.

    4. D. Lirellae erumpent; ascospores single, 90–120 × 25–35 μm [holotype: Brazil, Glaziou 5017b (W!)] ... Graphina nylanderiana Zahlbr. [non Patw. & C. R. Kulk.]

    5. E. Lirellae erumpent; ascospores single, 90–150 × 25–35 μm [holotype: Brazil, Malme 1227B (S, not seen; Redinger Reference Redinger1934)] ... Graphina heteroplacoides Redinger

13. 13(7) Lirellae prominent, lacking thalline margin (striatula-morph); thallus yellow-green to olive-yellow; Central America [holotype: Costa Rica, Lücking 16661 (USJ!; isotype: F!)] ... ... Graphis fournierii Lizano & Lücking

    Notes. If lirellae with distinct brown pruina along slit, cf. Glyphis substriatula (Nyl.) Staiger*.

    Lirellae erumpent to prominent, with lateral to complete thalline margin; thallus white-grey to rarely olive-grey ... 14

14. 14(13) Labia white-pruinose; lirellae erumpent, with lateral thalline margin, short and sparsely branched (albotecta-morph); mature ascospores not exceeding 45 μm; South America ... Graphis albotecta (Redinger) Staiger

    1. A. Ascospores 30–45 × 10–19 μm [holotype: Brazil, Malme 3532 (S 5528!)] ... ... Graphina albotecta Redinger

    2. B. Ascospores 30–45 × 10–15 μm [holotype: Brazil, Malme 1089 (S 2190!)] ... ... Graphina rimulosa Redinger

    Labia non-pruinose; lirellae erumpent to prominent, with lateral to complete thalline margin; mature ascospores exceeding 45 μm ... 15

15. 15(14) Mature ascospores exceeding 65 μm ... 16

16. Mature ascospores 30–65 μm long ... 17

17. 16(15) Lirellae immersed, with apically thick complete thalline margin, elongate and irregularly branched (consanguinea-morph); ascospores 20–30 μm broad; Eastern Palaeotropics (China) [lectotype (here selected, based on annotation label by Nakanishi in 1973): China, Chung 602b (W!)] ... Graphis plumbea (Zahlbr.) Lücking*

    Labia prominent, with apically thin complete thalline margin, short and sparsely branched (acharii-morph); ascospores 12–15 μm broad; Eastern Palaeotropics [holotype: Taiwan, Asahina 345 (W!; isotype: US!)] ... ... Graphis oxyspora (Zahlbr.) Lücking*

18. 17(15) Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched (tenella-morph); ascospores 10–15 μm broad; Neotropics and African Palaeotropics [lectotype (Wirth & Hale Reference Wirth and Hale1978): Brazil, Malme 998 (S 6508!)] ... ... Graphis plurispora (Redgr.) Lücking & Chaves

    Lirellae erumpent to prominent, with apically thin complete thalline margin; ascospores 15–23 μm broad ... 18

19. 18(17) Lirellae very short and unbranched (multisulcata-morph); Neotropics [holotype: Brazil, Puiggari 74 (G!)] ... Graphis multisulcata (Müll. Arg.) Lücking & Chaves

    Lirellae elongate to very long and irregularly to radiately branched (subradiata-morph); Neotropics [holotype: Colombia, Lindig 2725 (H-Nylander 7434!)] ... ... Graphis subradiata (Nyl.) Lücking*

    Notes. Graphis analoga var. subradiata Nyl. was considered a synonym of G. multisulcata by Lücking et al. (Reference Lücking and Rivas Plata2008), but the two differ markedly in their lirella morphology. The material referred to as G. multisulcata by Lücking et al. (Reference Lücking and Rivas Plata2008) belongs to G. subradiata.

Group 19: Labia striate, excipulum completely carbonized, hymenium inspersed, ascospores transversely septate

1. 1 Ascospores small (25–35 × 5–7 μm), 9–13-septate; stictic acid (K+ yellow); lirellae erumpent, lacking thalline margin (striatula-morph); Neotropics (Central America) [holotype: Costa Rica, Aptroot 60569 (INB!)] ... Graphis syzygii Aptroot*

   Notes. This new species was not recognized in the recent treatment by Lücking et al. (Reference Lücking and Rivas Plata2008).

   Ascospores large to very large (75–200 × 15–30 μm); norstictic acid (K+ yellow forming red crystals) or no substances (K−) ... 2

2. 2(1) No substances (K−); ascospores 100–200 × 18–30 μm; lirellae prominent, with apically thin complete thalline margin, short and sparsely branched (acharii-morph); Neotropics and Eastern Paleotropics [holotype: South America, s.col. (G!)] ... Graphis cinerea Fée

   Norstictic acid (K+ yellow forming red crystals); ascospores 75–100 × 15–22 μm; lirellae variable ... 3

3. 3(2) Lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Eastern Palaeotropics [holotype: Indonesia (Java), Groenhart 5190 (L!)] ... Graphis leucaenae Aptroot*

   Lirellae erumpent, lacking thalline margin, short and sparsely branched (striatula-morph); Eastern Palaeotropics [holotype: Papua New Guinea, Aptroot 37028 (B!)] ... ... Graphis inspersolongula Aptroot*

Group 20: Labia striate, excipulum completely carbonized, hymenium inspersed, ascospores muriform

1. 1 Ascospores terminally muriform; lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph); Neotropics (Central America) and Eastern Palaeotropics (Thailand) [holotype: Costa Rica, Chaves 423 (INB!)] ... Graphis subflexibilis Lücking & Chaves

   Ascospores regularly muriform; lirellae variable ... 2

2. 2(1) Norstictic acid (K+ yellow forming red crystals); ascospores small (20–45 × 9– 12 μm); lirellae erumpent, with lateral thalline margin, short and sparsely branched (tenella-morph); Neotropics [holotype: Guadeloupe, Duss 1590 (TUR-Vainio 27248!)] ... Graphis pseudoanaloga Vain.

   No substances (K−); ascospores medium-sized to large (50–180 × 9–35 μm); lirellae prominent, with apically thin complete thalline margin, elongate and irregularly branched (acharii-morph) ... 3

3. 3(2) Ascospores medium-sized (50–70 × 9–12 μm); Neotropics (Central America) [holotype: Costa Rica, Lücking 15235a (F!)] ... Graphis pseudocinerea Lücking & Umaña

   Ascospores large (80–180 × 10–35 μm) ... 4

4. 4(3) Ascospores very long and narrow (120–180 × 10–15 μm), more than 10 times as long as broad, with only some segments having longitudinal septa (submuriform); Neotropics (Central America) [holotype: Costa Rica, Sipman 48019a (INB!)] ... ... Graphis altamirensis Sipman & Lücking

   Ascospores shorter and broader (80–140 × 15–35 μm), less than 10 times as long as broad, with all segments having longitudinal septa (muriform) ... 5

5. 5(4) Ascospores 1–4 per ascus, 25–35 μm broad, about 3–4 times as long as broad (see also Group 11: 7); pantropical ... Graphis phaeospora Vain.

   Labia striate [original material: not designated (Staiger Reference Staiger2002)] ... ... Graphis subcinerea Staiger [nom. inval. (not validly published)]

   Ascospores 4–8 per ascus, 15–25 μm broad, about 5–6 times as long as broad (see also Group 11: 7); Neotropics [holotype: Costa Rica, Lücking 15269d (INB!)] ... ... Graphis argentata Lücking & Umaña

   Labia striate [original material: not designated (Lücking et al. Reference Lücking and Rivas Plata2008)] ... ... Graphis cinereoides Lücking & Chaves [nom. inval. (not validly published)]

Carbacanthographis cleitops (Fée) Lücking comb. nov

Graphis cleitops Fée, Suppl. Essai Crypt. Écorc.: 32 (1837); Graphina cleitops (Fée) Müll. Arg., Mém. Soc. Phys. Hist. Nat. Genève 29(8): 41 (1887); type: Unknown locality, s.col. (G!— holotype).

Carbacanthographis coccospora (Aptroot) Aptroot & Lücking comb. nov

Graphis coccospora Aptroot, Fungal Div. 9: 19 (2002); type: Brazil, Aptroot 41071 (ABL!—holotype).

Carbacanthographis induta (Müll. Arg.) Lücking comb. nov

Graphis induta Müll. Arg., Hedwigia 30: 185 (1891); type:): Vietnam, Balansa s.n. (G! —holotype).

Notes. The type material has Carbacanthographis-like lirellae and I− ascospores, which justifies recombination in Carbacanthographis. The only species in that genus with similar ascospores is C. candidata (Nyl.) Staiger & Kalb, which produces protocetraric acid. Norstictic acid as major substance was previously unknown in Carbacanthographis (Staiger Reference Staiger2002; Archer Reference Archer2006).

Carbacanthographis triphoroides (M. Wirth & Hale) Lücking comb. nov

Graphina triphoroides M. Wirth & Hale, Smithson. Contr. Bot. 40: 44 (1978); Graphis triphoroides Nyl., Flora 69: 103 (1886); nom. inval.; type: Cuba, Wright 15 (H-Nylander 7520!—holotype; US!—isotype).

Notes. This taxon was indicated as a synonym of Graphina crassa (Müll. Arg.) [≡ Carbacanthographis crassa (Müll. Arg.) Staiger & Kalb] by Nakanishi (annotation label in 1973). However, the two species differ markedly: C. crassa has a completely carbonized excipulum, single-spored asci, and the ascospores are I− (I+ weakly violet in C. triphoroides).

Glyphis atrofusca (Müll. Arg) Lücking

In Archer, Flora of Australia 57: 651 (2009); Graphina atrofusca Müll. Arg., Flora 70: 74 (1887); Graphis atrofusca (Müll. Arg.) Stizenb., Ber. Tät. St. Gallisch. Naturw. Gesellsch. 1889–1890: 186 (1891); type: South Africa, Wilms 70 (G!— lectotype, here selected).

Graphina polycarpa Müll. Arg., Flora 70: 63 (1887); Graphis polycarpa (Müll. Arg.) Stizenb., Ber. Tät. St. Gallisch. Naturw. Gesellsch. 1889–1890: 184 (1891); type: South Africa, Wilms 48 (G!—(holotype).

Graphina montoensis A. W. Archer, Mycotaxon 77: 168 (2001); Glyphis montoensis (A. W. Archer) Staiger, Biblioth. Lichenol. 85: 173 (2002); type: Australia, Streimann 9845 (CANB!—holotype).

Notes. Revision of type material of supposed Graphis species turned up two earlier names for the species named Glyphis montoensis by Staiger (Reference Staiger2002). Both names were described in the same paper and probably from the same population, but since the type material of Graphina atrofusca (Wilms 70) is best developed, the epithet atrofusca is here selected for the species. The Wilms collections underline the ecological preference of this species for rather dry, partially exposed habitats.

Glyphis substriatula (Nyl.) Staiger

Graphina sulcatula Müll. Arg., Rev. Mycol. 10: 119 (1888); type Paraguay, Balansa s.n. (G!—holotype).

Graphina sulcatula var. conglomerata Müll. Arg., Rev. Mycol. 10: 119 (1888); type: Paraguay, Balansa 4185 (G!—lectotype, selected here).

Graphina bakeri Zahlbr., Ann. Mycol. 2: 269 (1904); Graphina sulcatula var. bakeri (Zahlbr.) Redinger, Arkiv Bot. 26A(1): 31 (1934); type: Nicaragua, Baker 2372 (W!—holotype).

Notes. Revision of type material of Graphis species revealed three synonyms of this widespread neotropical taxon, although the synonymy of Graphina sulcatula var. conglomerata is provisional; the type material has strongly aggregate lirellae and might represent a separate species.

Graphis apoda (Zahlbr.) Lücking comb. et stat. nov

Graphis illinata var. apoda Zahlbr. in Magnusson & Zahlbruckner, Ark. Bot. 31A(1): 32 (1944); type: USA. (Hawaii), Rock 131 (W!—holotype).

Graphis brahmanensis Aptroot sp. nov

Sicut Graphis duplicata sed thallo acido stictico continente differt.

Typus: Papua New Guinea, Madang, Ramu valley, Brahman, along road to Bundi, on branches of Dipterocarpaceae tree in virgin lowland rain forest, 200 m, August 1992, Aptroot 31685 (B!—holotypus).

(Fig. 13A)

Fig. 13. New species in Graphis (all holotypes). A, G. brahmanensis; B, G. cupei; C, G. evirescens (new combination); D, G. funilina; E, G. leucenae; F, G. lourdesina. Scale = 1 mm.

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae erumpent, with basal thalline margin, elongate and irregularly branched, 1–5 × 0·1–0·2 mm; labia eventually striate. Excipulum laterally carbonized; hymenium clear. Ascospores 8 per ascus, transversely 9–11-septate, 20–30 × 4–6 μm, hyaline.

Secondary chemistry. Stictic and hypostictic acid.

Notes. This new species is a typical representative of the Graphis duplicata group and differs from G. duplicata by the presence of stictic acid and the less prominent lirellae. Graphis stenotera is very similar in morphology and chemistry but has an apically carbonized excipulum only.

Graphis cincta (Pers.) Aptroot

Aptroot in Archer, Fl. Australia 57: 651 (2009); Opegrapha cincta Pers., Ann. Wetter. Ges. 2: 15 (1811); type: Dominican Republic, s.col. (L!—holotype; L!—isotypes).

Graphis cremicolor (H. Magn.) Lücking & Archer comb. nov

Graphina cremicolor H. Magn., Ark. Bot., Ser. 2, 3(10): 267 (1955); type: USA (Hawaii), Faurie 720 (UPS—holotype, not seen; Magnusson Reference Magnusson1955).

Graphis cupei Vain. ex Lücking sp. nov

Sicut Graphis anfractuosa sed ascosporis majoribus differt.

Typus Cuba, Ekman 19 (TUR-Vainio 27371!—holotypus).

(Fig. 13B)

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae prominent, with basal thalline margin, elongate and irregularly branched, 1–5 × 0·3–0·4 mm; labia eventually striate. Excipulum completely carbonized; hymenium inspersed. Ascospores 4–8 per ascus, transversely 13–17-septate, 60–70 × 7–9 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. Graphis cupei is here validated since it represents a distinct taxon with no other name available, characterized by the rather robust lirellae with inspersed hymenium and medium-sized ascospores and lacking secondary substances. Most similar is G. anfractuosa but that species has much smaller ascospores and completely lacks a thalline margin.

Graphis enteroleuca (Ach.) Lücking comb. nov

Opegrapha enteroleuca Ach., Syn. Meth. Lich.: 78 (1814) [non Fée 1827 nec Nyl. 1853]; type: South America, unknown locality, s.col. (H-Acharius 658!—holotype).

Graphis galactoderma (Zahlbr.) Lücking comb. nov

Graphina galactoderma Zahlbr. in Handel-Mazzetti, Symb. Sin. 3: 54, 57 (1930); type: China, Handel-Mazzetti 5912 (W!).

Graphis evirescens (Redinger) Lücking comb. nov. (Fig. 13C)

Graphina evirescens Redinger, Ark. Bot. 26A(1): 10, 43 (1933); type Brazil, Malme 3527 (S-5540!—holotype).

Graphis funilina Aptroot sp. nov

Sicut Graphis noumeana sed ascosporis minoribus differt.

Typus: Brazil, Minas Gerais, Parque do Caraça, near Funil, on sandstone, 1350 m, September 1997, Aptroot 41306 (SP!—holotypus; ABL!—isotypus). Paratypes: same locality, same date, Aptroot 41305 (ABL, SP); same locality, along track to Varginha chapel, 1250 m, September 1997, Aptroot 41015 (ABL, SP).

(Fig. 13D)

Thallus corticate, smooth to uneven, white to pale yellowish grey.

Lirellae erumpent, with basal to lateral thalline margin, elongate and irregularly branched, 1–5 × 0·7–1·2 mm; labia striate. Excipulum completely carbonized; hymenium clear. Ascospores 8 per ascus, 7-septate, 30–40 × 10–12 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. Most similar among species with small, transversely septate ascospores and lirellae with striate labia and completely carbonized excipulum is G. aurita, which differs in the much narrower lirellae and the smaller ascospores.

Graphis ingarum (Vain.) Lücking comb. et stat. nov

Graphis angustata var. ingarum Vain., Ann. Acad. Sci. Fenn., Ser. A, 6(7): 158 (1915). type: Guadeloupe, Duss 1190 (TUR-Vainio 27848!—holotype).

Graphis inspersolongula Aptroot sp. nov

Sicut Graphis longula sed hymenio insperso et thallo acido norstictico continente differt.

Typus: Papua New Guinea, Central Province, Owen Stanley Range, Myola, on tree trunk at margin of primary montane forest, 2100 m, October 1995, Aptroot 37049 (B!—holotypus).

Thallus corticate, smooth to uneven, pale grey.

Lirellae erumpent, lacking thalline margin, short and sparsely branched, 1–3 × 0·4–0·7 mm; labia striate. Excipulum completely carbonized; hymenium inspersed (type A). Ascospores 8 per ascus, 13–19-septate, 75–100 × 15–20 μm, hyaline.

Secondary chemistry. Norstictic acid.

Notes. This species resembles Graphis longula but differs in the inspersed hymenium, the presence of norstictic acid, and the shorter lirellae.

Graphis isidiata (Hale) Lücking comb. nov

Thelotrema isidiatum Hale, Mycotaxon 3: 178 (1975); Type: Venezuela, Hale 42425 (US!—holotype).

Graphis japonica (Müll. Arg.) A. W. Archer & Lücking comb. nov

Graphina japonica Müll. Arg., Flora 74: 113 (1891); type: Japan, Miyoshi 23 (G!—holotype).

Graphis jeanmuelleri Lücking nom. nov

Graphina elegantula Müll. Arg., Flora 63: 23 (1880); non Graphis elegantula Zahlbr. in Handel-Mazzetti, Symb. Sin. 3: 39, 50 (1930); type: Brazil, Puiggari s.n. (G!—holotype).

Graphis kousyuensis (Horik. & M. Nakan.) Lücking comb. nov

Graphina kousyuensis Horik. & M. Nakan. in Nakanishi, Journal of Science of the Hiroshima University, Series B, 2, 11: 93 (1966), 265 (1967); type: Japan, Nakanishi 1902 (HIRO—holotype, not seen).

Graphis leucaenae Aptroot sp. nov

Sicut Graphis cinerea sed ascosporis minoribus et acidum norsticticum continente differt.

Typus: Indonesia (Java), Malang, Agricultural School gardens, on trunk of Leucaena glauca, March 1940, Groenhart 5190 (L!—holotype; B!—isotype).

(Fig. 13E)

Thallus corticate, smooth to uneven, pale grey.

Lirellae erumpent to prominent, with apically thin complete thalline cover (thick laterally), elongate and irregularly branched, 1–5 × 0·7–1·2 mm; labia striate. Excipulum completely carbonized; hymenium inspersed (type B). Ascospores 8 per ascus, 13–19-septate, 75–100 × 15–22 μm, hyaline.

Secondary chemistry. Norstictic acid.

Notes. This species is most similar to Graphis cinerea, but differs in the smaller ascospores (100–200 × 18–30 μm in the latter) and the rather thick lateral part of the thalline cover, as well as the presence of norstictic acid.

Graphis lourdesina Aptroot sp. nov

Sicut Graphis ruiziana sed ascosporis majoribus differt.

Typus Brazil, Minas Gerais, Parque do Caraça, near Gruta de Lourdes, on bark of Eucalyptus, 1450 m, September 1997, Aptroot 41545 (SP!—holotypus; B!—isotypus).

(Fig. 13F)

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae prominent to sessile, lacking thalline margin, very short and unbranched, 0·5–2·5 × 0·4–0·6 mm; labia entire. Excipulum completely carbonized; hymenium clear. Ascospores 2 per ascus, muriform, 100–120 × 30–35 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. This new species is closely related to Graphis ruiziana and G. subruiziana, but differs in the large ascospores, which measure 35–65 × 10–20 μm in G. ruiziana (4–8 per ascus) and 60–80 × 20–30 μm in G. subruiziana (2–4 per ascus).

Graphis myolensis Aptroot sp. nov

Sicut Graphis lapidicola sed lirellis prominentibus margine thallino basali differt.

Typus: Papua New Guinea, Central Province, Owen Stanley Range, Myola, along Iora creek, on tree trunk in primary montane forest, 2100 m, October 1995, Aptroot 37428 (B!).

Thallus corticate, smooth to uneven, white to pale yellowish grey.

Lirellae prominent, with basal thalline margin, short and sparsely branched, 1–4 × 0·2–0·4 mm; labia entire. Excipulum laterally carbonized; hymenium clear. Ascospores 8 per ascus, muriform, 60–70 × 11–13 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. This new species is most similar to Graphis lapidicola and G. xylophaga, which differ chiefly in their erumpent lirellae with rather thick lateral to complete thalline margin.

Graphis nadurina Aptroot sp. nov

Sicut Graphis cleistomma sed ascosporis minoribus differt.

Typus: Papua New Guinea, Central Province, Owen Stanley Range, surroundings of Naduri village, on tree trunk in secondary vegetation, 1600 m, October 1995, Aptroot 38161 (B!—holotypus). Paratypes: Papua New Guinea, Madang, Manam Isl., 450 m, Jul 1992, Aptroot 30513 (ABL); ibid., Aug 1979, Iserentant 9364aa (ABL, LG).

(Fig. 14A)

Fig. 14. New species in Graphis (all holotypes). A, G. nadurina; B, G. norstictica; C, G. sarawakensis; D, G. slendrae; E, G. subintermedians; F, G. syzygii. Scale = 1 mm.

Thallus corticate, smooth to uneven, pale yellowish grey.

Lirellae prominent to sessile, with apically thick complete thalline margin, very short and unbranched, 0·5–2·5 × 0·4–0·6 mm; labia entire. Excipulum (laterally to) completely carbonized; hymenium clear. Ascospores 2 per ascus, muriform, 40–50 × 14–16 μm, hyaline.

Secondary chemistry. Norstictic and connorstictic acids.

Notes. This new species is most similar to Graphis cleistomma, which differs in having much larger ascospores.

Graphis negrosina (Vain.) Lücking comb. et stat. nov

Graphis duplicata var. negrosina Vain., Ann. Acad. Sci. Fenn., Ser. A, 15(6): 245 (1921); type: Philippines, Merrill 6790 (TUR-Vainio 27630!—holotype).

Graphis neoelongata Lücking nom. nov

Graphis elongata Vain., Acta Soc. Fauna Fl. Fenn. 7: 107 (1890); non Graphis elongata Zenker in Goebel & Kunze, Pharmaceutische Waarenkunde 1: 165 (1829); type: Brazil, s.col. (TUR-Vainio 27202!—holotype).

Graphis norstictica Archer & Lücking sp. nov

Sicut Graphis borealis sed ascosporis angustioribus differt.

Typus: Philippines, Baker 1253 (TUR-Vainio 27260!—holotypus).

(Fig. 14B)

Thallus corticate, smooth to uneven, white to pale yellowish grey.

Lirellae erumpent, with lateral thalline margin, elongate and irregularly branched, 1–5 × 0·2–0·3 mm; labia entire. Excipulum laterally carbonized; hymenium clear. Ascospores 8 per ascus, muriform, 35–50 × 17–25 μm, hyaline.

Secondary chemistry. Norstictic acid.

Notes. The type material of G. nanodes Vain. consists of two syntypes, one containing norstictic acid (Baker 1253) and the other no lichen substances (Copeland 1382). Since Vainio (1921) reported the thallus of G. nanodes as K−, the latter was selected as lectotype of that species. For the well-developed collection Baker 1253, no name is available, and it is here described as new. Graphis norstictica belongs in a group of three closely related species with similar lirella morphology and anatomy (lineola-morph) and similar chemistry, differing chiefly in ascospore size: G. renschiana with small ascospores 20–35 × 8–14 μm, G. norstictica with larger, narrow ascospores 35–50 × 10–16 μm, and G borealis with larger, broad ascospores 35–50 × 17–25 μm. The species is also known from Kenya and Papua New Guinea.

Graphis novopalmicola A. W. Archer & Lücking nom. nov

Graphina palmicola Müll. Arg., Flora 70: 402 (1887); non Graphis palmicola Makhija & Adaw.; type: Australia, Sayer s.n. (G!—holotype).

Graphis oxyspora (Zahlbr.) Lücking comb. nov

Graphina oxyspora Zahlbr., Fedde Repert. 31: 214 (1933); type: Taiwan, Asahina 345 (W!—lectotype; US!—isolectotype).

Graphis paralleloides Cáceres & Lücking nom. nov

Graphis rimulosa var. parallela Müll. Arg., J. Linn. Soc. Bot. 29: 224 (1882); Graphis parallela (Müll. Arg.) Cáceres & Lücking in Cáceres, Libri Bot. 22: 84 (2007) [nom. illeg.; non Graphis parallela Müll. Arg.]; type: India, Watt s.n. (G!—holotype).

Graphis plumbea (Zahlbr.) Lücking comb. nov

Graphina plumbea Zahlbr. in Handel-Mazzetti, Symb. Sin. 3: 54, 55 (1930); type: China, Chung 602b (W!—lectotype).

Graphis riopiedrensis (Fink) Lücking comb. nov

Graphina riopiedrensis Fink, Mycologia 19: 217 (1927); type: Puerto Rico, Fink 2167 (MICH—holotype, not seen; FH!—isotype).

Graphis sarawakensis Hale ex Lücking sp. nov

Sicut Graphis subturgidula sed ascosporis regulariter muriformibus latioribusque et thallo verrucoso differt.

Typus: Indonesia (Borneo: Sarawak), Hale 30435 (US!—holotype).

(Fig. 14C)

Thallus corticate, verrucose, white to pale grey.

Lirellae sessile, with thick complete, laterally verrucose thalline cover, elongate and irregularly branched, 1–5 × 0·4–0·6 mm; labia entire. Excipulum completely carbonized; hymenium clear. Ascospores 2–4 per ascus, muriform, 80–100 × 15–20 μm, hyaline.

Secondary chemistry. Stictic acid.

Notes. This species is most similar to the recently described Graphis subturgidula but differs in several features, notably the verrucose thallus, the sessile lirellae with rather thick thalline cover, and the regularly muriform, broader ascospores. Graphis granulosa is morphologically similar but has indistinctly striate labia and contains hypostictic acid.

Graphis semirigida (Müll. Arg.) Lücking comb. nov

Graphina semirigida Müll. Arg., J. Linn. Soc. London, Bot., 29: 226 (1892); type: India, Watt 83 (G!—isotype).

Graphis slendrae Hale ex Lücking sp. nov

Sicut Graphis dupaxana sed lirellis sessilibus radiatis differt.

Typus: Malaysia (Borneo: Sabah), Hale 28009 (US!—holotypus).

(Fig. 14D)

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae prominent, with basal thalline margin, very long and radiately branched, 1–10 × 0·1–0·2 mm; labia striate. Excipulum completely carbonized; hymenium clear. Ascospores 8 per ascus, 5–7-septate, 25–35 ×5–7 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. Graphis slendrae is most closely related to G. dupaxana, with which it shares lirellae with completely carbonized excipulum and striate labia, as well as small, transversely septate ascospores and lack of secondary substances. However, the sessile, very long and radiately branched lirellae of G. slendrae look very different from the prominent, comparatively short and sparsely branched lirellae of G. dupaxana.

Graphis subalbostriata Lücking nom. nov

Graphis angustata var. albostriata Vain., Ann. Acad. Sci. Fenn., Ser. A., 15(6): 158 (1915); non Graphis albostriata Vain., Acta Soc. Fauna Fl. Fenn. 7: 103 (1890) ; typus Guadeloupe, Duss 517 (TUR-Vainio 27847!—holotypus).

Graphis subintermedians Hale ex Lücking sp. nov

Sicut Graphis pseudoserpens sed ascosporis majoribus differt; type Malaysia (Borneo: Sabah), Hale 29240 (US!—holotype).

(Fig. 14E)

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae erumpent (to prominent), with basal thalline margin, short and sparsely branched, 1–3 × 0·3–0·4 mm; labia striate. Excipulum laterally carbonized; hymenium clear. Ascospores 2 per ascus, muriform, 40–75 × 20–30 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. This new species belongs in a small group of taxa with laterally carbonized excipulum, striate lirellae, clear hymenium, and muriform ascospores. Within this group, it is the only species lacking secondary substances and with ascospores exceeding 60 × 16 μm.

Graphis subradiata (Nyl.) Lücking comb. et stat. nov

Graphis analoga var. subradiata Nyl., Acta Soc. Sci. Fenn. 7: 465 (1863); Nylander, Ann. Sci. Nat., Ser. 4, 19: 360 (1863); Graphina analoga var. subradiata (Nyl.) Zahlbr., Cat. Lich. Univ. 2: 395 (1923); type: Colombia, Lindig 2725 (H-Nylander 7434!—holotype).

Graphis subserpens Staiger sp. nov

Sicut Graphis bettinae sed ascosporis latioribus differt.

Typus: Brazil, Kalb 31118 (hb. Kalb!—holotypus).

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae prominent, with thick complete thalline margin, short and sparsely branched, 1–3 × 0·6–0·8 mm; labia entire. Excipulum completely carbonized; hymenium inspersed. Ascospores 4–8 per ascus, 9–19-septate, 80–120 × 20–30 μm, hyaline.

Secondary chemistry. No substances detected.

Notes. This species was treated as nomen nudum in Staiger (Reference Staiger2002); it represents a distinct taxon, distinguished from the closely related G. bettinae by the much broader ascospores, and hence is validated here.

Graphis subtecta (Nyl.) Lücking comb. et stat. nov

Graphis analoga var. subtecta Nyl., Acta Soc. Sci. Fenn. 7: 465 (1863); type: Colombia, Lindig 2725 (H-Nylander 7408b!—holotype).

Notes. As can be seen from the key, this taxon has very little in common with G. analoga.

Graphis subvernicosa Lücking nom. nov

Opegrapha vernicosa Fée, Suppl. Essai Crypt. Écorc.: 24 (1837) [non Graphis vernicosa Nyl., Acta Soc. Sci. Fenn. 7: 466 (1863)]; Graphina vernicosa (Fée) Müll. Arg., Flora 63: 41 (1880); Müller Argoviensis, Mém. Soc. Phys. Hist. Nat. Genève 29(8): 41 (1887); type: America, s.col. (G—holotype, not seen).

Graphis sulphurella (Zahlbr.) Lücking comb. nov

Graphina sulphurella Zahlbr., Ann. Mycol. 10: 366 (1912); type: Hawaii, Rock 135 (W!—holotype).

Graphis syzygii Aptroot sp. nov

Sicut Graphis stenotera sed excipulo lateraliter et basaliter carbonisato et hymenio insperso differt.

Typus: Costa Rica, Guanacaste, Volcán Tenerio National Park, 25 km NNW of Tilarán, along access road to station near Bijagua, on Syzygium tree in meadow, 700 m, March 2004, Aptroot 60569 (INB!—holotypus; B!—isotypus).

(Fig. 14F)

Thallus corticate, smooth to uneven, white to pale grey.

Lirellae erumpent, without or with basal thalline margin, elongate and irregularly branched, 1–5 × 0·2–0·3 mm; labia striate. Excipulum completely carbonized; hymenium inspersed. Ascospores 8 per ascus, transversely 9–13-septate, 25–35 × 5–7 μm, hyaline.

Secondary chemistry. Stictic acid.

Notes. This species closely resembles Graphis stenotera morphologically and chemically but differs in the completely carbonized excipulum and inspersed hymenium. The few other species with striate labia, completely carbonized excipula, inspersed hymenia, and transversely septate ascospores, viz. G. cinerea, G. leucaenae, and G. inspersolongula, have much larger ascospores (75–200 × 15–30 μm).