Introduction
Molecular phylogenies have dramatically changed the understanding of generic classification within Graphidaceae, the largest family of tropical crustose lichens (Mangold et al. Reference Mangold, Martín, Lücking and Lumbsch2008; Rivas Plata & Lumbsch Reference Rivas Plata and Lumbsch2011; Rivas Plata et al. Reference Rivas Plata, Lumbsch and Lücking2012; Lücking et al. Reference Lücking, Johnston, Aptroot, Kraichak, Lendemer, Boonpragob, Cáceres, Ertz, Ferraro and Jia2014). Quantitative extrapolations and molecular data for species complexes in tandem with new fieldwork in underexplored areas of the tropics suggest that the family Graphidaceae may contain up to 3500 species, that is 1400 undescribed species in addition to the 2100 currently known (Lücking et al. Reference Lücking, Johnston, Aptroot, Kraichak, Lendemer, Boonpragob, Cáceres, Ertz, Ferraro and Jia2014; Joshi et al. Reference Joshi, Upreti, Divakar, Lumbsch and Lücking2018). The family has already been treated in several studies focusing on material from Sri Lanka, revealing a large number of taxa new to science (Awasthi & Singh Reference Awasthi and Singh1978; Singh & Awasthi Reference Singh and Awasthi1979; Hale Reference Hale1981; Nagarkar & Hale Reference Nagarkar and Hale1989; Breuss & Brunnbauer Reference Breuss and Brunnbauer1997; Papong et al. Reference Papong, Lücking, Thammathaworn and Boonpragob2009; Weerakoon et al. Reference Weerakoon, Rivas Plata, Lumbsch and Lücking2012a, Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbschb, Reference Weerakoon, Lücking and Lumbsch2014, Reference Weerakoon, Jayalal, Wijesundara, Karunaratne and Lücking2015; Rivas Plata et al. Reference Rivas Plata, Parnmen, Staiger, Mangold, Frisch, Weerakoon, Hernández, Cáceres, Kalb and Sipman2013; Li et al. Reference Li, Hyde, Zhao, Hongsanan, Abdel-Aziz, Abdel-Wahab, Alvarado, Alves-Silva, Ammirati and Ariyawansa2016; Jatnika et al. Reference Jatnika, Weerakoon, Arachchige, Noer, Voytsekhovich and Lücking2019).
A recent analysis of new collections from Sri Lanka found c. 250 species including c. 170 new records and led to the discovery of c. 30 new species in Graphidaceae. The most speciose group was Graphis sensu Staiger (Reference Staiger2002) with over 100 taxa, including six new taxa described by Weerakoon et al. (Reference Weerakoon, Rivas Plata, Lumbsch and Lücking2012a, Reference Weerakoon, Wijeyaratne, Wolseley, Rivas Plata, Lücking and Lumbschb). Graphis sensu Staiger was recently divided into the two genera Graphis s. str. and Allographa (Kalb et al. Reference Kalb, Lücking and Kalb2018; Lücking & Kalb Reference Lücking and Kalb2018). Therefore, in the present paper, in addition to describing six species new to science, we also provide an updated key to all species known from Sri Lanka in the two genera, with the taxa redispositioned into Graphis s. str. and Allographa. As noted by Lücking & Kalb (Reference Lücking and Kalb2018), it is currently not possible to readily separate Graphis and Allographa at the genus level phenotypically, despite the two genera being very distantly related. The bulk of the species in both genera differ in lirellae morphology (delicate vs. robust), excipulum development (thin vs. massive), hamathecium inspersion when present (type A vs. type B), and the often anastomosing lateral paraphyses in Allographa; however, in both genera there are a number of exceptions to these rules, particularly concerning species with black, striate labia. Therefore, keying out the taxa at species level is necessary.
In addition, an attempt is made to place the richness and composition of Graphis and Allographa in Sri Lanka into context, discussing a possible signature of the ‘biotic ferry’ species interchange, by a comparison with two other well-sampled countries, namely Costa Rica in the Neotropics and Thailand in the eastern Paleotropics.
Materials and Methods
Specimens were examined using Olympus SZX7, Leica MS5 and Motic K400 dissecting microscopes and Olympus BX50 with Nomarski, Zeiss Axioscop 2 and VistaVision VWR V036 compound microscopes, in part connected to Nikon Coolpix E995, Jenoptic ProgRes C3 and C5 digital microscope cameras. Specimens are deposited at BM and PDA. Anatomical measurements refer to specimens mounted in water; for iodine reactions, we used Fluka 62650 Lugol's solution. We performed thin-layer chromatography (TLC) using solvent C (Orange et al. Reference Orange, James and White2010). Named lirellae-types follow Lücking et al. (Reference Lücking, Archer and Aptroot2009) and Kalb et al. (Reference Kalb, Lücking and Kalb2018).
We compared the species inventories of Graphis and Allographa for four countries, two in the Neotropics (Costa Rica, Mexico) and two in the eastern Paleotropics (Sri Lanka, Thailand). To assess compositional similarity, we computed pairwise Sørensen similarity values between all four countries in PC-Ord 6.0 (McCune & Mefford Reference McCune and Mefford2011). To statistically compare species sampling between Sri Lanka and Thailand and its effect on the Sørensen metrics, we performed 50% jackknifing with 1000 repeats in each case, and for each repeat we computed the Sørensen similarity value with the full sample from Costa Rica by multiplying the subsample result by two and randomly correcting by 0, +1 or –1 to avoid exclusively even numbers.
Results and Discussion
The total number of species of Allographa and Graphis now recorded for Sri Lanka is 124, thus surpassing the previously reported numbers for Costa Rica (115; Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008), Thailand (115; Kalb et al. Reference Kalb, Lücking and Kalb2018) and Mexico (71; Wirth & Hale Reference Wirth and Hale1963, specimens in US revised by R. Lücking; Bárcenas-Peña et al. Reference Barcenas Peña, Lücking, Miranda-González and Herrera-Campos2014; Fig. 1). This is remarkable as Sri Lanka covers an area of only 65 000 km2, compared to Thailand with 513 000 km2 (almost eight times the size). On the other hand, the area of Costa Rica (51 000 km2) is comparable with that of Sri Lanka and species density is about the same, with between 1·9 (Sri Lanka) and 2·3 (Costa Rica) species per 1000 km2, whereas Thailand has 0·22 species per 1000 km2. In addition to area, these differences might be explained partly by topography, especially as most species of Graphis and Allographa favour mid elevations. The effect of area can be roughly balanced by log-transformed area values, since species richness relates logarithmically to area (Lücking et al. Reference Lücking, Seavey, Common, Beeching, Breuss, Buck, Crane, Hodges, Hodkinson and Lay2011). Thus, a better estimate is richness per logarea, which results in 26 for Sri Lanka, 24 for Costa Rica, 20 for Thailand and 11 for Mexico. It can be assumed that Costa Rica and Sri Lanka are comparatively well sampled, whereas sampling effort in Thailand is lower and for Mexico it is still quite low. To obtain similar species per logarea scores as Costa Rica and Sri Lanka, the total richness for Thailand would have to amount to 140 species, and for Mexico to 160 species, which implies that currently 80% of the expected species richness in these two genera is known from Thailand, and only little more than 40% from Mexico. The only country with a higher number of reported names in Graphis and Allographa is Brazil, but no updated inventory is yet available for that country.
Fig. 1. Comparison of the richness and composition of Graphis and Allographa species from Sri Lanka with Costa Rica and Mexico in the Neotropics and Thailand in the eastern Paleotropics. Numbers in parentheses indicate the total recorded species of Allographa plus Graphis for each country. Increased line thickness reflects greater pairwise Sørensen similarity values for the two countries being compared, with the exact values reported next to each line. In colour online.
A pairwise comparison between species composition of the four countries yielded Sørensen similarity values of 37% between Mexico and Costa Rica (Neotropics) and 44% between Thailand and Sri Lanka (eastern Paleotropics), the difference probably explained by the low sampling effort in Mexico. Cross-comparison between the neotropical and paleotropical countries showed notable differences: between Costa Rica (or Mexico) and Thailand, similarity amounted to 29 and 31%, respectively, whereas between Costa Rica (or Mexico) and Sri Lanka, it ranged between 38% and 41%, c. 10% higher in both cases (Fig. 1). Since all four inventories are based on the same modern species concept and are largely assembled by the same group of taxonomists, this difference cannot be considered a bias of taxonomic concept. Furthermore, randomized subsampling of species in Sri Lanka and Thailand demonstrated a highly significant difference in the central tendency of the similarity values of both countries with Costa Rica (Fig. 2; Mann-Whitney U test, P = 0·001). The individually observed value for Thailand (0·29) was outside the 99·9% range obtained for Sri Lanka (Fig. 2; P = 0·001), whereas the individually observed value for Sri Lanka (0·41) was outside the 99·6% range obtained for Thailand (Fig. 2; P = 0·01).
Fig. 2. Random subsampling of the species composition of Allographa and Graphis in Thailand and Sri Lanka, with 1000 repeats and subsequent comparison of each repeat to a random subsample of Costa Rica. The resulting Sørensen pairwise similarity values with Costa Rica exhibit significant differences between Thailand and Sri Lanka. The observed central tendency values are indicated for each country. In colour online.
One explanation for the higher similarity of Sri Lanka (as compared to Thailand) with Costa Rica is that this is a signature of the so-called ‘biotic ferry’ species interchange. Sri Lanka being compared forms part of the Indian geological block, which until 160 mya was connected to Gondwana. After the break-up of Gondwana this block drifted slowly across the Tethys and eventually collided with the Eurasian Plate around 55 mya, carrying both Gondwana and endemic elements and thus acting as a ‘biotic ferry’ (McKenna Reference McKenna, Tarling and Runcorn1973; Ashton & Gunatilleke Reference Ashton and Gunatilleke1987; Macey et al. Reference Macey, Schulte, Larson, Ananjeva, Wang, Pethiyagoda, Rastegar-Pouyani and Papenfuss2000; Morley Reference Morley2000; Rust et al. Reference Rust, Singh, Rana, McCann, Singh, Anderson, Sarkar, Nascimbene, Stebner and Thomas2010). For this hypothesis to hold, species composition in areas of tropical Africa should be similar to those in the Neotropics, which is currently not possible to assess due to the lack of data from tropical Africa. However, a major contradiction to that hypothesis would be the assumed age of the species, at over 100 my. According to various molecular clock studies, the genera Graphis and Allographa did not diversify until c. 30–50 mya, and the age of most species is probably not greater than 5–10 my (Lücking et al. Reference Lücking, Tehler, Bungartz, Rivas Plata and Lumbsch2013; Kraichak et al. Reference Kraichak, Divakar, Crespo, Leavitt, Nelsen, Lücking and Lumbsch2015), meaning that the genera and their species originated long after the Gondwana break-up and even after the collision of the Indian subcontinent with the Eurasian plate. Thus, even if one assumes that taxa currently identified as ‘species’ in these inventories represent species complexes, the timeline to postulate a Gondwana origin of selected lineages present in both the Neotropics and Sri Lanka would be wrong. A more likely explanation would be geologically recent mid- to long-distance dispersal facilitated by paleoclimatic conditions combined with the relative proximity of India (and Sri Lanka) to the African Paleotropics, as compared to other parts of tropical Asia. This has, for example, been shown for Melastomataceae, with evidence of multiple dispersal between Africa, Madagascar and India during the Miocene, 23–5 mya (Renner Reference Renner2004).
Certainly, to further elucidate this intriguing issue, good sampling across all tropics and a solid phylogeny are needed for these and other lichen genera with similar distribution and richness patterns.
Key to Allographa and Graphis in Sri Lanka
Key to species groups
1 Hamathecium inspersed ………2
Hamathecium not inspersed ………4
2(1) Ascospores muriform ………Group A
Ascospores only transversely septate ………3
3(2) Thallus K− ………Group B
Thallus K+ yellow > red or K+ red ………Group C
4(1) Thallus UV+ yellow (lichexanthone), hamathecium not inspersed ………Group D
Thallus UV−, hamathecium variable ………5
5(4) Ascospores muriform (at least terminally) ………6
Ascospores only transversely septate ………7
6(5) Thallus K+ yellow or yellow > red ………Group E
Thallus K− ………Group F
7(5) Thallus K+ yellow, in section forming a persistently yellow efflux (stictic acid)……… ………Group G
Thallus K+ yellow > red, K+ red or K− ………8
8(7) Thallus K+ yellow > red or K+ red, in section forming a yellow efflux and subsequently either conspicuous, red, needle-shaped crystals (norstictic acid) or very small, red crystals (salazinic acid) ………Group H
Thallus K− ………9
9(8) Labia becoming striate ………Group I
Labia entire ………Group J
Group A. Hamathecium inspersed, ascospores muriform
1 Thallus K− ………2
Thallus K+ yellow or yellow > red ………4
2(1) Ascospores 12–17 µm wide; lirellae with apically thick complete thalline cover ……… ……… Allographa elixii
Ascospores 15–35 µm wide; lirellae with apically thin complete thalline cover ………3
3(2) Labia becoming striate ………Allographa argentata
Labia entire ………Allographa phaeospora
4(1) Thallus K+ yellow (stictic acid); ascospores 50–130 × 15–25 µm……… ………Graphis diplocheila
Thallus K+ yellow > red (norstictic acid); ascospores variable ………5
5(4) Ascospores 8 per ascus, 25–35 × 7–8 µm ………Allographa knucklensis
Ascospores 1–2 per ascus, >60 µm long ………6
6(5) Excipulum completely carbonized ………7
Excipulum laterally carbonized ………8
7(6) Ascospores 1 per ascus, 100–145 × 25–40 µm; lirellae erumpent, elongate and irregularly branched (subserpentina morph) ………Graphis novopalmicola
Ascospores 1–2 per ascus, 60–110 × 16–30 µm; lirellae prominent, short and unbranched (dussii morph) ………Allographa leprographa
8(6) Marginal lirellae elongate, dispositioned radially and typically curved, central lirellae short and straight; thalline margin of lirellae thin, labia appearing roughened and not sharply delimited from thalline margin ………Graphis flosculifera
Lirellae irregularly arranged, without marked difference between marginal and central lirellae and central lirellae usually longer; thalline margin of lirellae thick, labia largely covered or when visible, rather well delimited from thalline margin ……… ………Graphis insulana
Group B. Hamathecium inspersed, ascospores transversely septate, thallus K−
1 Lirellae prominent, thalline margin absent or basal ………2
Lirellae erumpent, thalline margin lateral ………3
2(1) Ascospores 25–45 × 6–9 µm ………Graphis anfractuosa
Ascospores 60–70 × 7–9 µm ………Graphis cupei
3(1) Ascospores 13–15-septate, 40–55 × 9–12 µm ………Graphis intermedians
Ascospores 5–9-septate, up to 40 µm long ………4
4(3) Disc exposed, white pruinose ………Graphis submarginata
Disc concealed ………5
5(4) Lirellae radiately branched, up to 10 mm long ………Graphis arbusculiformis
Lirellae sparsely branched, up to 3 mm long ………Graphis lineola
Group C. Hamathecium inspersed, ascospores transversely septate, thallus K+ yellow or yellow > red
1 Thallus K+ yellow, in section forming a persistently yellow efflux (stictic acid) ………2
Thallus K+ yellow > red, in section forming a yellow efflux and subsequently conspicuous, red, needle-shaped crystals (norstictic acid) ………4
2(1) Excipulum completely carbonized; lirellae with complete thalline cover……… ………Graphis luluensis
Excipulum laterally carbonized; lirellae with lateral thalline margin ………3
3(2) Ascospores 40–60 × 10–14 µm ………Graphis kelungana
Ascospores 20–40 × 6–8 µm ………Graphis leptocarpa
4(1) Labia becoming striate, with thin complete thalline cover ………Allographa leucaenae
Labia entire, with basal or lateral thalline margin ………5
5(4) Ascospores 50–70 × 8–10 µm; thalline margin lateral ………Graphis mikuraensis
(syn.: G. srilankensis Weerakoon et al.)
Ascospores <50 µm long; thalline margin variable 6
6(5) Excipulum laterally carbonized; thalline margin lateral ………7
Excipulum completely carbonized; thalline margin variable ………8
7(6) Disc exposed, white pruinose ………Graphis handelii
Disc concealed ………Graphis cincta
8(6) Thalline margin absent or at most basal ………Graphis desquamescens
Thalline margin lateral ………9
9(8) Ascospores 11–13-septate, 35–50 × 7–8 µm ………Graphis rajapakshana
Ascospores 5–7-septate, 20–35 × 5–9 µm ………Graphis gonimica
Group D. Hamathecium not inspersed, thallus UV+ yellow (lichexanthone)
1 Ascospores 15–20 × 5–8 µm; excipulum laterally carbonized; thallus K+ yellow > red (norstictic acid in addition to lichexanthone) ………Graphis stipitata
Ascospores 45–60 × 10–14 µm; excipulum completely carbonized; thallus K− (no other substances) ………Allographa sauroidea
Group E. Hamathecium not inspersed, ascospores muriform, thallus UV−, K+ yellow or yellow > red
1 Thallus K+ yellow, in section forming a persistently yellow efflux (stictic acid) ………2
Thallus K+ yellow > red, in section forming a yellow efflux and subsequently conspicuous, red, needle-shaped crystals (norstictic acid) ………3
2(1) Ascospores 2–4 per ascus, 50–70 × 18–25 µm ………Graphis japonica
Ascospores 1 per ascus, 70–95 × 20–30 µm ………Graphis streblocarpa
3(1) Ascospores 8 per ascus, 25–35 × 9–12 µm ………Graphis renschiana
Ascospores 1 per ascus, >80 µm long ………4
4(3) Lirellae elongate and irregularly branched ………Graphis subserpentina
Lirellae short and unbranched ………Graphis hiascens
Group F. Hamathecium not inspersed, ascospores muriform, thallus K− and UV−
1 Ascospores up to 70 µm long ………2
Ascospores generally >70 µm long ………8
2(1) Lirellae short and unbranched, sessile and lacking thalline cover (nuda-morph) ………3
Lirellae at least partly branched, erumpent (to prominent), with lateral to apically thin complete thalline margin ………4
3(2) Ascospores 4–8 per ascus, 35–65 × 10–20 µm ………Allographa ruiziana
Ascospores 8 per ascus, 25–40 × 10–20 µm ………Allographa nuda
4(2) Labia becoming striate; excipulum apically carbonized ………5
Labia entire; excipulum laterally or completely carbonized ………7
5(4) Ascospores 1–2 per ascus, >16 µm wide; thalline margin lateral ………Graphis disserpens
Ascospores 2–8 per ascus, <16 µm wide; thalline margin variable ………6
6(5) Ascospores 2–6 per ascus; thalline margin complete but apically thin ……… ………Graphis paraserpens
Ascospores 8 per ascus; thalline margin lateral ………Graphis galactoderma
7(4) Ascospores 20–40 × 10–12 µm; thalline margin complete but apically thin.……… ………Graphis subtecta
Ascospores 50–70 × 15–22 µm; thalline margin lateral ………Graphis lapidicola
8(1) Ascospores only terminally muriform ………9
Ascospores muriform throughout ………10
9(8) Lirellae with yellow-orange pruina ………Allographa jayatilakana
Lirellae lacking pigmented pruina ………Allographa vestitoides
10(8) Excipulum laterally carbonized ………11
Excipulum completely carbonized ………13
11(10) Ascospores 2–4 per ascus; thallus thinly verrucose ………Allographa bambusicola
Ascospores 1 per ascus; thallus not verrucose ………12
12(11) Thalline margin complete ………Graphis subhiascens
Thalline margin basal ………Allographa mahaeliyensis
13(10) Labia becoming striate ………14
Labia entire ………17
14(13) Ascospores 2–6 per ascus ………Allographa acharii
Ascospores 1–2 per ascus ………15
15(14) Ascospores 1–2 per ascus; thalline margin lateral ………Graphis myrtacea
Ascospores 1 per ascus; thalline margin complete, although apically usually thin ………16
16(15) Lirellae prominent ………Allographa macella
Lirellae immersed ………Allographa consanguinea
17(13) Ascospores 2–4 per ascus; lirellae immersed ………Graphis dolichographa
Ascospores 1 per ascus; lirellae prominent to sessile ………18
18(17) Thalline margin lateral, conspicuously thickened; lirellae sessile, very short ……… ………Allographa plagiocarpa
Thalline margin complete; lirellae prominent, distinctly elongate ……… ………Allographa illinata
Group G. Hamathecium not inspersed, ascospores transversely septate, thallus K+ yellow (stictic acid)
1 Thallus with isidia; lirellae with complete thalline cover ………Graphis isidiza
Thallus without isidia; lirellae with basal or lateral thalline cover ………2
2(1) Disc exposed, often pruinose ………3
Disc closed ………4
3(2) Ascospores 60–100 × 8–12 µm ………Graphis longiramea
Ascospores 25–45 × 6–10 µm ………Graphis sundarbanensis
4(2) Labia white pruinose; lirellae usually radiating ………Graphis dendrogramma
Labia not pruinose; lirellae variable ………5
5(4) Thallus and/or thalline margin verrucose ………6
Thallus and thalline margin not verrucose ………8
6(5) Ascospores 35–40 × 7–8 µm ………Graphis allugallenensis
Ascospores longer ………7
7(6) Thalline margin verrucose; ascospores 15–21-septate ……… ………Allographa weerasooriyana
Thalline margin not verrucose; ascospores 11–17-septate ………Allographa rustica
8(5) Labia becoming striate ………9
Labia remaining entire ………11
9(8) Carbonization lateral; thalline margin basal; duplicata-morph ……… ………Graphis brahmanensis
Carbonization usually only apical; thalline margin lateral or absent; tenella- or striatula-morph ………10
10(9) Thalline margin absent ………Graphis stenotera
Thalline margin lateral ………Graphis vittata
11(8) Ascospores usually >40 µm ………12
Ascospores usually <40 µm ………13
12(11) Excipulum laterally to completely carbonized; thalline margin lateral, thick ……… ………Graphis crassilabra
Excipulum only laterally carbonized; thalline margin lateral, thin ……… ………Graphis leptogramma
13(11) Excipulum completely carbonized; thalline margin lateral, thick ……… ………Graphis descissa
Excipulum only laterally carbonized; thalline margin lateral, thin ……… ………Graphis immersella
Group H. Hamathecium not inspersed, ascospores transversely septate, thallus K+ yellow > red (norstictic acid) or K+ red (salazinic acid)
1 Thallus K+ yellow then quickly red, in section forming a yellow efflux and rapidly very small, red crystals (salazinic acid) ………2
Thallus K+ yellow then slowly red, in section forming a yellow efflux and subsequently conspicuous, red, needle-shaped crystals (norstictic acid) ………4
2(1) Labia orange; disc exposed, ochraceous ………Graphis ferruginea
Labia not pigmented; disc concealed ………3
3(2) Ascospores 5–7-septate; only salazinic acid present (TLC) ………Graphis bakeri
Ascospores 7–9-septate; protocetraric acid also present (TLC) ………Graphis litoralis
4(1) Thalline margin absent; ascospores thickly double-walled ………Graphis elegans
Thalline margin present; ascospores not double-walled ………5
5(4) Disc exposed, white pruinose ………Graphis pyrrhocheiloides
Disc concealed ………6
6(5) Labia white pruinose ………7
Labia not pruinose ………8
7(6) Excipulum completely carbonized; labia with thin complete thalline margin ……… ………Graphis caesiocarpa
Excipulum laterally carbonized; labia with lateral thalline margin ………Graphis caesiella
8(6) Ascospores 20–30 × 5–8 µm ………Graphis librata
Ascospores >50 µm 9
9(8) Labia becoming striate, with thin complete thalline margin ………Allographa leptospora
Labia entire, with lateral thalline margin ………Allographa marginata
Group I. Thallus K− and UV−, hamathecium not inspersed, ascospores transversely septate, labia becoming striate
1 Thalline margin absent to basal ………2
Thalline margin complete to lateral ………15
2(1) Thalline margin absent ………3
Thalline margin basal ………9
3(2) Ascospores mostly >50 µm long ………4
Ascospores never >50 µm long ………5
4(3) Excipulum laterally carbonized ………Allographa striatula
Excipulum apically carbonized ………Graphis caribica
5(3) Excipulum apically to laterally carbonized ………6
Excipulum completely carbonized ………8
6(5) Excipulum laterally carbonized; lirellae erumpent ………Graphis duplicata
Excipulum apically carbonized; lirellae prominent ………7
7(6) Lirellae unbranched; ascospores 5–7-septate ………Graphis granulocarpa
Lirellae branched; ascospores 7–11-septate ………Graphis tsunodae
8(5) Lirellae erumpent ………Allographa rimulosa
Lirellae prominent ………Graphis dupaxana
9(2) Ascospores >50 µm long ………10
Ascospores never >40 µm long ………12
10(9) Ascospores 50–110 × 4–8 µm; excipulum laterally carbonized ……… ………Allographa polystriata
Ascospores broader, 7–12 µm wide; excipulum apically carbonized ………11
11(10) Ascospores 60–100 × 10–12 µm ………Graphis dotalugalensis
Ascospores 50–80 × 7–10 µm ………Graphis thunsinhalayensis
12(9) Excipulum completely carbonized ………Graphis slendrae
Excipulum apically to laterally carbonized ………13
13(12) Lirellae sparsely branched (tenella-morph) ………Graphis chlorotica
Lirellae irregularly branched (striatula-morph) ………14
14(13) Thallus verrucose ………Graphis verrucoserpens
Thallus smooth to uneven ………Graphis proserpens
15(1) Thalline margin complete ………16
Thalline margin lateral ………19
16(15) Lirellae immersed; thallus rimose, Fissurina-like ………Graphis rimosothallina
Lirellae erumpent to prominent; thallus smooth to uneven, Graphis-like ………17
17(16) Ascospores <50 µm long; excipulum laterally carbonized ………Allographa ingarum
Ascospores mostly >50 µm long; excipulum completely carbonized ………18
18(17) Lirellae prominent; ascospores 70–120 × 12–18 µm ………Allographa angustata
Lirellae erumpent; ascospores 50–80 × 8–12 µm ………Allographa glauconigra
19(15) Ascospores mostly >40 µm long ………20
Ascospores <40 µm long ………23
20(19) Thallus verrucose; lirellae prominent, more or less stellately branched; excipulum completely carbonized ………Allographa asterizans
Thallus smooth; lirellae erumpent, irregularly branched; excipulum variable ………21
21(20) Ascospores 80–125 × 8–12 µm; excipulum apically carbonized ……… ………Graphis subalbostriata
Ascospores <70 µm long ………22
22(21) Ascospores 50–70 × 10–13 µm; excipulum completely carbonized ……… ………Allographa longula
Ascospores 30–50 × 7–10 µm; excipulum laterally carbonized ………Graphis leptoclada
23(19) Labia white pruinose; excipulum apically carbonized ………24
Labia not pruinose; excipulum variable ………25
24(23) Thallus ecorticate ………Graphis glaucescens
Thallus corticate ………Graphis chloroalba
25(23) Excipulum apically carbonized; ascospores >30 µm ………Graphis subtenella
Excipulum laterally or completely carbonized; ascospores <30 µm ………26
26(25) Excipulum laterally carbonized ………Graphis tenella
Excipulum completely carbonized ………Graphis aurita
Group J. Thallus K− and UV−, hamathecium not inspersed, ascospores transversely septate, labia entire
1 Ascospores at least partly >45 µm long ………2
Ascospores never >45 µm long ………11
2(1) Ascospores at least partly >80 µm long ………3
Ascospores never >80 µm long ………5
3(2) Excipulum laterally carbonized; thalline margin basal ………Allographa hossei
Excipulum completely carbonized; thalline margin complete or absent ………4
4(3) Thalline margin absent; lirellae very short and unbranched (nuda type) ……… ………Allographa bifera
Thalline margin complete; lirellae elongate and branched (rhizicola type) ……… ………Allographa rhizicola
5(2) Labia or disc white pruinose ………6
Labia nor disc pruinose; disc concealed ………8
6(5) Disc open, pruinose ………Graphis scripta s. lat.
Disc concealed ………7
7(6) Thallus ecorticate; excipulum laterally carbonized ………Allographa pavoniana
Thallus corticate; excipulum apically carbonized ………Allographa sayeri
8(5) Thalline margin absent; lirellae very short and unbranched (nuda type). ……… ………Allographa adpressa
Thalline margin complete to lateral; lirellae elongate and branched ………9
9(8) Thalline margin complete ………Allographa anguilliradians
Thalline margin lateral ………10
10(9) Excipulum laterally carbonized ………Allographa elongata
Excipulum completely carbonized ………Allographa flavens
11(1) Thalline margin absent; lirellae very short and mostly unbranched ………12
Thalline margin present; lirellae branched ………13
12(11) Ascospores 15–25 × 5–8 µm; excipulum laterally carbonized ………Allographa nana
Ascospores 25–45 × 7–13 µm; excipulum completely carbonized ……… ………Graphis conferta
13(11) Excipulum apically or laterally carbonized ………14
Excipulum completely carbonized ………17
14(13) Excipulum apically carbonized ………15
Excipulum laterally carbonized ………16
15(14) Ascospores 30–40 × 8–10 µm; labia white pruinose ………Allographa xanthospora
Ascospores 25–35 × 6–8 µm; labia not pruinose ………Graphis oligospora
16(14) Thallus partly ecorticate; lirellae partly white pruinose ………Graphis furcata
Thallus corticate; lirellae not pruinose ………Graphis pinicola
17(13) Thallus ecorticate; lirellae white pruinose ………Allographa sitiana
Thallus corticate; lirellae not pruinose ………18
18(17) Ascospores 20–30 × 5–8 µm ………19
Ascospores 30–45 × 7–9 µm ………20
19(18) Thalline margin basal; lirellae usually stellately branched ………Graphis dracaenae
Thalline margin lateral; lirellae irregularly branched ………Graphis immersicans
20(18) Thalline margin complete, apically thin ………Graphis negrosina
Thalline margin lateral and thickened ………Graphis oxyclada
The New Species
Allographa bambusicola Weerakoon, Lücking & Aptroot sp. nov.
MycoBank No.: MB 832538
Bambusicolous Allographa with entire labia, laterally carbonized excipulum, 80–100 × 15–17 µm large, muriform ascospores and rather thick, irregularly verrucose lateral thalline margin of the lirellae.
Type: Sri Lanka, Central Province, Galhiriya Estate, 5 km NNW of Hunasgiriya, 7°26′N, 80°42′E, 1241 m, high altitude, cardamom plantation, December 1989, Cloonan 56 (PDA—holotype).
(Fig. 3A & B)
Thallus bambusicolous, up to 5 cm diam., continuous; surface thinly verrucose, creamish to light green-grey; prothallus not observed. Thallus in section 80–120 µm thick, with cortex 10–20 µm thick, photobiont layer 20–30 µm thick, and medulla 50–70 µm thick, containing large clusters of calcium oxalate crystals.
Lirellae more or less straight, short, unbranched, prominent, rhizicola-type, with a rather thick, protruding, irregularly verrucose lateral thalline margin, 0·5–1·0 mm long, 0·2–0·3 mm wide, 0·25–0·30 mm high; disc concealed, labia entire. Excipulum laterally carbonized, 50–70 µm wide; hypothecium prosoplectenchymatous, 10–15 µm high, pale olive; hymenium 150–180 µm high, colourless, not inspersed; epithecium granulose, 5–10 µm high, brownish. Paraphyses unbranched; asci fusiform, 150–160 × 25–30 µm. Ascospores 2–4 per ascus, fusiform to oblong, muriform, 80–100 × 15–17 µm, 5–6 times as long as wide, colourless, I+ amyloid.
Fig. 3. Habitus and ascospores of Allographa and Graphis species, all holotypes. A & B, A. bambusicola; C, A. weerasooriyana; D–F, G. flosculifera. Scales: A, C, D & E = 1 mm; B & F = 20 µm. In colour online.
Secondary chemistry
No substances detected by TLC.
Etymology
The name refers to the bambusicolous growth.
Distribution and habitat
The new species was collected in montane forest in the central region of Sri Lanka. It is thus far known only from the type locality.
Discussion
This species belongs to a group of taxa with entire labia, a laterally carbonized excipulum and large, muriform ascospores. Allographa atrocelata (A. W. Archer) Lücking & Kalb is similar in lirella morphology, except that the thalline margin is smooth, but its ascospores are much larger (100–140 × 30–40 µm). Graphis xylophaga (R. C. Harris) Lendemer has erumpent lirellae with a smooth, lateral thalline margin with apically jet black labia and distinctly broader ascospores (20–30 µm), and G. lapidicola Fée also differs in the erumpent lirellae, with an apically rather thick, smooth complete margin and relatively broader ascospores (3–4 times as long as wide).
Allographa weerasooriyana Weerakoon, Arachchige & Lücking sp. nov.
MycoBank No.: MB 832539
Corticolous Allographa resembling A. rustica Kremp. in overall anatomy and chemistry, but with verrucose thalline margins of the lirellae, and labia not distinctly raised above the thalline margin.
Type: Sri Lanka, Sabaragamuwa hill range, 10 km from Yathiyanthota, Halgolla Tea Estate, 7°02′N, 80°23′E, 1241 m, high altitude, tea estate, July 1988, Cloonan 686 (PDA—holotype).
(Fig. 3C)
Thallus corticolous, up to 10 cm diam., continuous; surface marmorate and coarsely verrucose, light yellowish grey. Thallus in section 100–150 µm thick, with prosoplectenchymatous cortex 10–15 µm thick, irregular algal layer 30–50 µm thick, intermingled with and partially covered by large clusters of calcium oxalate crystals, and medulla 50–80 µm thick, filled with numerous grey crystals.
Lirellae flexuose, irregularly branched, erumpent to prominent, marginata-type, with protruding, irregularly verrucose, rather thick lateral thalline margins but with the labia not distinctly raised, 3–10 mm long, 0·4–0·6 mm wide, 0·30–0·35 mm high; disc concealed, labia entire. Excipulum completely carbonized, 50–120 µm wide; hypothecium prosoplectenchymatous, 20–30 µm high, pale yellowish; hymenium 150–170 µm high, colourless, not inspersed; epithecium indistinct. Paraphyses unbranched; asci fusiform, 120–140 × 20–30 µm. Ascospores (mostly young in the type, with few asci with mature ascospores observed) 4–8 per ascus, oblong, 15–21-septate, 70–90 × 10 µm, 7–9 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
Stictic acid.
Etymology
This new species is named in honour of Aruna Weerasooriya, Professor at PVAMU (Texas A & M University System), a pioneer botanist in Sri Lanka.
Distribution and habitat
The new species was collected in a high elevation tea plantation in the Sabaragamuwa mountain range of Sri Lanka, where it is known only from the type locality.
Discussion
This species most closely resembles Allographa rustica (Kremp.) Lücking & Kalb in overall anatomy and chemistry but differs in the coarsely verrucose to marmorate thallus and the verrucose thalline margin of the lirellae, as well as the upper part of the labia being level with the thalline margin. Graphis schroederi Zahlbr. can be separated by the erumpent lirellae with smooth margins that mostly cover the labia (Lücking et al. Reference Lücking, Archer and Aptroot2009).
Additional specimen examined. Sri Lanka:Sabaragamuwa: mountain range, Coolbone Tea Estate, 7°01′N, 80°28′E, 860 m, mid altitude, 1988, Cloonan 077 (PDA).
Graphis flosculifera Weerakoon, Lücking & Aptroot sp. nov.
MycoBank No.: MB 832540
Corticolous Graphis resembling G. insulana but differing in the unique disposition of the lirellae and the slightly more elongate ascospores.
Type: Sri Lanka, Western Province, Gampaha, 2 km from Divulapitya, 7°30′N, 80°62′E, 130 m, low altitude, coconut plantation, September 1988, on bark of Cocos nucifera, Cloonan 988 (PDA—holotype).
(Fig. 3D–F)
Thallus corticolous, up to 8 cm diam., pruinose, white to ash grey; prothallus not observed. Thallus in section 80–100 µm thick, lacking a cortex and entirely formed by an irregular photobiont layer intermingled with large clusters of calcium oxalate crystals.
Lirellae flexuose, unbranched to very sparsely branched, in the central portion of the thallus very short and irregularly arranged, in the periphery much longer and distinctly radiating, erumpent, subserpentina-type, with lateral thalline margins, 1–5 mm long, 0·3–0·4 mm wide, 0·20–0·25 mm high; disc concealed, labia entire, appearing roughened and not sharply delimited from the rather thin thalline margin. Excipulum laterally carbonized, 50–80 µm wide; hypothecium prosoplectenchymatous, 10–15 µm high, pale olive; hymenium 100–120 µm high, colourless, inspersed (type A sensu Lücking et al. Reference Lücking, Archer and Aptroot2009); epithecium indistinct, olive brown. Paraphyses unbranched; asci clavate, 80–100 × 20–25 µm. Ascospores 1 per ascus, muriform, 80–100 × 20–25 µm, 3·5–4·5 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
Norstictic acid.
Etymology
The name refers to the flower-like arrangement of the lirellae.
Distribution and habitat
The new species was collected in a low elevation coconut plantation in the Western Province of Sri Lanka. It grew on almost every coconut tree in the area of the type locality.
Discussion
This new species is close to Graphis insulana (Müll. Arg.) Lücking and differs mainly in the unique disposition of the lirellae; in G. insulana, the lirellae are typically irregularly arranged and of similar length throughout, as in most species of the genus. Considering the ontogeny of the lirellae, one would usually expect those in the thallus centre to be larger or longer, as they are older, but in G. flosculifera the opposite is the case. In addition, in G. insulana the thalline margins of the lirellae are rather thick and the black labia, when visible, are rather sharply delimited from the latter. There are also slight differences in ascospore shape with those in G. insulana typically 3·0–3·5 times as long as wide (Lücking et al. Reference Lücking, Archer and Aptroot2009). In addition, G. insulana has a thicker thallus and thalline margin of the lirellae, with the black labia rather sharply delimited although mostly covered by the thallus. In G. flosculifera the thallus and thalline margins of the lirellae are thin and the labia appear dark grey and roughened, and not sharply delimited from the thalline margin.
Graphis rajapakshana Weerakoon, Lücking & Aptroot sp. nov.
MycoBank No.: MB 832541
Corticolous Graphis resembling G. desquamescens, including in ascospore size, but having lirellae with a distinct lateral thalline margin.
Type: Sri Lanka, central mountain region, 20 km from Nuwaraeliya, Pattipola, 6°50′N, 80°50′E, 1850 m, high altitude, Pattipola Tea Plantation, September 1988, Cloonan 195 (PDA—holotype).
(Fig. 4A & B)
Fig. 4. Habitus, ascospores and lirella anatomy of Graphis species, all holotypes. A & B, G. rajapakshana; C & D, G. rimosothallina; E & F, G. thunsinhalayensis. Scales: A, C, D & E = 1 mm; B & F = 20 μm. In colour online.
Thallus corticolous, up to 5 cm diam., continuous; surface smooth to uneven, pale white to green-white; prothallus not observed. Thallus in section 100–150 µm thick, with prosoplectenchymatous upper cortex, 10–15 µm thick, irregular algal layer 30–60 µm thick, intermingled with and partially covered by large clusters of calcium oxalate crystals; medulla 50–80 µm thick, encrusted with large clusters of calcium oxalate crystals following the outlines of the dead periderm cells of the bark, and also filled with small, grey crystals.
Lirellae flexuose, sparsely branched, erumpent, lineola-type, with a lateral thalline margin, 1–3 mm long, 0·20–0·25 mm wide, 0·15–0·20 mm high; disc concealed, labia entire. Excipulum completely carbonized, 30–50 µm wide; hypothecium prosoplectenchymatous, 10–15 µm high, pale yellowish; hymenium 100–120 µm high, colourless, densely inspersed (type A sensu Lücking et al. Reference Lücking, Archer and Aptroot2009); epithecium granulose, 5–10 µm high, olive brown. Paraphyses unbranched; asci fusiform, 100–110 × 20 µm. Ascospores 8 per ascus, oblong and tapering, 11–13-septate, 35–50 × 7–8 µm, 5–6 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
Norstictic acid.
Etymology
This new species is named in honour of Dr Ranil Rajapaksha, senior lecturer at the University of Peradeniya, a pioneer pteridologist in Sri Lanka.
Distribution and habitat
The new species was collected in a high elevation tea planation in the central mountain range of Sri Lanka.
Discussion
This new species belongs to a small group centred around Graphis desquamescens (Fée) Zahlbr., with entire labia, a completely carbonized excipulum, an inspersed hymenium, transversely septate ascospores and containing norstictic acid. Among these, Graphis centrifuga Räsänen corresponds in the lateral thalline margin but has very long, radiately branched lirellae and very small ascospores (15–25 µm long). Graphis cervinonigra Zahlbr. and G. gonimica Zahlbr., also have a lateral thalline margin and small ascospores (20–30 µm long) and, at first, the lirellae are stellately branched. Graphis desquamescens (Fée) Zahlbr. agrees with the new species in ascospore size but its lirellae lack a thalline margin. Graphis mikuraensis Y. Ohmura & M. Nakan. (syn.: G. srilankensis Weerakoon et al.; see below) also belongs to this group but differs from the new species in the prominent lirellae and larger ascospores (50–70 µm long).
Additional material examined. Sri Lanka:Sabaragamuwa: mountain range, Coolbone Tea Estate, 7°01′N, 80°28′E, 860 m, mid elevation, July 1988, unknown collector 630. Central Province: Mahagasthota upper division, 6°56′N, 80°47′E, 1760 m, high elevation, February 1989, unknown collector 198; Nawanagala Tea Estate, 7°19′N, 80°51′E, 1180 m, high elevation, December 1989, Cloonan 228 (all PDA).
Graphis rimosothallina Weerakoon, Lücking & Aptroot sp. nov.
MycoBank No.: MB 832542
Corticolous Graphis with a thick, uneven, rimose thallus and Fissurina-like lirellae, a completely carbonized excipulum and transversely 7-septate ascospores of 32–37 × 8–10 µm.
Type: Sri Lanka, Horton Plains, 8 March 2015, Weerakoon NE224 (PDA—holotype; isotype: BM).
(Fig. 4C & D)
Thallus corticolous, up to 10 cm diam., continuous; surface uneven and rimose (deeply broken in parts), pale greenish; prothallus not observed. Thallus in section 0·5–1·0 mm thick, with prosoplectenchymatous upper cortex, 20–35 µm thick, irregular algal layer 50–100 µm thick, intermingled with, and partially covered by, large clusters of calcium oxalate crystals, and medulla 0·5–1·0 mm thick, encrusted with large clusters of calcium oxalate crystals.
Lirellae flexuose, often curved, sparsely branched, immersed, with a complete thalline margin, most similar to the consanguinea-type but superficially resembling a Fissurina, 1·0–3·5 mm long, 0·35–0·50 mm wide; disc concealed, labia striate, covered by a thin thalline margin or thick pruina. Excipulum completely carbonized, 50–100 µm wide; hypothecium not observed; hymenium 150–500 µm high, colourless, not inspersed; epithecium granulose, 5–10 µm high, olive brown. Paraphyses unbranched; asci fusiform, 100–180 × 20 µm. Ascospores 8 per ascus, fusiform, 7-septate, 32–37 × 8–10 µm, 3·5–4·0 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
No substances detected by TLC.
Etymology
The name refers to the deeply fissured thallus.
Distribution and habitat
The new species was collected in the southern mountain range of Sri Lanka.
Discussion
Graphis rimosothallina has striate labia, a completely carbonized excipulum, a clear hymenium and transversely septate ascospores, and thus keys out in group 17 in the world key by Lücking et al. (Reference Lücking, Archer and Aptroot2009). In that group, it differs from other species with small ascospores and lacking secondary compounds by its rather unique morphology, strongly resembling a species of Fissurina. All the other species in that group have a Graphis-like thallus and at best a lateral thalline margin, with the striate labia clearly exposed. Graphis rimosothallina is distinguished from species of Fissurina by the strongly carbonized excipulum and the multiseptate, amyloid ascospores.
Graphis thunsinhalayensis Weerakoon, Arachchige & Lücking sp. nov.
MycoBank No.: MB 832543
Corticolous Graphis resembling G. subalbostriata but with smaller ascospores and lacking white lines between the striae of the labia.
Type: Sri Lanka, central mountain region, 20 km from Nuwaraeliya, Pattipola, 6°50′N, 80°50′E, 1760 m, high altitude, Mahagasthota Tea Estate, upper division, September 1988, Cloonan 72 (PDA— holotype).
(Fig. 4E & F)
Thallus corticolous, up to 5 cm diam., continuous; surface smooth to uneven, white to light yellowish grey; prothallus not observed. Thallus in section 100–120 µm thick, with loose upper cortex, 5–10 µm thick, irregular algal layer 30–50 µm thick, intermingled with and partially covered by large clusters of calcium oxalate crystals, and medulla 40–60 µm thick.
Lirellae flexuose, irregular to radiately branched, erumpent, with a thin, lateral thalline margin, tenella- to dichotoma-type, especially when young, older lirellae with a basal thalline margin, 1–10 mm long, 0·2–0·3 mm wide, 0·15–0·25 mm high; disc concealed, labia finely striate. Excipulum crenulate, apically carbonized, 25–50 µm wide; hypothecium prosoplectenchymatous, 10–15 µm high, hyaline to pale yellowish; hymenium 100–120 µm high, colourless, not inspersed; epithecium granulose, 5–10 µm high, olive brown. Paraphyses unbranched; asci fusiform, 100–110 × 20–25 µm. Ascospores 8 per ascus, oblong, 13–17-septate, 50–80 × 7–10 µm, 7–8 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
No substances detected by TLC.
Etymology
The name refers to Thun Sinhalaya, which was the last kingdom in the Kandy area and lost its autonomy in 1817 (Pieris Reference Pieris1945).
Distribution and habitat
The new species was collected in a high elevation tea planation in the central mountain range of Sri Lanka.
Discussion
Graphis thunsinhalayensis is part of a small group with apically carbonized, striate labia and medium-sized to large ascospores, lacking inspersion and secondary substances. The neotropical Graphis subalbostriata Vain. ex Lücking has larger ascospores and distinct white lines between the striae, and G. olivacea Redinger, also with larger ascospores, has lirellae with an apically thin, complete thalline margin. Graphis supertecta Müll. Arg. corresponds in ascospore size but differs in the immersed lirellae with pruinose labia. Graphis caribica Lücking differs in the greenish thallus and the prominent lirellae lacking or with a basal thalline margin only.
Additional material examined. Sri Lanka: central mountain region, Galhiriya Tea Estate, 7°26′N, 80°42′E, 1241 m, high elevation, December 1989, unknown collector 055; Cottaganga Ella, 7°22′N, 80°48′E, 1471 m, high elevation, December 1989, unknown collector 008; Galhiriya Tea Estate, upper division, 7°25′N, 80°42′E, 1260 m, high elevation, December 1989, unknown collector 291; road to Horton Plains, 6°50′N, 80°48′E, 1869 m, high elevation, February 1989, unknown collector 602; Pedro Tea Estate, 7°00′N, 80°44′E, 1920 m, high elevation, February 1989, unknown collector 074 & 087 & Cloonan 962 (all PDA).
Name Validation
Graphis verrucoserpens Lücking ex Lücking sp. nov.
MycoBank No.: MB 832544
Graphis verrucoserpens Lücking in Lücking et al., Lichenologist 41: 421 (2009) [nom. inval.].
Corticolous Graphis resembling G. proserpens but with a distinctly and rather coarsely verrucose thallus surface.
Type: Jamaica, parish of St. Thomas, Monkey Hill to south slope of Mossmans Peak, Blue Mts., 1494 m, 19 February 1953, Imshaug 14579 (MSC-0029035!—holotype; NYBG!—isotype).
(Fig. 5)
Fig. 5. Habitus of Graphis verrucoserpens, holotype. Scale = 1 mm. In colour online.
Thallus corticolous, up to 5 cm diam., continuous; surface rather coarsely verrucose, greenish grey; prothallus not observed. Thallus in section 50–150 µm thick, with dense upper cortex, 5–10 µm thick, irregular algal layer 20–50 µm thick, intermingled with and partially covered by large clusters of calcium oxalate crystals (particularly in the verrucae), and medulla 20–60 µm thick.
Lirellae flexuose, irregularly branched, erumpent, with a basal thalline margin, striatula-type, 1–5 mm long, 0·1–0·2 mm wide, 0·10–0·15 mm high; disc concealed, labia finely striate. Excipulum crenulate, apically (to laterally) carbonized, 20–30 µm wide; hypothecium prosoplectenchymatous, 10–15 µm high, hyaline to pale yellowish; hymenium 80–100 µm high, colourless, not inspersed; epithecium granulose, 5–10 µm high, olive brown. Paraphyses unbranched; asci fusiform, 80–90 × 15–20 µm. Ascospores 8 per ascus, oblong, 7–11-septate, 25–35 × 5–7 µm, 4·5–5·5 times as long as wide, colourless, I+ amyloid.
Secondary chemistry
No substances detected by TLC.
Discussion
This taxon had already been keyed out in Lücking et al. (Reference Lücking, Archer and Aptroot2009) but has not yet been validly described, an oversight corrected here. It is in most aspects identical to Graphis proserpens but differs clearly in the rather coarsely verrucose thallus.
Additional specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 964 (BM).
Other Species
Allographa acharii (Fée) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 15 (2018).—Graphis acharii Fée, Essai Crypt. Écorc.: 39 (1824).—Graphina acharii (Fée) Müll. Arg., Mém. Soc. Phys. Genève 29(8): 38 (1887); type: South America, unknown locality, s. dat., s. col. (G, not seen).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, acharii-type, 1–10 × 0·5–1 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 2–6 per ascus, muriform, 80–170 × 15–30 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 944 (BM).
Allographa adpressa (Vain.) Lücking & Kalb
Herzogia 31: 546 (2018).—Graphis adpressa Vain., Acta Soc. Fauna Fl. Fenn. 7: 119 (1890); type: Brazil, Vainio s. n. (TUR-Vainio 27851! Vainio, Lich. Bras. Exs. 1289—lectotype, Wirth & Hale Reference Wirth and Hale1978).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, without a clear thalline margin, nuda-type, 0·5–1·5 × 0·25–0·35 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–17-septate, 50–70 × 13–15 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nawalapitiya, 2015, Weerakoon 10020 (BM).
Allographa anguilliradians Lücking ex Lücking
In Lücking & Kalb, Herzogia 31: 544 (2018).—Graphis anguilliradians Lücking, in Lücking et al., Lichenologist 41: 408 (2009) [nom. inval.]; type: Trinidad, Imshaug & Imshaug 31858 (MSC-0024122!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae prominent, very long-radiately branched, with a complete (thin) thalline margin, anguilliradians-type, 3–10 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 6–8 per ascus, transversely 9–13-septate, 50–70 × 8–11 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 319 (BM).
Allographa angustata (Eschw.) Lücking & Kalb
Herzogia 31: 547 (2018).—Graphis angustata Eschw., in Martius, Fl. Bras. 1(1): 73 (1833); type: Brazil, Martius s. n. (M, not seen).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, acharii-type, 1–7 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 11–15-septate, 50–100 × 12–18 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nawalapitiya, 2015, Weerakoon 10019 (BM).
Allographa argentata (Lücking & Umaña) Lücking & Kalb
Herzogia 31: 547 (2018).—Graphis argentata Lücking & Umaña, in Lücking et al., Fieldiana 46: 59 (2008); type: Costa Rica, Puntarenas, La Amistad International Park, Altamira Station (La Amistad Pacífico Conservation Area), Talamanca Ridge, Lücking 15269d (INB!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, acharii-type, 1–6 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium inspersed. Ascospores 4–6 per ascus, muriform, 90–120 × 15–25 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 518C (BM).
Allographa asterizans (Nyl.) Lücking & Kalb
Herzogia 31: 547 (2018).—Graphis asterizans Nyl., Acta Soc. Sci. Fenn. 7(2): 467 (1863); type: China, Hong Kong, s. col. (H-Nyl 7781!—holotype).
Thallus corticate, verrucose, greenish grey. Lirellae prominent, stellately branched, with a lateral (thick) thalline margin, asterizans-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 15–23-septate, 40–70 × 7–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Imbulpitiya, 2015, Weerakoon Im72 (BM).
Allographa bifera (Zahlbr.) Lücking & Kalb
Herzogia 31: 547 (2018).—Graphis bifera Zahlbr., in Handel-Mazzetti, Symbol. Sinic. 3: 47 (1930); type: China, Yunnan, Handel-Mazzetti 12262 (W!—holotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, without a clear thalline margin, nuda-type, c. 3 × 0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 15–21-septate, 80–100 × 8–15 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 619 (BM).
Allographa consanguinea (Müll. Arg.) Lücking & Kalb
Herzogia 31: 549 (2018).—Graphina consanguinea Müll. Arg., Nuov. Giorn. Bot. Ital. 21: 362 (1889).—Graphis consanguinea (Müll. Arg.) Lücking, in Lücking et al., Fieldiana 46: 6 (2008); type: Brazil, Glaziou s. n. (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae immersed, elongate-irregularly branched, with complete (thick) thalline margin, consanguinea-type, 1–7 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 110–125 × 30–35 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 558 (BM).
Allographa elixii (A. W. Archer) Lücking & Kalb
Herzogia 31: 550 (2018).—Phaeographina elixii A. W. Archer, Biblioth. Lichenol. 78: 14 (2001).—Graphis elixiana A. W. Archer, Telopea 11: 73 (2005); type: Australia, Queensland, Elix 16321 (CANB—holotype, photograph seen).
Thallus corticate, smooth, greenish grey. Lirellae prominent, short-sparsely branched, with a complete (thick) thalline margin, illinata-type, 1–3 × 0·4–0·6 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 1 per ascus, muriform, 15–21 × 1–3-septate, 90–130 × 12–17 µm, hyaline.
Secondary chemistry
Hirtifructic acid.
Selected specimen examined. Sri Lanka: Horton Plains National Park, 2015, Weerakoon 76A (BM).
Allographa elongata (Zenker) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 18 (2018).—Graphis elongata Zenker, in Goebel & Kunze, Pharmaceutische Waarenkunde (Eisenach) 1: 165, tab. XXII, fig. 1 (1827); type: not seen.
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 13–17-septate, 70–80 × 11–15 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 982 (BM).
Allographa flavens (Müll. Arg.) Lücking & Kalb
Herzogia 31: 550 (2018).—Graphis flavens Müll. Arg., Flora 65: 334 (1882); type: Indonesia, Java, Junghuhn 446 (L—holotype!; G—isotype!).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a lateral (thick) thalline margin, marginata-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 11–15-septate, 50–70 × 9–11 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 224 (BM).
Allographa glauconigra (Vain.) Lücking & Kalb
Herzogia 31: 550 (2018).—Graphis glauconigra Vain., Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 242 (1921); type: Philippines, Merrill 6791 (TUR-Vainio 27884!—lectotype, Lücking et al. Reference Lücking, Archer and Aptroot2009).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, with a complete (thin) thalline margin, acharii-type, 1–3 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 4 per ascus, transversely 9–13-septate, 40–80 × 8–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 300 (BM).
Allographa hossei (Vain.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 18 (2018).—Graphis hossei Vain., Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo 1(no. 3): 53 (1921); type: Thailand [Siam], Hosseus s. n. (TUR-Vainio 27806!—lectotype, Lücking et al. Reference Lücking, Archer and Aptroot2009).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a basal thalline margin, hossei-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 2–8 per ascus, transversely 11–17-septate, 50–110 × 8–14 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 674 (BM).
Allographa illinata (Eschw.) Lücking & Kalb
Herzogia 31: 551 (2018).—Graphis illinata Eschw., in Martius, Fl. Bras. 1: 82 (1833).—Graphina illinata (Eschw.) M. Wirth & Hale, Smiths. Contr. Bot. 40: 37 (1978); type: Brazil, Martius s. n. (M—lectotype, Wirth & Hale (Reference Wirth and Hale1978), not seen).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a complete thalline margin, illinata-type, 1–7 × 0·4–0·6 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 100–150 × 18–30 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nuwaraeliya, 2015, Weerakoon Ne476 (BM).
Allographa ingarum (Vain.) Lücking & Kalb
Herzogia 31: 551 (2018).—Graphis ingarum (Vain.) Lücking, in Lücking et al., Lichenologist 41: 436 (2009).—Graphis angustata var. ingarum Vain., Ann. Acad. Sci. Fenn., Ser. A 6(no. 7): 158 (1915); type: Guadeloupe, Duss 1190 (TUR-Vainio 27848!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, acharii-type, 1·5–7·0 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–13-septate, 38–48 × 7–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Adam's Peak, 2015, Weerakoon 352 (BM).
Allographa jayatilakana Weerakoon, Arachchige & Lücking
In Jatnika et al., Lichenologist 51: 230 (2019); type: Sri Lanka, Western Province, Colombo City, Hokandara, G. Weerakoon GW 100 (PDA!—holotype; B!, BM!—isotypes).
Thallus corticate, uneven, white-grey. Lirellae prominent, unbranched to sparsely branched, pigmented, chrysocarpa-type, 1–5 × 0·3–0·4 mm; disc closed, labia mostly entire but becoming striate in older lirellae, apically exposed, black, laterally covered by a yellow to yellow-orange pruina. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, terminally muriform, with 15–19 transverse septa and usually one or two of the (sub)terminal cells on each side with a longitudinal septum, 80–120 × 9–12 µm, hyaline.
Secondary chemistry
Lirellae covered with yellow anthraquinone, K+ yellow slowly turning violet, in microscopic section with vanishing K+ yellow efflux and pigment granules, then turning purple-violet.
Selected specimen examined. Sri Lanka: type.
Allographa knucklensis (Weerakoon, Wijey. & Wolseley) Lücking & Kalb
Herzogia 31: 552 (2018).—Graphis knucklensis Weerakoon et al., in Weerakoon et al., Bryologist 115: 79 (2012); type: Sri Lanka, Knuckles conservation area, Weerakoon & Wijeyaratne 971 (PD!—holotype).
Thallus ecorticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, albotecta-type, 1–3 × 0·2–0·3 mm; disc partly exposed, white, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, muriform 7–9 × 0–2-septate, 25–35 × 7–8 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: type.
Allographa leprographa (Nyl.) Lücking & Kalb
Herzogia 31: 552 (2018).—Graphis leprographa Nyl., Acta Soc. Sci. Fenn. 26(no. 10): 21 (1900); type: Sri Lanka [Ceylon], 1879, Almquist (H-Nyl 7398!—lectotype, Staiger Reference Staiger2002).
Thallus corticate, smooth, creamish to white-grey. Lirellae prominent, short-sparsely branched, with a lateral (thick) thalline margin, marginata-type, 1–3 × 0·25–0·35 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 1–2 per ascus, muriform, 60–110 × 16–30 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: type.
Allographa leptospora (Vain.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 19 (2018).—Graphis leptospora Vain., Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo 1(no. 3): 53 (1921); type: Thailand [Siam], Hosseus s. n. (TUR-Vainio 27807!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, with a complete (thin) thalline margin, acharii-type; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–23-septate, 80–120 × 6–10 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Nawalapitiya, 2015, Weerakoon 10023 (BM).
Allographa leucaenae (Aptroot) Lücking & Kalb
Herzogia 31: 552 (2018).—Graphis leucaenae Aptroot, in Lücking et al., Lichenologist 41: 437 (2009); type: Indonesia, Java, Groenhart 5190 (L!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, with a complete (thin) thalline margin, acharii-type, 1–5 × 0·7–1·2 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 13–19-septate, 75–100 × 15–22 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 817 (BM).
Allographa longula (Kremp.) Lücking & Kalb
Herzogia 31: 552 (2018).—Graphis longula Kremp., Flora 59: 380 (1876); type: Brazil, Glaziou 5497 (G!—lectotype, Wirth & Hale Reference Wirth and Hale1978).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–10 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely to laterally carbonized. Hymenium clear. Ascospores 6–8 per ascus, transversely 11–17-septate, 50–70 × 10–13 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles mountain range, 2015, Weerakoon 243 (BM).
Allographa macella (Kremp.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 20 (2018).—Graphis macella Kremp., Flora 59: 380 (1876).—Graphina macella (Kremp.) Müll. Arg., Flora 63: 39 (1880); type: Brazil, Glaziou 6289b (M!—lectotype, Wirth & Hale Reference Wirth and Hale1978).
Thallus corticate, smooth, greenish grey. Lirellae prominent, short-sparsely branched, with a complete (thin) thalline margin, acharii-type, 1–3 × 0·5–0·7 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely to laterally carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 70–170 × 20–40 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 900 (BM).
Allographa mahaeliyensis (Weerakoon, Jayalal & Lücking) Lücking & Kalb
Herzogia 31: 553 (2018).—Graphis mahaeliyensis Weerakoon et al., in Weerakoon et al., Nova Hedwigia 101: 82 (2015); type: Sri Lanka, Central Province, Horton Plains, Jayalal B.6.3.2.5 (PD!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a basal thalline margin, striatula-type, 1–3 × 0·4–0·6 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 90–100 × 20–25 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: type.
Allographa marginata (Raddi) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 20 (2018).—Graphis marginata Raddi, Memor. Soc. Ital. Sc. 18: 344 (1820); type: Brazil, s. col. (not seen).
Thallus corticate, smooth, creamish to white-grey. Lirellae prominent, elongate-irregularly branched, with a lateral (thick) thalline margin, marginata-type, 1–8 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 11–25-septate, 50–120 × 9–18 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 841 (BM).
Allographa nana (Fée) Lücking & Kalb
Herzogia 31: 553 (2018).—Graphis nana (Fée) Spreng., Syst. Veget. 4(1): 249 (1827).—Opegrapha nana Fée, Essai Cryptog. Écorc. Offic. 1: 26 (1824); type: South America, s. col. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, without a clear thalline margin, nuda-type; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 15–25 × 5–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nawalapitiya, 2015, Weerakoon 10013 (BM).
Allographa nuda (H. Magn.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 75 (2018).—Graphis nuda (H. Magn.) Staiger & Lücking, in Lücking et al., Fieldiana 46: 93 (2008).—Graphina nuda H. Magn., Ark. Bot. 3(10): 266 (1955) [non Graphis nuda Staiger, Biblioth. Lichenol. 85: 245 (2002), nom. inval., ICBN, Art. 32, 35–37]; type: Hawaii, Faurie 1025b (UPS!—holotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, without a clear thalline margin, nuda-type; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, muriform, 25–40 × 10–20 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Horton Plains, 2015, Weerakoon 19215 (BM).
Allographa pavoniana (Fée) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 21 (2018).—Graphis pavoniana Fée, Essai Crypt Écorc. Offic. 1: 40 (1824); type: unknown locality, s. dat., s. col. (G!—lectotype, Staiger Reference Staiger2002).
Thallus partly ecorticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, caesiella-type, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 7–17-septate, 30–60 × 7–11 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 600 (BM).
Allographa phaeospora (Vain.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 21 (2018).—Graphis phaeospora Vain., Acta Soc. Fauna Fl. Fenn. 7: 97 (1890); type: Brazil, Vainio s. n. (TUR-Vainio 27166!—holotype; Vainio, Lich. Bras. Exs. 682).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, negrosina-type, 1–7 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 1–4 per ascus, muriform, 110–130 × 25–35 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 119 (BM).
Allographa plagiocarpa (Fée) Lücking & Kalb
Herzogia 31: 554 (2018).—Graphis plagiocarpa Fée, Essai Crypt. Écorc. Offic. 1: 38 (1824); type: unknown locality, s. col. (G!—isotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, with a lateral (thick) thalline margin, dussii-type, 0·5–1·5 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 80–120 × 10–18 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Sinharaja, 2015, Weerakoon MO72 (BM).
Allographa polystriata (Makhija, A. Dube, Adaw. & Chitale) Lücking & Kalb
Herzogia 31: 555 (2018).—Graphis polystriata Makhija et al., Geophytology 36: 65 (2006); type: India, Maharashtra, Makhija & Mantri 00.278 (AMH—holotype, not seen).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a basal thalline margin, striatula-type, 1–7 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 11–23-septate, 50–110 × 4–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 1006 (BM).
Allographa rhizicola (Fée) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 22 (2018).—Opegrapha rhizicola Fée, Essai Crypt. Écorc. Offic. 1: 33 (1824).—Graphis rhizicola (Fée) Lücking & Chaves, in Lücking et al., Fieldiana 46: 102 (2008); type: South America, unknown locality (G—holotype, not seen).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a complete thalline margin, illinata-type, 2–10 × 0·4–0·6 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 19–25-septate, 70–130 × 11–20 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 303 (BM).
Allographa rimulosa (Mont.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 22 (2018).—Opegrapha rimulosa Mont., Ann. Sci. Nat. Bot., Ser. 2 18: 271 (1842).—Graphis rimulosa (Mont.) Trevis., Spighe et Paglie: 11 (1853).—Graphis rimulosa (Mont.) Müll. Arg., Bull. Soc. Roy. Bot. Belg. 30: 79 (1891) [nom. illeg., ICBN Art. 52.1]; type: Guyana, Leprieur 200 (PC—lectotype, Wirth & Hale (Reference Wirth and Hale1978), not seen).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, without a clear thalline margin, striatula-type, 1–3 × 0·2–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 7–11-septate, 30–50 × 7–13 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 583 (BM).
Allographa ruiziana (Fée) Lücking & Kalb
Herzogia 31: 555 (2018).—Opegrapha ruiziana Fée, Essai Crypt. Écorc. Offic. 1: 27 (1824).—Graphis ruiziana (Fée) A. Massal., Mem. Lichenogr.: 111 (1853).—Graphina ruiziana (Fée) Müll. Arg., Mém. Soc. Phys. Genève 29(8): 38 (1887); type: South America, unknown locality, Humboldt & Bonpland s. n. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very short-unbranched, without a clear thalline margin, nuda-type, 0·5–1·5 × 0·2–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, muriform, 35–65 × 10–20 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 352 (BM).
Allographa rustica (Kremp.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 23 (2018).—Graphis rustica Kremp., Nuovo Giorn. Bot. Ital. 7: 61 (1875); type: Singapore, Beccari 258 (M—lectotype, Archer (Reference Archer2006), not seen).
Thallus corticate, bumpy, greenish grey. Lirellae prominent, elongate-irregularly branched, with a lateral (thick) thalline margin, marginata-type, 1–7 × 0·4–0·6 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 11–17-septate, 65–85 × 8–11 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 812 (BM).
Allographa sauroidea (Leight.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 23 (2018).—Graphis sauroidea Leight., Trans. Linn. Soc. London 25: 452 (1866); type: Brazil, Spruce 331 (BM—holotype, not seen).
Thallus corticate, smooth, pale greenish grey. Lirellae prominent, elongate-irregularly branched, without a clear thalline margin, hossei-type, 1–5 × 0·3–0·4 mm; disc exposed, dark, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–15-septate, 45–60 × 10–14 µm, hyaline.
Secondary chemistry
Lichexanthone.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 013 (BM).
Allographa sayeri (Müll. Arg.) Lücking & Kalb
Herzogia 31: 555 (2018).—Graphis sayeri Müll. Arg., Flora 70: 401 (1887); type: Australia, Sayer s. n. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae immersed, elongate-irregularly branched, with a lateral (thick) thalline margin, caesiella-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, white pruinose, entire. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–11-septate, 30–55 × 6–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 179 (BM).
Allographa sitiana (Vain.) Lücking & Kalb
Herzogia 31: 555 (2018).—Graphis sitiana Vain., Acta Soc. Fauna Fl. Fenn. 7(2): 120 (1890); type: Brazil, Vainio 533 (TUR-Vainio 27854!—holotype).
Thallus ecorticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a lateral (thick) thalline margin, farinulenta-type, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, white pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 3–5-septate, 20–30 × 7–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Jaffna, 2015, Weerakoon JF27 (BM).
Allographa striatula (Ach.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 26 (2018).—Graphis striatula (Ach.) Spreng., Syst. Veg., Ed. 16 4(1): 250 (1827).—Opegrapha striatula Ach., Syn. Meth. Lich.: 74 (1814); type: Guinea, s. col. (H-Acharius 629!—holotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, short-sparsely branched, without a clear thalline margin, striatula-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 9–15-septate, 40–60 × 8–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 626 (BM).
Allographa vestitoides (Fink) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 27 (2018).—Graphis vestitoides (Fink) Staiger, Biblioth. Lichenol. 85: 263 (2002).—Graphina vestitoides Fink, Mycologia 19: 218 (1927); type: Puerto Rico, Fink 1986 (MICH—lectotype, Wirth & Hale (Reference Wirth and Hale1978), not seen).
Thallus corticate, smooth, greenish grey. Lirellae prominent, elongate-irregularly branched, with a complete (thin) thalline margin, acharii-type, 1–8 × 0·5–0·8 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 2–6 per ascus, terminally muriform, 19–23 × 0–2-septate, 80–140 × 12–16 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 636 (BM).
Allographa xanthospora (Müll. Arg.) Lücking & Kalb
In Kalb et al., Phytotaxa 377: 29 (2018).—Graphis xanthospora Müll. Arg., Bull. Herb. Boissier 3: 320 (1895); type: Australia, Sayer s. n. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae immersed, elongate-irregularly branched, with a lateral (thick) thalline margin, caesiella-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, white pruinose, entire. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 30–40 × 8–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 095 (BM).
Graphis allugallenensis Weerakoon, Wijey. & Rivas Plata
In Weerakoon et al., Bryologist 115: 77 (2012); type: Sri Lanka, Central Province, Allugallena nature trail, Weerakoon & Wijeyaratne 255A (PD!—holotype).
Thallus corticate, verrucose, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–5 × 0·2–0·3 mm; disc closed, labia verrucose, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–11-septate, 35–40 × 7–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: type.
Graphis anfractuosa (Eschw.) Eschw.
In Martius, Fl. Bras. 1(1): 86 (1833).—Scaphis anfractuosa Eschw., Syst. Lich.: 25 (1824); type: Brazil, Martius s. n. (M—lectotype, Wirth & Hale (Reference Wirth and Hale1978), not seen; G!—isolectotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, short-sparsely branched, without a clear thalline margin, hossei-type, 1–3 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 7–11-septate, 25–45 × 6–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Hakgala, 2015, Weerakoon Hg51 (BM).
Graphis arbusculiformis (Vain.) Lücking
In Lücking et al., Lichenologist 44: 391 (2012).—Graphis subdisserpens f. arbusculiformis Vain., Bot. Tidsskr. 29: 132 (1909); type: Thailand, Schmidt s. n. (TUR-Vainio 27563!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, very long-radiately branched, with a lateral thalline margin, centrifuga-type, 1–10 × 0·20–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely to laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 7–9-septate, 25–35 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Hakgala, 2015, Weerakoon 299 (BM).
Graphis aurita Eschw.
In Martius, Fl. Bras. 1: 90 (1833); type: Brazil, s. col. (M—holotype, not seen).
Thallus corticate, smooth, white-grey. Lirellae prominent, short-sparsely branched, with a lateral thalline margin, tenella-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–13-septate, 17–25 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 238 (BM).
Graphis bakeri Vain.
Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 253 (1921); type: Philippines, Baker 547 (TUR-Vainio 27874!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, caesiella-type, 1–5 × 0·20–0·25 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–7-septate, 20–40 × 6–9 µm, hyaline.
Secondary chemistry
Salazinic acid.
Selected specimen examined. Sri Lanka: Jaffna, 2015, Weerakoon JF26 (BM).
Graphis brahmanensis Aptroot
In Lucking et al., Lichenologist 41: 434 (2009); type: Papua New Guinea, Madang, Ramu Valley, Brahman, along road to Bundi, Aptroot 31685 (B!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a basal thalline margin, striatula-type, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–11-septate, 20–30 × 4–6 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Monaragala, 2015, Weerakoon Mn16 (BM).
Graphis caesiella Vain.
Acta Soc. Fauna Fl. Fenn. 7: 122 (1890); type: Brazil, Vainio s. n. (TUR-Vainio 27730!—holotype; Vainio, Lich. Brasil Exs. 45).
Thallus corticate, smooth, white-grey. Lirellae immersed, elongate-irregularly branched, with a lateral (thick) thalline margin, caesiella-type, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 20–40 × 6–8 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 553 (BM).
Graphis caesiocarpa Redinger
Ark. Bot. 27A(3): 23 (1935); type: Brazil, Malme 3602 (S 6507!—lectotype, Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thin) thalline margin, caesiella-type, 1–7 × 0·15–0·25 mm; disc closed, labia smooth, white pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 30–40 × 7–9 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon Kn27 (BM).
Graphis caribica Lücking
In Lumbsch et al., Phytotaxa 18: 59 (2011); type: Dominica, Imshaug & Imshaug 32777-A2 (MSC-0006670!—holotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, without a clear thalline margin, striatula-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–15-septate, 50–65 × 9–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 820 (BM).
Graphis chloroalba Makhija & Adaw.
Mycotaxon 91: 373 (2005); type: India, Andaman Islands, Nagarkar & Patwardhan 86.529 (AMH—holotype, not seen).
Thallus corticate, smooth, white-grey. Lirellae immersed, elongate-irregularly branched, with a lateral thalline margin, chloroalba-type; disc closed, labia smooth, white pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 20–25 × 5–7 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Imbulpitiya, 2015, Weerakoon Im70 (BM).
Graphis chlorotica A. Massal.
In Kremp., Verh. K. K. Zool.-Bot. Gesellsch. Wien 21: 865 (1871); type: Indonesia, Java, s. col. (W—holotype, not seen).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, with a basal thalline margin, tenella-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–13-septate, 30–40 × 5–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Wilpattu National Park, 2015, Weerakoon WL17 (BM).
Graphis cincta (Pers.) Aptroot
In Archer, Fl. Australia 57: 651 (2009); type: Dominican Republic, s. col. (L!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 7–9-septate, 25–35 × 6–8 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 030 (BM).
Graphis conferta Zenker
Pharmaceutische Waarenkunde (Eisenach) 1(3): 166 (1829); type: not seen.
Thallus corticate, smooth, white-grey. Lirellae sessile, short-sparsely branched, without a clear thalline margin, hossei-type; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 25–45 × 7–13 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nuwaraeliya, 2015, Weerakoon Ne126 (BM).
Graphis crassilabra Müll. Arg.
Flora 65: 502 (1882); type: Australia, Hartmann 78 (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin; disc closed, labia smooth, not pruinose, entire. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–15-septate, 45–65 × 7–12 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 217 (BM).
Graphis cupei Vain. ex Lücking
In Lücking et al., Lichenologist 41: 436 (2009); type: Cuba, Ekman 19 (TUR-Vainio 27371!—holotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a basal thalline margin, hossei-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. 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 by TLC.
Selected specimen examined. Sri Lanka: Imbulpitiya, 2015, Weerakoon 70A (BM).
Graphis dendrogramma Nyl.
In Crombie, J. Linn. Soc. London, Bot. 16: 226 (1877); type: Malaysia, Admiralty Islands, 1875, Moseley (H-Nyl 7165!—holotype).
Graphis irradians Nyl., Acta Soc. Sci. Fenn. 26(10): 21 (1900) [nom. illeg., ICBN Art. 53.1, non Graphis irradians Fée]—Graphis ceylandica Zahlbr., Cat. Lich. Univ. 2: 297 (1923); type: Sri Lanka [Ceylon], Almquist s. n. (H-Nyl 7900!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae immersed, very long-radiately branched, with a lateral (thick) thalline margin, dendrogramma-type, 1–7 × 0·15–0·25 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 20–30 × 6–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimens examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon Kn66 (BM); type of Graphis irradians Nyl.
Graphis descissa Müll. Arg.
Bull. Herb. Boissier 3: 318 (1895); type: Australia, Knight 339 (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–5 × 0·15–0·30 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 25–40 × 6–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Sinharaja World Heritage Site, 2015, Weerakoon Si05 (BM).
Graphis desquamescens Fée
Bull. Soc. Bot. Fr. 21: 24 (1874); type: Brazil, Glaziou 5082 (M!—lectotype, Wirth & Hale Reference Wirth and Hale1978).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, without a clear thalline margin, hossei-type, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 5–9-septate, 25–50 × 5–9 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Ambewella, 2015, Weerakoon Ne102 (BM).
Graphis diplocheila Vain.
Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 204 (1921); type: Philippines, McGregor 8605 (TUR-Vainio 27239!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thick) thalline margin, subserpentina-type, 1–5 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 1–2 per ascus, muriform, 50–130 × 15–25 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 1027 (BM).
Graphis disserpens Nyl.
Bull. Soc. Linn. Normandie, Sér. 2 7: 175 (1874) [non Graphis disserpens Vain., nom. illeg.]—Graphina disserpens (Nyl.) Müll. Arg., Flora 63: 23 (1880); type: Colombia [Nova Granata], Lindig 93 (H-Nyl 7415!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, very long-radiately branched, with a lateral thalline margin, dichotoma-type, 5–20 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 1–2 per ascus, muriform, 35–45 × 17–24 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 63A (BM).
Graphis dolichographa Nyl.
Acta Soc. Sci. Fenn. 7(2): 465 (1863); type: Colombia [Nova Granata], Lindig 866 (H-Nyl 6052!—holotype).
Thallus corticate, smooth, white-grey. Lirellae immersed, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–10 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 2–4 per ascus, muriform, 40–90 × 16–28 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 668A (BM).
Graphis dotalugalensis Weerakoon, Wijey. & Rivas Plata
In Weerakoon et al., Bryologist 115: 77 (2012); type: Sri Lanka, Central Province, Knuckles conservation area, Dotalugala Mountain peak, Weerakoon & Wijeyaratne 214A (PD!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a basal thalline margin, striatula-type, 1–6 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 11–15-septate, 60–100 × 10–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: type.
Graphis dracaenae Vain.
Cat. Welwitsch Afr. Pl. 2: 439 (1901); type: Africa, Welwitsch 397 (TUR-Vainio 27561!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, stellately branched, with a basal thalline margin, geraensis-type, 1–3 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–7-septate, 20–30 × 5–7 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 326 (BM).
Graphis dupaxana Vain.
Ann. Acad. Sci. Fenn., Ser. A 15(6): 241 (1921); type: Philippines, McGregor 14313 (TUR-Vainio 27869!—lectotype, Lücking et al. Reference Lücking, Archer and Aptroot2009).
Thallus corticate, smooth, white-grey. Lirellae prominent, short-sparsely branched, without a clear thalline margin, striatula-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 25–40 × 7–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 822 (BM).
Graphis duplicata Ach.
Syn. Meth. Lich.: 81 (1814); type: South America, unknown locality, s. col. (H-Ach 586!—lectotype, Staiger Reference Staiger2002).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, without a clear thalline margin, striatula-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 25–45 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Adam's Peak, 2015, Weerakoon AD11 (BM).
Graphis elegans (Sm.) Ach.
Syn. Meth. Lich.: 85 (1814).—Opegrapha elegans Sm., in Sowerby & Smith, Engl. Bot.: 26 (1808).—Aulacographa elegans (Sm.) Leight., Ann. Mag. Nat. Hist., Ser. 2 13: 389 (1854); type: Great Britain, England, Borrer s. n. (BM!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, without a clear thalline margin, striatula-type, 1–3 × 0·5–0·7 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–13-septate, 35–60 × 9–13 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 122 (BM).
Graphis ferruginea Vain.
Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 257 (1921); type: Philippines, Host 15028 (TUR-Vainio 27880!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, scripta-type, 1–5 × 0·15–0·25 mm; disc partly exposed, ochraceous, labia orange pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 4-septate, 20–25 × 5–7 µm, hyaline.
Secondary chemistry
Salazinic acid and anthraquinone.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 351 (BM).
Graphis furcata Fée
Essai Crypt. Écorc. Offic. 1: 40 (1824); type: South America, unknown locality, Humboldt & Bonpland s. n. (G!—holotype).
Thallus partly ecorticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, caesiella-type, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 6–8 per ascus, transversely 5–9-septate, 25–35 × 7–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 554 (BM).
Graphis galactoderma (Zahlbr.) Lücking
In Lücking et al., Lichenologist 41: 436 (2009).—Graphina galactoderma Zahlbr., in Handel-Mazzetti, Symbol. Sinic. 3: 57 (1930); type: China, Yunnan, Handel-Mazzetti 5912 (W—holotype, not seen).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–2 × c. 0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, muriform, 20–45 × 9–16 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Rilhena, 2015, Weerakoon Ri52 (BM).
Graphis glaucescens Fée
Essai Crypt. Écorc. Offic. 1: 36 (1824); type: South America, Humboldt & Bonpland s. n. (G—lectotype, Wirth & Hale (Reference Wirth and Hale1978), not seen; S 2184—isolectotype!).
Thallus ecorticate, smooth, bluish grey. Lirellae immersed, elongate-irregularly branched, with a lateral thalline margin, glaucescens-type, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, white pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 25–35 × 5–7 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 106 (BM).
Graphis gonimica Zahlbr.
Annls Mycol. 30: 431 (1932); type: China, Yunnan, Chung 591a (W! —lectotype, Lücking et al. Reference Lücking, Archer and Aptroot2009).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, with a lateral (thick) thalline margin, subserpentina-type, 1–3 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 5–7-septate, 20–35 × 5–9 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 530 (BM).
Graphis granulocarpa Redinger
Ark. Bot. 27A(3): 7, 34, tab. II, fig. 23 (1935); type: Brazil, Malme 3680 (S-6502—lectotype, Lücking et al. Reference Lücking, Archer and Aptroot2009).
Thallus corticate, smooth, white-grey. Lirellae prominent, very short-unbranched, without a clear thalline margin, granulocarpa-type, 0·5–1·0 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–7-septate, 30–40 × 9–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Illukkubura, 2015, Weerakoon 10145 (BM).
Graphis handelii Zahlbr.
In Handel-Mazzetti, Symbol. Sinic. 3: 44 (1930); type: China (Yunnan), Handel-Mazzetti 11403, 12788 (W!—syntypes).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, scripta-type, 0·7–3·0 × 0·10–0·25 mm; disc exposed, white, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 5–11-septate, 20–45 × 5–9 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 852 (BM).
Graphis hiascens (Fée) Nyl.
Ann. Sci. Nat. Bot., Sér. 4 11: 226 (1859).—Opegrapha hiascens Fée, Suppl. Essai Crypt. Écorc.: 25 (1837).—Graphina hiascens (Fée) Müll. Arg., Mém. Soc. Phys. Nat. Hist. Genève 29: 42 (1887); type: South America, unknown locality, s. col. (G—holotype, not seen).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, with a lateral (thick) thalline margin, subserpentina-type, 1–3 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 80–120 × 15–32 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 992 (BM).
Graphis immersella Müll. Arg.
Bull. Herb. Boissier 3: 319 (1895); type: Australia, Shirley 1793 (G!—lectotype, Archer Reference Archer1999).
Thallus corticate, smooth, greenish grey. Lirellae immersed, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·10–0·15 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 25–40 × 6–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Kalupahana, 2015, Weerakoon 014 (BM).
Graphis immersicans A. W. Archer
Aust. Syst. Bot. 14: 262 (2001); type: Australia, Queensland, Streimann 31317 (CANB—holotype, photograph seen).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–7-septate, 20–30 × 5–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Jaffna, 2015, Weerakoon JF176 (BM).
Graphis insulana (Müll. Arg.) Lücking & Sipman
In Lücking et al., Fieldiana 46: 84 (2008).—Graphina insulana Müll. Arg., Bot. Jahrb. 4: 56 (1883); type: Fiji, s.col. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–5 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 1–2 per ascus, muriform, 60–90 × 20–30 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Rilhena, 2015, Weerakoon Ri89 (BM).
Graphis intermedians Vain.
Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo 1(no. 3): 50 (1921); type: Thailand [Siam], Hosseus s. n. (TUR-Vainio 27214!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 13–15-septate, 40–55 × 9–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 538 (BM).
Graphis isidiza Adaw. & Makhija
Lichenologist 36: 361 (2004); type: India, Tamil Nadu, Makhija 01.106 (AMH—holotype, not seen).
Thallus corticate, with isidia, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete thalline margin, consanguinea-type, 2–8 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 20–35 × 4–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 07 (BM).
Graphis japonica (Müll. Arg.) A. W. Archer & Lücking
In Lücking et al., Lichenologist 41: 437 (2009).—Graphina japonica Müll. Arg., Flora 74: 113 (1891); type: Japan, Miyoshi 23 (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, with a lateral (thick) thalline margin, subserpentina-type, 1–3 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 2–4 per ascus, muriform, 50–70 × 18–25 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 214 (BM).
Graphis kelungana Zahlbr.
Feddes Repert. 31: 208 (1933); type: Taiwan [Formosa], Faurie 30 (W!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, subserpentina-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 11–13-septate, 40–60 × 10–14 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Adam's Peak, 2015, Weerakoon AD74 (BM).
Graphis lapidicola Fée
Bull. Soc. Bot. Fr. 21: 28 (1874); type: Brazil, Glaziou 3303 (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–5 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 2–4 per ascus, muriform, 50–70 × 15–22 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 226 (BM).
Graphis leptocarpa Fée
Essai Crypt. Écorc. Offic. 1: 36 (1824); type: South America, unknown locality, Humboldt & Bonpland s. n. (G!—lectotype, Wirth & Hale Reference Wirth and Hale1978).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, lineola-type, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 7–11-septate, 20–40 × 6–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Nuwaraeliya, 2015, Weerakoon 763 (BM).
Graphis leptoclada Müll. Arg.
Flora 65: 335 (1882).—Opegrapha rimulosa Mont. & Bosch, Lich. Java: 4 (1857) [nom. illeg., ICBN Art. 53.1; non Opegrapha rimulosa Mont.]; type: Indonesia, Java, Junghuhn s. n. (G!—lectotype, Archer Reference Archer2001).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–13-septate, 30–50 × 7–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 268 (BM).
Graphis leptogramma Nyl.
Bull. Soc. Linn. Normandie, Sér. 2 2: 111 (1868); type: New Caledonia, Deplanche s. n. (H-Nyl 7751!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 11–15-septate, 40–60 × 7–9 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 659 (BM).
Graphis librata C. Knight
Trans. Proc. New Zeal. Inst. 16: 404 (1884); type: New Zealand, Knight 67:23 (WELT—lectotype, Hayward (Reference Hayward1977), not seen).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally to completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 20–30 × 5–8 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 795 (BM).
Graphis lineola Ach.
Lichenogr. Univ.: 264 (1810); type: West Indies, unknown locality, Swartz s. n. (H-Ach 584!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 7–9-septate, 20–40 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 788 (BM).
Graphis litoralis Lücking, Sipman & Chaves
In Lücking et al., Fieldiana 46: 88 (2008); type: Costa Rica, Limón, Gandoca-Manzanillo Wildlife Refuge, Manzanillo Section (La Amistad Caribe Conservation Area), Manzanillo, Lücking 17098b (CR!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, caesiella-type, 1–5 × 0·1–0·2 mm; disc closed, labia smooth, white pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–9-septate, 30–35 × 6–8 µm, hyaline.
Secondary chemistry
Salazinic acid.
Selected specimen examined. Sri Lanka: Nuwaraeliya, 2015, Weerakoon 795 (BM).
Graphis longiramea Müll. Arg.
J. Linn. Soc. London, Bot. 29: 225 (1892); type: India, Watt 78 (G!—paratype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, very long-radiately branched, with a lateral thalline margin, longiramea-type, 5–15 × 0·4–0·5 mm; disc partly exposed, dark, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 7–23-septate, 60–100 × 8–12 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Namunukula, 2015, Weerakoon 162 (BM).
Graphis luluensis A. W. Archer
Mycotaxon 89: 326 (2004); type: Solomon Islands, Hill 8667 (BM!—holotype).
Thallus corticate, verrucose, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thin) thalline margin, negrosina-type; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 5–7-septate, 25–35 × 6–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 026 (BM).
Graphis mikuraensis Y. Ohmura & M. Nakan.
In Ohmura et al., Bull. Natl. Mus. Nat. Sci., Taiwan 42: 5 (2016); type: Japan, Izu Islands (Tokyo Metropolis), along the route between Sato and Ieno-sawa, Mikura-jima Island, on tree branch, elevation 500 m, 15 July 1981, Higuchi 10474 (TNS—holotype, not seen).
Graphis srilankensis Weerakoon et al., in Weerakoon et al., Bryologist 115: 79 (2012); holotype: Sri Lanka, Central Province, Manigala Mountain, Weerakoon & Wijeyaratne 812 (PD!).
Thallus corticate, smooth, creamish to white-grey. Lirellae prominent, elongate-irregularly branched, with a lateral (thick) thalline margin, marginata-type, 1–5 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 9–13-septate, 50–70 × 8–10 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: type of G. srilankensis.
Graphis myrtacea (Müll. Arg.) Lücking
In Lücking et al., Fieldiana 46: 93 (2008).—Graphina myrtacea Müll. Arg., Hedwigia 34: 33 (1895); type: Brazil, Ule 127 (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, tenella-type, 1–7 × 0·4–0·5 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 1–2 per ascus, muriform, 75–150 × 25–45 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 248 (BM).
Graphis negrosina (Vain.) Lücking
In Lücking et al., Lichenologist 41: 438 (2009).—Opegrapha negrosina Vain., Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 271 (1921); type: Philippines, Merrill 6790 (TUR-Vainio 27630!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thin) thalline margin, negrosina-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 9–11-septate, 35–45 × 7–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Nawalapitiya, 2015, Weerakoon 10014 (BM).
Graphis novopalmicola A. W. Archer & Lücking
In Lücking et al., Lichenologist 41: 439 (2009).—Graphina palmicola Müll. Arg., Flora 70: 402 (1887); type: Australia, Queensland, Sayer s. n. (G!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–5 × 0·5–0·6 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely to laterally carbonized. Hymenium inspersed. Ascospores 1 per ascus, muriform, 100–145 × 25–40 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 682 (BM).
Graphis oligospora Zahlbr.
In Handel-Mazzetti, Symbol. Sinic. 3: 45 (1930); type: China, Yunnan, Handel-Mazzetti 2451 (W!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, entire. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 25–35 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 545A (BM).
Graphis oxyclada Müll. Arg.
Flora 68: 512 (1885); type: Kenya, Hildebrandt 2540 (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, with a lateral thalline margin, marginata-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 30–45 × 7–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 331 (BM).
Graphis paraserpens Lizano & Lücking
In Lücking et al., Fieldiana 46: 96 (2008); type: Costa Rica, San José, Leonel Oviedo Ecological Reserve (Cordillera Volcánica Central Conservation Area), Lücking 15667 (USJ!—holotype; F!—isotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thin) thalline margin, symplecta-type, 2–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 2–6 per ascus, muriform, 30–40 × 9–11 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Namunukula, 2015, Weerakoon 639 (BM).
Graphis pinicola Zahlbr.
In Handel-Mazzetti, Symbol. Sinic. 3: 43 (1930); type: China, Yunnan, Handel-Mazzetti 2829 (W!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–11-septate, 25–35 × 10–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Sinharaja World Heritage Site, 2015, Weerakoon MO16 (BM).
Graphis proserpens Vain.
Bot. Tidsskr. 29: 132 (1909).—Graphis disserpens Vain., Acta Soc. Fauna Fl. Fenn. 7: 123 (1890) [nom. illeg., ICBN Art. 53.1; non Graphis disserpens Nyl.]; type: Brazil, Vainio s. n. (TUR-Vainio 27577A!—holotype; Vainio, Lich. Bras. Exs. 1091).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a basal thalline margin, striatula-type, 1–5 × 0·2–0·4 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 20–40 × 6–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 217 (BM).
Graphis pyrrhocheiloides Zahlbr.
Cat. Lich. Univers. 2: 321 (1924).—Graphis pyrrhocheila Vain., Hedwigia 46: 179 (1907) [non Mont. & Bosch 1855]; type: Indonesia, Malacca, Schmidt 9 (TUR-Vainio 27556!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, scripta-type, 1–5 × 0·2–0·3 mm; disc exposed, white, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 5–13-septate, 30–45 × 6–10 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Horton Plains National Park, 2015, Weerakoon HO02 (BM).
Graphis renschiana (Müll. Arg.) Stizenb.
Ber. Tät. St Gall. Naturw. Ges. 1891: 184 (1891).—Graphina renschiana Müll. Arg., Flora 68: 512 (1885); type: Madagascar, Rensch 969 (G!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, lineola-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, muriform, 25–35 × 9–12 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 363 (BM).
Graphis scripta (L.) Ach.
Kgl. Vetensk.-Akad. Nya Handl. 1809: 145 (1809).—Lichen scriptus L., Spec. Plant. 2: 1140 (1753); type: tab. XVIII and fig. 1 in Dillenius, Hist. Musc.—lectotype, Jørgensen et al. (Reference Jørgensen, James and Jarvis1994); Sweden, Malme s. n. (UPS!—epitype; Malme, Lich. Suec. Exs. 47).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, scripta-type, 1–7 × 0·2–0·4 mm; disc partly exposed, white, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–15-septate, 25–70 × 6–10 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 684 (BM).
Graphis slendrae Hale ex Lücking
In Lücking et al., Lichenologist 41: 440 (2009); type: Sabah, Hale 28009 (US!—holotype).
Thallus corticate, smooth, white-grey. Lirellae sessile, very long-radiately branched, with a basal thalline margin, slendrae-type, 1–10 × 0·1–0·2 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum completely carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–7-septate, 25–35 × 5–7 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 732 (BM).
Graphis stenotera Vain.
Ann. Acad. Sci. Fenn., Ser. A 15(no. 6): 243 (1921); type: Philippines, Fénix 12786 (TUR-Vainio 27887!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, short-sparsely branched, without a clear thalline margin, striatula-type, 1–3 × 0·15–0·30 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–15-septate, 20–40 × 6–9 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 906 (BM).
Graphis stipitata A. W. Archer
Mycotaxon 80: 368 (2001); type: Australia, Queensland, Streimann 16826 (CANB—holotype, photograph seen).
Thallus corticate, smooth, pale greenish grey. Lirellae erumpent, short-sparsely branched, with a basal thalline margin, hossei-type, 1–3 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 15–20 × 5–8 µm, hyaline.
Secondary chemistry
Lichexanthone.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 549A (BM).
Graphis streblocarpa (Bél.) Müll. Arg.
Flora 65: 502 (1882).—Opegrapha streblocarpa Bél., Voy. Indes Or., Bot. 2(Cryptog.): 134 (1846); type: not seen.
Graphis fissofurcata Leight., Trans. Linn. Soc. London 27: 177 (1869); type: Sri Lanka, Thwaites CL75 (BM!—lectotype, Archer Reference Archer2006).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 1–6 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum apically carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 70–95 × 20–30 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: type of Graphis fissofurcata.
Graphis subalbostriata Lücking
In Lücking et al., Lichenologist 41: 440 (2009).—Graphis angustata var. albostriata Vain., Ann. Acad. Sci. Fenn., Ser. A 6(no. 7): 158 (1915); type: Guadeloupe, Duss 517 (TUR-Vainio 27847!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–5 × 0·25–0·35 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 4–8 per ascus, transversely 9–10-septate, 80–125 × 8–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Kalupahana, 2015, Weerakoon 105A (BM).
Graphis subhiascens (Müll. Arg.) Lücking
In Lücking et al., Fieldiana 46: 111 (2008).—Graphina subhiascens Müll. Arg., Bot. Jahrb. 20: 2811 (1894); type: Tanzania, Holst 696 (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thick) thalline margin, subserpentina-type, 1–5 × 0·3–0·5 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 80–100 × 20–30 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 625A (BM).
Graphis submarginata Lücking
In Lücking et al., Fieldiana 46: 112 (2008).—Graphis marginata G. Mey. & Flot., Lich. Nova Acta Acad. Leopold.-Carol. 19(Suppl.): 229 (1843) [nom. illeg., ICBN Art. 53.1; non G. marginata Raddi].—Graphis lineola var. marginata (G. Mey. & Flot.) Zahlbr., Cat. Lich. Univ. 2: 317 (1924); type: Indonesia, Java, s. col. (not seen).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, scripta-type, 1–3 × 0·1–0·2 mm; disc exposed, white, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium inspersed. Ascospores 8 per ascus, transversely 5–9-septate, 25–35 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Jaffna, 2015, Weerakoon JF19 (BM).
Graphis subserpentina Nyl.
Acta Soc. Sci. Fenn. 7(2): 465 (1863); type: Sri Lanka [Ceylon], Hooker 8012 (G!—holotype).
Thallus corticate, smooth, creamish to white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral (thick) thalline margin, subserpentina-type, 4–8 × 0·3–0·4 mm; disc closed, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 1 per ascus, muriform, 80–130 × 20–35 µm, hyaline.
Secondary chemistry
Norstictic acid.
Selected specimen examined. Sri Lanka: type.
Graphis subtecta (Nyl.) Lücking
In Lücking et al., Lichenologist 41: 441 (2009).—Graphis analoga var. subtecta Nyl., Acta Soc. Sci. Fenn. 7(2): 465 (1863); type: Colombia [Nova Granata], Lindig 2725 (H-Nyl 7408b!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a complete (thin) thalline margin, negrosina-type; disc closed, labia smooth, not pruinose, entire. Excipulum completely carbonized. Hymenium clear. Ascospores 4–8 per ascus, muriform, 20–40 × 10–12 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 556 (BM).
Graphis subtenella Müll. Arg.
Flora 70: 400 (1887); type: Australia, Sayer s. n. (G!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, short-sparsely branched, with a lateral thalline margin, tenella-type, 1–3 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–13-septate, 30–40 × 5–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 595 (BM).
Graphis sundarbanensis Jagadeesh & G. P. Sinha
In Jagadeesh Ram et al., Lichenologist 39: 231 (2007); type: India, West Bengal, Jagadeesh Ram 828 (CAL—holotype, not seen).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, scripta-type; disc exposed, white, labia smooth, not pruinose, entire. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–11-septate, 25–45 × 6–10 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 546 (BM).
Graphis tenella Ach.
Syn. Meth. Lich.: 81 (1814); type: Guinea, s. col. (H-Acharius 585!—holotype).
Thallus corticate, smooth, white-grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–5 × 0·15–0·25 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 5–9-septate, 20–30 × 6–8 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 876 (BM).
Graphis tsunodae Zahlbr.
Annls Mycol. 14: 47 (1916); type: Japan, Tsunoda 182 (W!—holotype).
Thallus corticate, smooth, white-grey. Lirellae prominent, elongate-irregularly branched, without a clear thalline margin, striatula-type, 1–5 × 0·25–0·35 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 20–40 × 6–9 µm, hyaline.
Secondary chemistry
No substances detected by TLC.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 653 (BM).
Graphis vittata Müll. Arg.
Flora 65: 335 (1882); type: Indonesia, Java, Junghuhn 111 (G!—holotype).
Thallus corticate, smooth, greenish grey. Lirellae erumpent, elongate-irregularly branched, with a lateral thalline margin, tenella-type, 1–5 × 0·2–0·3 mm; disc closed, labia smooth, not pruinose, becoming striate. Excipulum apically to laterally carbonized. Hymenium clear. Ascospores 8 per ascus, transversely 7–11-septate, 30–50 × 7–8 µm, hyaline.
Secondary chemistry
Stictic acid.
Selected specimen examined. Sri Lanka: Knuckles Mountain Range, 2015, Weerakoon 907 (BM).
GW is grateful for research funding from Dilmah Conservation and the National Geographic Society (CS-R001-17 and WW-001R-17) and the opportunity to use the research facilities at the Natural History Museum of London and the Field Museum, Chicago. She also extends her gratitude to the Lichen Research Team (LRT): Pat Wolseley, Dulan Ranga Vidanapathirana, Dushantha Wasala and Chamara Udayanga. Thorsten Lumbsch is thanked for advice and support. Two anonymous reviewers are also thanked for comments that helped to improve the manuscript.
