Introduction
The lichen genus Fissurina Fée was established by Fée (Reference Fée1824) for the species with fissurine, lirelline ascomata and ovoid ascospores with a halo. The name Fissurina was used by Vainio (Reference Vainio1921) at the subgeneric level or for sections of the genus Graphis by Redinger (Reference Redinger1935) and Zahlbruckner (Reference Zahlbruckner1923). It was Kalb & Hafellner (Reference Kalb and Hafellner1992) who described a species in Fissurina as F. quadrispora Kalb & Hafellner and thus resurrected the name Fissurina at the generic level. Although the distinction of Fissurina from Graphis was recognized by Aptroot et al. (in Hawksworth Reference Aptroot, Karnefelt, Tibell and Hawksworth1994), stating that Fissurina seems to be a natural group which merits reintroduction, no systematic conclusions were drawn at that time. Eriksson & Hawksworth (Reference Eriksson and Hawksworth1998), in “Outline of the ascomycetes”, listed Fissurina as a synonym of Graphis following the system of Müll. Arg. (Reference Müller Argoviensis1880, Reference Müller Argoviensis1882) with spore-based genera. Later Fissurina was reintroduced by Staiger & Kalb (Reference Staiger and Kalb1999), and in phylogenetic studies (Staiger et al. Reference Staiger, Kalb and Grube2006; Mangold et al. Reference Mangold, Martín, Lücking and Lumbsch2008) it stands distinct amongst the known genera within Graphidaceae (including the Thelotremataceae).
The genus Hemithecium Trevis. was reintroduced by Staiger (Reference Staiger2002) for taxa with non-carbonized and convergent, often crenate, exciples. It was variable as to secondary chemistry, such as presence of the stictic acid complex, hymenium inspersion and ascospore type (spores hyaline and I+ blue-violet or brown and I+ reddish brown). Although chemistry and inspersion of hymenia may vary in other genera as well, the ascospore colour and iodine reaction are a constant character in most other graphidaceaen genera. Therefore, Staiger (Reference Staiger2002) proposed a subgeneric division of Hemithecium based on ascospore coloration with subgenus Hemithecium with hyaline ascospores and subgenus Leucogramma Staiger with brown ascospores. Recently the genus Pallidogramme Staiger et al. was introduced to accommodate the species in Hemithecium subgenus Leucogramma Staiger (Lücking et al. Reference Lücking, Chaves, Sipman, Umaña and Aptroot2008).
Materials and Methods
About 2000 specimens of lirelline Graphidaceae, mainly from the South Western Ghats, Eastern Himalaya, Western Himalaya, and the Andaman and Nicobar Islands of India, were studied from the AMH herbarium.
The species of Fissurina and Hemithecium were identified by comparison with types and protologues, or based on descriptions and photographs in recent treatments (e.g. Staiger & Kalb Reference Staiger and Kalb1999; Staiger Reference Staiger2002; Archer Reference Archer2006, Reference Archer2007, Reference Archer and McCarthy2009; Makhija & Adawadkar Reference Makhija and Adawadkar2007). Specimens are deposited in the Ajrekar Mycological Herbarium (AMH).
Sections of thalli and ascomata were mounted in water, 10% KOH (K), Lugol's solution (I), and lactophenol cotton-blue (LPCB). All measurements were made on material mounted in water. Secondary products were identified by thin-layer chromatography using standardized methods (Culberson & Kristinsson Reference Culberson and Kristinsson1970; Culberson Reference Culberson1972; White & James Reference White and James1985) using the solvent systems benzene-dioxane-acetic acid (180:45:5), hexane-diethylether-formic acid (130:80:20) and toluene-ethyl acetate-formic acid (139:83:8). The specimens were also examined under UV light (365 nm).
Taxonomy
The lichen genus Fissurina Fée
The lichen genus Fissurina, is widespread in tropical areas, and characterized by the following features: ascocarps fissurine; exciples poorly developed, non-carbonized or weakly carbonized; paraphyses tips warty or not; periphysoids present or absent; asci 1–8-spored; hyaline, ascospores rounded, usually thick-walled, trans-septate or muriform, often with a halo.
The similar genus Acanthothecis Clem., with warty paraphyses and periphysoids, can be differentiated from Fissurina by the ascospores without a thick jelly-like spore wall and with cylindrical spore locules. Graphis Adans. can be separated from Fissurina by its carbonized, well-developed exciple (labia) and ascospores without a halo.
After the comprehensive revisionary study of Graphidaceae by Staiger (Reference Staiger2002), many lichenologists reinvestigated this family and several changes have been made. As a result many species were added to Fissurina. Noteworthy contributions include the publications recording several species by Staiger (Reference Staiger2002), Nakanishi et al. (Reference Nakanishi, Kashiwadani and Moon2003), Staiger & Kalb (Reference Staiger, Kalb, Nash, Ryan, Diederich, Gries and Bungartz2004), Archer (Reference Archer2006, Reference Archer2007, Reference Archer and McCarthy2009), Makhija & Adawadkar (Reference Makhija and Adawadkar2007), Lendemer (Reference Lendemer2007), Lumbsch et al. (Reference Lumbsch, Ahti, Altermann, Amo de Paz, Aptroot, Arup, Bárcenas Peña, Bawingan, Benatti and Betancourt2011), and Lücking et al. (Reference Lücking, Seavey, Common, Beeching, Breuss, Buck, Crane, Hodges, Hodkinson and Lay2011).
In an earlier publication (Makhija & Adawadkar Reference Makhija and Adawadkar2007), 16 species of Fissurina with hyaline, trans-septate ascospores were described. Further studies on the family Graphidaceae have revealed the occurrence of 17 additional species, of which nine are new to science, and seven are new records to India. One species is not formally described as new due to scanty material and remains unnamed. Staiger (Reference Staiger2002) distinguished five lirella types for the identification of the ascomatal structures: dumastii-type, comparilis-type, incrustans-type, globulifica-type and subcontexta-type, based on their transition and formation of fruit bodies; the same types have been used in the present treatment.
Key to the species of Fissurina from India
1 Ascospores muriform ... 2
Ascospores trans-septate ... 16
2(1) Ascospores >150 µm long; ascomata 0·5–1·0 mm long, simple, round; asci 1-spored; ascospores 95–200 × 40–50 µm; no lichen substances present ... ... F. monospora C. Knight
Ascospores <150 µm long ... 3
3(2) Ascospores usually >100 µm long ... 4
Ascospores usually <100 µm long ... 5
4(3) Ascomata, 0·5–1·0 mm long, simple to branched; asci 1-spored; ascospores 70–100 (–130) × 20–50 µm; no lichen substances present ... ... F. submonospora B. O. Sharma et al.
Ascomata 0·1–0·4 mm long, simple to branched, asci 1-spored; ascospores 87–125 × 25–37 µm; stictic acid present ... ... F. microcarpa B. O. Sharma et al.
5(3) Ascospores usually > 40 µm long ... 6
Ascospores <40 µm long ... 8
6(5) Lichen substances present;.ascomata 0·5–1·5 mm long, simple, straight to curved, asci 1–4-spored; ascospores 70–78 × 20–25 µm; stictic, hypostictic acids present ... F. simplex B. O. Sharma et al.
Lichen substances absent ... 7
7(6) Ascomata 1·5–3·0 mm long, simple to rarely branched; asci 8-spored; ascospores 42·5–57·5 × 17·5–25·0 µm; no lichen substances present ... ... F. sporolata B. O. Sharma et al.
Ascomata simple to branched; ascospores muriform, 32–42 × 12–20 µm; no lichen substances present ... F. indica B. O. Sharma et al.
8(5) Ascospores <20 µm long ... 9
Ascospores >20 µm long ... 14
9(8) Hypothecium very thick, up to 40 µm; psoromic acid present; ascomata 1–3 mm long; asci 8-spored; ascospores 3-trans-septate, occasionally with 1 vertical septum, 9–14 × 4–6 µm ... F. globulifica (Nyl.) Staiger
Hypothecium thin, less than 40 µm thick; norstictic acid present or no substances ... 10
10(9) Norstictic acid present ... 11
Lichen substances absent ... 12
11(10) Ascomata branched, intermingled, short, up to 0·5–1·0 mm long of incrustans-type; ascospores submuriform, 7·5–12·5 × 5·0–7·5 µm; norstictic acid present ... ... Fissurina sp. 1
Ascomata simple, 0·2–1·5 mm long of dumastii-type; ascospores submuriform, 12–13 × 3–5 µm; norstictic acid present ... ... F. immersa B. O. Sharma et al.
12(10) Ascomata more than 5 mm long; ascomata 1–10 mm long, asteroidly branched, very rarely simple, semi-emergent; ascospores submuriform, 10–15 × 6–7 µm ... ... F. nicobarensis B. O. Sharma et al.
Ascomata <5 mm long ... 13
13(12) Ascomata 0·5–2·0 mm long, simple, immersed; ascospores muriform, 10–18 ×5–10 µm, with a halo of c. 2·5–5 µm ... F. nitidescens (Nyl.) Nyl.
Ascomata 0·2–0·5 mm long, branched, very close to each other, immersed to slightly raised; ascospores submuriform, 10–12 × 5–6 µm ... ... F. disposita B. O. Sharma et al.
14(8) Ascomata <1 mm long, simple to branched; ascospores muriform, 17–30 ×10–15 µm; no lichen substances present ... F. rubiginosa (Fée) Staiger
Ascomata >1 mm long ... 15
15(14) Ascospores submuriform, with 5–6 transverse septa and 1–2 longitudinal septa, 20–25 × 5–10 µm; no lichen substances present ... F. comparimuralis Staiger
Ascospores muriform, with 7–9 transverse septa and 2–3 longitudinal septa, 27–35 × 10–18 µm; no lichen substances present ... F. cingalina (Nyl.) Staiger
16(1) Thallus saxicolous; warty, cracked; ascomata 1–3 mm long; 17–25 × 6–8 µm, stictic acid present in the thallus ... F. saxicola Makhija & Adaw.
Thallus corticolous ... 17
17(16) Thalline margin with swollen tissue ... 18
Thalline margin without swollen tissue ... 20
18(17) Lichen substances present; ascomata short, 0·5–1·0 mm long, unbranched, of subcontexta-type; ascospores 18–28(–35) × 10–12 µm; stictic acid present in the thallus ... F. cf triticea (Nyl.) Staiger
Lichen substances absent ... 19
19(18) Ascomata 0·5–2·5 mm long, simple to branched, of subcontexta-type; exciple yellowish brown; ascospores 11–16 × 4–5 µm, with a conspicuous, 3–5 µm thick halo ... F. capsulata Makhija & Adaw.
Ascomata very long, thin, ribbon-like, intricate and anastomosed, of subcontexta-type; exciple bright reddish orange to dark red; ascospores 8–16 × 3–4 µm, without a conspicuous thick halo ... F. taeniocarpoides Makhija & Adaw.
20(17) Ascomata > 10 mm ( 8–13 mm) long; ascospores trans-septate, 8–16 (–21) × 3–4 (–6) µm; no lichen substances present ... F. longiramea Makhija & Adaw.
Ascomata less than 10 mm long ... 21
21(20) Ascospores >20 µm long ... 22
Ascospores <20 µm long ... 26
22(21) Ascomata up to 5 mm long ... 23
Ascomata >5 mm long ... 24
23(22) Ascomata 0·5–3·5 mm long; ascospores 15–25 × 8–14 µm; stictic acid present in the thallus ... F. canlaonensis (Vain.) Staiger
Ascomata 1–5 mm long; ascospores 12–25 × 8–10 µm; no lichen substances present ... F. insidiosa C. Knight & Mitt.
24(22) Lichen substances absent; ascomata 3–7 mm long; ascospores 60–70 µm ... ... F. dumastii Fée
Lichen substances present ... 25
25(24) Ascomata 1–6 mm long; ascospores 14–21 × 3–5 µm; protocetraric and fumarprotocetraric acids present ... F. karnatakensis Makhija & Adaw.
Ascomata 1–6 mm long; ascospores 20–27 × 7–10 µm; salazinic acid present ... ... F. andamanensis B. O. Sharma et al.
26(21) Ascomata >2 mm long ... 27
Ascomata <2 mm long ... 29
27(26) Salazinic acid present; ascomata 0·5–4·0 mm long, simple to branched; ascospores 10–12 × 4–6 µm ... F. dumastioides var. salazinica Makhija & Adaw.
Salazinic acid absent ... 28
28(27) Ascomata 2–4 mm long; ascospores 16–19 × 8–10 µm, stictic and constictic acid present ... F. khasiana Makhija & Adaw.
Ascomata 0·5–3·5 mm long; ascospores 10–12 × 4–6 µm, stictic acid present ... ... F. dumastoides (Fink) Staiger
29(26) Lichen substances present ... 30
Lichen substances absent ... 31
30(29) Thallus distinctly verrucose; ascomata 0·2–0·3 mm long; ascospores 7–12 ×3–4 µm; protocetraric and fumarprotocetraric acids present ... ... F. verrucosa Makhija & Adaw.
Thallus smooth; ascomata 0·2–0·4 mm long; ascospores 13–15 × 4–5 µm, stictic acid present ... F. inquinata C. Knight & Mitt.
31(29) Ascomata more than 1 mm long; ascomata 0·5–1·5 mm long; ascospores 10–13 × 5–7 µm ... F. subnitidula (Nyl.) Staiger
Ascomata up to 1 mm long ... 32
32(31) Ascomata of comparilis-type; ascospores 13–16 × 3–5 µm ... F. rugosa C. Knight
Ascomata of dumastii-type; ascospores 14–18 × 6–8µm ... ... F. coarctata Makhija & Adaw.
The Species
Fissurina andamanensis B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561853
Species F. dumastioidis var. salazinici similis ob acidum salazinicum continentem et ascosporas majores differt.
Typus: India, Andaman Islands, Middle Andaman, Betapur Range, Pitcher Nala, 26 December 1985, M. B. Nagarkar & P. K. Sethy 85.2462 (AMH—holotypus).
Fig. 1. Fissurina species, habitus. A, F. andamanensis (85.2462—holotype, AMH); B, F. disposita (77.1062—holotype, AMH); C, F. immersa (80.92—holotype, AMH); D, F. indica (76.67—holotype, AMH); E, F. microcarpa (83.382—holotype, AMH); F, F. nicobarensis (87.132—holotype, AMH); G, F. simplex (81.775—holotype, AMH); H, F. sporolata (80.510—holotype, AMH); I, F. submonospora (83.187—holotype, AMH); J, F. rubiginosa (83.40, AMH); K, Fissurina sp. 1 (83.228, AMH), Scales: A–J = 1 mm. 139 × 180 mm. In colour online.
Fig. 2. Fissurina species, sections of ascocarps. A, F. andamanensis (85.2462—holotype, AMH); B, F. disposita (77.1062—holotype, AMH); C, F. immersa (80.92—holotype, AMH); D, F. indica (76.67—holotype, AMH); E, F. microcarpa (83.382—holotype, AMH); F, F. nicobarensis (87.132—holotype, AMH). Scales: A–C = 50 µm; D–F = 100 µm. In colour online.
Thallus corticolous, yellowish brown, glossy, cracked, slightly verrucose.
Ascomata lirelline, fissurine, 1–6 mm long, simple to irregularly branched, concolorous with the thallus, immersed to semi-emergent, curved, thin, slender, with acute to subacute ends, structure of dumastii-type. Disc slit-like, epruinose. Exciple entire, dark blackish brown, orange-brown to non-carbonized, thin at the base, convergent, covered by the thalline margin up to the top, with a distinct, reddish orange prosoplectenchymatous upper corticiform layer. Hymenium hyaline, clear, 100–112 µm high, I−, KI−. Hypothecium light orange-brown, 12–14 µm high. Paraphyses simple. Periphysoids short, indistinctly warty at the tips. Asci 8-spored. Ascospores hyaline, 3-trans-septate, 20–27 × 7–10 µm, with a thin halo, I−, KI−.
Chemistry. Salazinic acid present.
Remarks. The occurrence of salazinic acid is rather rare amongst Fissurina species and so far it is known only in F. dumastioides (Fink) Staiger var. salazinica Makhija & Adawadkar. This can easily be differentiated from F. adamanensis in having smaller ascospores, 10–12 × 4–6 µm. The new species also resembles Fissurina rufula (Mont.) Staiger, and F. cf. triticea (Nyl.) Staiger in having similar ascomatal structure, but differs in having salazinic acid. Fissurina rufula has no lichen substances while F. triticea contains stictic acid. Fissurina elaiocarpa (A. W. Archer) A. W. Archer also has similar ascomatal structure but has muriform ascospores.
The species has been collected from the evergreen forest of Middle Andaman, where the vegetation is typically tropical.
Additional specimen examined. India: Andaman Islands: Middle Andaman, Betapur Range, Pitcher Nala, 1985, M. B. Nagarkar & P. K. Sethy 85.1854 (AMH).
Fissurina canlaonensis (Vain.) Staiger
Biblioth. Lichenol. 82: 161 (2002).—Graphis canlaonensis Vain. Ann. Acad. Sci. Fenn., ser. A, 15 (6): 261(1921).
Thallus corticolous, orange-brown, cracked, uneven, glossy, delimited by black hypothallus.
Ascomata lirelline, 0·5–1·0 mm long, slender, simple, very rarely branched, curved, immersed, terminally acute, structure of subcontexta-type. Disc slit-like, epruinose. Exciple entire, non-carbonized, orange-brown, present at the base, converging at the apical region, covered by the thalline margin up to the top, with a distinct upper corticiform layer, studded with crystals. Hymenium hyaline, sometimes with light yellow tinge, clear, 112–162 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple with branched tips. Periphysoids absent. Asci 8-spored. Ascospores hyaline, 3–4-trans-septate, 10–17 × 4–9 µm, with a 2·5–5·0 µm thick halo, I−, KI−.
Chemistry. Stictic, and constictic acids (sometimes) present.
Remarks. Fissurina canlaonensis has been collected from the moist deciduous forest in the Andaman Islands and is reported here for the first time from India. The species was earlier reported from the Philippines.
The specimens referred under Graphis canlaonensis Vain. (Patwardhan & Kulkarni Reference Patwardhan and Kulkarni1976) were later identified as Fissurina dumastioides Makhija & Adawadkar (Makhija & Adawadkar Reference Makhija and Adawadkar2007) and Graphis canlaonensis was erroneously reported from India.
Specimens examined. India: Andaman Islands: North Andaman, Tugapur Range, Pathar Tikari, moist deciduous forest, 1985, P. G. Patwardhan & M. B. Nagarkar 85.2599, 85.2683 (AMH).
Fissurina cingalina (Nyl.) Staiger
Biblioth. Lichenol. 85: 128 (2002).—Graphis cingalina Nyl. Acta Soc. Sci. Fenn., 26(10): 21 (1900).
Thallus corticolous, greenish grey, cracked, smooth, glossy, delimited by black hypothallus.
Ascomata lirelline, 1–4 mm long, branched, light brown, immersed, terminally acute, structure of dumastii-type. Disc slit-like, epruinose. Exciple entire, non-carbonized, orange, convergent, present at the base, covered by the thalline margin up to the top with a 10–12 µm thick, orange-brown upper corticiform layer studded with crystals. Hymenium hyaline, clear, 100–125 µm high, I−, KI−. Hypothecium indistinct, hyaline, 10–12 µm. Paraphyses simple. Periphysoids absent. Asci 8-spored. Ascospores hyaline, muriform, 7–9-trans-septate and 2–3 vertical septa, 27–35 × 10–18 µm, with a 2·5–5·0 µm thick halo, I−, KI−.
Chemistry. No lichen compounds present.
Remarks. This species has been recorded from the moist deciduous and semi-evergreen forests of Kerala, Tamil Nadu and Maharashtra in India. It was previously known from Sri Lanka, Venezuela and Brazil and has now been reported for the first time from India.
Specimens examined. India: Kerala: Munnar-Kumily road, 30 km, Cardamom hills, 1976, P. G. Patwardhan & A. V. Prabhu 76.739, 76.740. Maharashtra: Mahabaleshwar, Dhobi Ghat, 1974, M. B. Nagarkar, 74.640; Amboli, 1974, C. R. Kulkarni & A. V. Prabhu 74.1427; Shirgaonkar Point, 2000, U. V. Makhija 00.496. Tamil Nadu: Kodaikanal, Silver Cascade, 1975, P. G. Patwardhan & A. V. Prabhu 75.109; Naduvattum, Nilgiri hills, in Shola forest, elev. approx. 5000 ft., 1978, P. G. Patwardhan 78.105; Kollimalai, 1985, P. G. Patwardhan & M. B. Nagarkar 85.1531, 85.1536, 85.1537, 85.1539, 85.1572, 85.1609, 85.1633, 85.1647 (AMH).
Fissurina comparimuralis Staiger
Biblioth. Lichenol. 85: 134 (2002).
Thallus corticolous, brown, cracked, verrucose, slightly glossy, delimited by thin black hypothallus.
Ascomata lirelline, 1–4 mm long, slender, all over the thallus, flexuous, simple to irregularly branched, curved, immersed, terminally acute, structure of comparilis-type; thalline margin slightly raised. Disc slit-like, immersed. Exciple entire, non-carbonized, orange, convergent, present at the base, covered by the thalline margin up to the top, with a distinct prosoplectenchymatous upper corticiform layer studded with crystals. Hymenium hyaline, clear, 87–112 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple. Periphysoids short. Asci 8-spored. Ascospores hyaline, submuriform, 5–6 transverse and 1–2 longitudinal septa, 20–25 ×5–10 µm, with 1–2 µm thick halo, I+ weak blue.
Chemistry. No lichen compounds present.
Remarks. Fissurina comparimuralis was previously known from Brazil and is now reported for the first time from India. The species has been collected from the evergreen forest of Kerala in the Western Ghats of India.
Specimens examined. India: Kerala: Thekadi, 1973, P. G. Patwardhan & M. B. Nagarkar 73.2460 (AMH).
Fissurina disposita B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561854
Similis F. egenae et F. nitidescentis ob structuram ascomatium et ascosporas minores et submuriformes differt.
Typus: India, Meghalaya, Mawsmai, near Cherapunji, 28 October 1977, P. G. Patwardhan & M. B. Nagarkar 77.1062 (AMH—holotypus).
Thallus corticolous, brown, verrucose, cracked, flaking, glossy.
Ascomata lirelline, 0·2–0·5 mm long, immersed to slightly raised arising as the swelling which then cracks and gapes, branched, present all over the thallus, crowded, terminally acute, structure of comparilis-type. Disc creamish, sunken, epruinose. Exciple entire, non-carbonized, light brown, present at the base. Hymenium hyaline, clear, 62–75 µm high, I−, KI−. Paraphyses simple. Periphysoids short. Asci 8-spored. Ascospores hyaline, submuriform, 3–4 transverse and 1–2 longitudinal septa, 10–12 × 5–6 µm, I−, KI−.
Chemistry. No lichen substances present.
Remarks. Fissurina disposita differs from the most closely allied F. egena (Nyl.) and F. nitidescens (Nyl.) Nyl. in having smaller ascospores of 10–12 × 5–6 µm. Fissurina egena has 15–23 × 7–10 µm large, muriform ascospores. Fissurina cingalina (Nyl.) Staiger and F. columbina (Tuck.) Staiger also lack lichen substances, but they have much larger ascospores, 20–32 × 9–17 µm in F. cingalina, and 20–30 × 10–15 µm in F. columbina.
Fissurina disposita has been collected from the subtropical forest of Mawsmai near Cherrapunji and from the shola forest of Karnataka. The sholas are sheltered depressions in the landscape.
Additional specimens examined. India: Karnataka: Baba Budangiri road, in shola forest, 1980, P. G. Patwardhan & M. B. Nagarkar 80.42, 80.46, 80.47, 80.48 (AMH).
Fissurina immersa B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561855
Similis F. inabensis sed differt ascosporis minoribus et acido norstictico continente.
Typus: India, Karnataka, Mudigiri, 26 January 1980, P. G. Patwardhan 80.92 (AMH— holotypus).
Thallus corticolous, brown, uneven, finely cracked, glossy, delimited by the black hypothalloidal region at the periphery.
Ascomata lirelline, concolorous with the thallus, immersed to slightly raised arising as swelling which then cracks and gapes, simple to very rarely branched, 0·2–1·5 mm long, straight to curved, terminally acute, structure of dumastii-type. Disc sunken, slit-like, epruinose. Exciple entire, non-carbonized, present at the base, orange-brown, covered by the slightly raised thalline margin with a distinct prosoplectenchymatous corticiform layer. Hymenium hyaline, clear, 50–75 µm high, I−, KI−. Paraphyses thin, simple with indistinct warty tips. Periphysoids short, indistinctly warty at the tips. Asci 8-spored. Ascospores hyaline, submuriform, 3–4 transverse and 1 vertical septa, 12–13 × 3–5 µm, I+ weak blue.
Chemistry. Norstictic acid present.
Remarks. Fissurina immersa belongs in a group of species having dumastii type ascomata and is clearly distinguished from the other species of this group in having short lirellae, submuriform ascospores and norstictic acid in the thallus.
Fissurina inabensis (Vain.) Nakan. & Kashiw., a species from Japan having submuriform ascospores and somewhat similar ascomatal structure, can be differentiated from the present new species in having larger ascospores, 21–30 × 10–14 µm, and stictic acid. Fissurina elaiocarpa (A. W. Archer) A. W. Archer, with similar ascomatal structure and muriform ascospores, has large ascospores, 21–28 µm long.
Fissurina immersa has been collected from the evergreen forest of Karnataka.
Additional specimen examined. India: Karnataka: Mudigiri, 1980, P. G. Patwardhan 80.635 (AMH).
Fissurina indica B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561856
Similis Fissurinae cingalinae sed ascosporis majoribus differt.
Typus: India, Kerala, Wynad forest, on Gudalur-Nilambur Road, 20 January 1976, P. G. Patwardhan & A. V. Prabhu 76.67 (AMH—holotypus).
Thallus corticolous, greenish, cracked, verrucose, glossy, delimited by black hypothalloidal region at the periphery.
Ascomata small, 0·2–0·4 mm long, concolorous with the thallus, immersed, simple to branched, acute ends, structure of dumastii-type, present all over the thallus. Disc slit-like. Exciple entire, non-carbonized, orange-brown, convergent, indistinctly present at the base, covered by the distinct thalline margin up to the top with a prosoplectenchymatous corticiform layer, studded with crystals. Hymenium hyaline, clear, 137–162 µm high, I−, KI−. Hypothecium indistinct, hyaline. Paraphyses simple. Periphysoids short, indistinct. Asci 6–8-spored. Ascospores hyaline, muriform, multilocular, 32–42 ×12–20 µm, with an indistinct halo, I−, KI−.
Chemistry. No lichen substances present.
Remarks. The new species can be distinguished from other species having no lichen substances, viz. Fissurina cingalina (Nyl.) Staiger, F. columbina (Tuck.) Staiger and F. marginata Staiger, by its larger ascospores.
Additional specimens examined. India: Assam: Maniknagar, 1977, P. G. Patwardhan & M. B. Nagarkar 77.1220; Darugiri Reserved forest, 1978, M. B. Nagarkar 78.356. Kerala: Cardamom hills, Devicolam, 5 km on Kumily road, 1976, P. G. Patwardhan & A. V. Prabhu 76.750; Munnar-Kumily road, on 78 km from Cardamom hills, 1976, C. R. Kulkarni 76.842; Paranthal, near Trivendrum, 1976, P. G. Patwardhan & A. V. Prabhu 76.878, 76.903 (AMH).
Fissurina microcarpa B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561857
Similis Fissurinae undulatae sed ascosporis majoribus differt.
Typus: India, Tamil Nadu, Upper Kodayar, 24 January 1983, P. G. Patwardhan 83.382 (AMH—holotypus).
Thallus corticolous, brownish, cracked, warty, glossy.
Ascomata lirelline, all over the thallus, short, 0·1–0·4 mm long, immersed to slightly raised arising as swelling which then cracks and gapes, simple to branched, concolorous with the thallus, terminally acute, structure of dumastii-type. Disc slit-like. Exciple entire, non-carbonized, indistinct at the base, orange-brown, covered with the thalline margin up to the top with a distinct corticiform layer. Hymenium hyaline, clear, 112–150 µm high, I−, KI−. Hypothecium light yellow, 10–15 µm. Paraphyses simple. Periphysoids short, indistinctly warty at the tips. Asci 1-spored. Ascospores hyaline, muriform, multilocular, 87–125 × 25–37 µm, with a 5–8 µm thick halo, I−, KI−.
Chemistry. Stictic acid present.
Remarks. Fissurina microcarpa is morphologically somewhat similar to F. submonospora (described below). However, Fissurina submonospora has no lichen compounds in the thallus. Fissurina undulata (Müll. Arg.) M. Nakan. & Kashiw., a species from Japan, also has stictic acid and monosporate asci but that species has smaller ascospores, 30–38 × 15–18 µm.
Additional specimens examined. India: Karnataka: Shringeri, 2 km from Shringeri-Balehonur road, 1974, C. R. Kulkarni 74.3221, 74.3627. Tamil Nadu: Upper Kodayar, 1983, P. G. Patwardhan 83.571 (AMH).
Fissurina monospora C. Knight
Transact. New Zealand Instit. 15: 354 (1883).
Thallus corticolous, brown, cracked, verrucose, glossy, delimited by the black hypothallus.
Ascomata lirelline, 0·5–1·0 mm long, simple, round, immersed, terminally obtuse or rounded. Disc narrow to broad, epruinose, sunken. Exciple entire, non-carbonized, orange-brown, rudimentary at the base, covered by the crystal-studded thalline margin up to the top, with orange-brown corticiform layer. Hymenium hyaline, clear, laterally I−, 90–125 µm high. Hypothecium light yellow, 8–10 µm. Paraphyses simple. Asci 1-spored. Periphysoids short, not warty at the tips. Ascospores hyaline, muriform, 95–200 × 40–50 µm, with an indistinct halo, I+ weakly blue.
Chemistry. No lichen substances present in the thallus.
Remarks. Fissurina monospora is known from New Zealand and has now been reported for the first time from Karnataka in the Western Ghats of India.
Specimens examined. India: Karnataka: Balehonur-Shringeri road, 1974, P. G. Patwardhan & M. B. Nagarkar 74.3168; South Canara, Koppa forest, 5 km from Koppa on Shimoga road, 1976, C. R. Kulkarni 76.1076, 76.1178 (AMH).
Fissurina nicobarensis B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561858
Similis Fissurinae globulificae sed lirellis longioribus et acidis deficientibus differt.
Typus: India, Nicobar Islands, Great Nicobar, Campbell Bay to Laful Bay, 2 January 1987, P. K. Sethy & P. G. Patwardhan 87.132 (AMH—holotypus).
Thallus corticolous, yellowish brown, glossy, warty, delimited by black hypothalloidal region.
Ascomata lirelline, 1–10 mm long, branched, concolorous with the thallus, semi-emergent, terminally acute, structure of globulifica-type. Disc creamish, narrow to slightly broad, epruinose. Exciple entire, non-carbonized, orange-brown, convergent, thin at base, covered by the thalline margin up to the top, with a distinct, prosoplectenchymatous upper corticiform layer, studded with crystals. Hymenium hyaline, clear, 62–87 µm high, I−, KI−. Hypothecium distinct, light yellow, 10–15 µm thick. Paraphyses simple, long. Periphysoids short, thin, without warts. Asci 8-spored. Ascospores hyaline, submuriform, ellipsoid, 10–15 × 6–7 µm, with a thin halo, I+ weak violet.
Chemistry. No lichen substances present.
Remarks. The new species shows the globulifica-type of ascomatal structure. Fissurina globulifica (Nyl.) Staiger, with similar ascomatal structure, differs by having short lirellae of c. 1–3 mm long and containing psoromic acid.
Fissurina albocinerea (Vain.) Staiger and F. instabilis (Nyl.) Nyl. also have a similar ascomatal structure. They differ because F. albocinerea has 4-locular, trans-septate ascospores and F. instabilis has muriform ascospores and psoromic acid.
Fissurina nicobarensis has been collected in tropical rainforest from the Great Nicobar Island.
Additional specimens examined. India: Nicobar Islands: Great Nicobar, Campell Bay to Laful Bay, 1986, P. G. Patwardhan & P. K. Sethy 86.856; ibid., 1987, P. K. Sethy & P. G. Patwardhan 87.216 (AMH).
Fissurina nitidescens (Nyl.) Nyl
Lichenes Japoniae: 108 (1890)—Graphis nitidescens Nyl. In Tuckerman, N. Amer. Lich., 2: 123 (1888).
Thallus corticolous, light yellow, cracked, uneven, glossy.
Ascomata lirelline, 0·5–2 mm long, brown, simple, straight, immersed, terminally acute, structure of dumastii-type. Disc narrow to broad, brown, epruinose. Exciple entire, non-carbonized, orange, convergent, present below, covered by the raised thalline margin up to the top, with a thin corticiform layer, studded with crystals. Hymenium hyaline, clear, 87–100 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple. Periphysoids absent or indistinct. Asci 8-spored. Ascospores hyaline, muriform, 10–18 × 5–10 µm, with a halo of c. 2·5–5 µm, I−.
Chemistry. No lichen substances present.
Remarks. Fissurina nitidescens is known from Africa, Australia, USA and Brazil and has now been reported for the first time from India.
Specimens examined. India: Karnataka: Ganga Mulla, to Nagartirtha, on the way to Kudremukh, 1981, P. G. Patwardhan & D. W. Rane 81.77. Kerala: Wyanad, 1973, C. R. Kulkarni & P. D. Badhe 73.2810 (AMH).
Fissurina rubiginosa (Fée) Staiger
Biblioth. Lichenol. 85: 148 (2002).—Graphis rubiginosa Fée, Essai Sur les Cryptogames des écorces exotiques officinales: 47 (1825).
Fig. 3. Fissurina species, sections of ascocarps. A, F. simplex (81.775—holotype, AMH); B, F. sporolata (80.510—holotype, AMH); C, F. submonospora (83.187—holotype, AMH); D, F. rubiginosa (83.40, AMH). Scales: A–D = 100 µm. In colour online.
Thallus corticolous, greyish brown, cracked, rough, verrucose, slightly glossy.
Ascomata lirelline, very short, 0·4–0·6 mm, brownish, concolorous with the thallus, simple to branched, thin, immersed, terminally acute, structure of dumastii-type. Disc very narrow, not visible. Exciple entire, non-carbonized, convergent, present at the base, covered by the thalline margin up to the top, with a distinct yellowish upper corticiform layer, studded with crystals. Hymenium hyaline, clear, 100–120 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple. Periphysoids short, indistinctly warty at the tips. Asci 8-spored. Ascospores hyaline, muriform, round to elongate, 17–30 × 10–15 µm, with a 3–5 µm thick halo, I−.
Chemistry. No lichen substances present.
Remarks. Fissurina rubiginosa is well differentiated from F. elaiocarpa (A. W. Archer) A. W. Archer and F. incrustans Fée by having minute lirellae. Both species have muriform ascospores and no lichen compounds but have large ascomata, 1–6 (–8) mm long in Fissurina elaiocarpa and 1–4 mm long in F. incrustans. Fissurina coarctata Makhija & Adawadkar is also comparable with F. rubiginosa in its short, crowded lirellae and chemistry, but it has trans-septate ascospores.
Fissurina rubiginosa is known from French-Guayana and the USA and has now been reported for the first time from India.
Specimen examined. India: Tamil Nadu: Chitteri, 1983, P. G. Patwardhan 83.40 (AMH).
Fissurina simplex B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561859
Fissurina simplex distincte differt inter omnibus Fissurinae speciebus ob praesentiam acidi stictici et ascosporas majores.
Typus: India, Kerala, Silent valley, 31 December 1981, P. K. Sethy & S. P. Kekre 81.775 (AMH—holotypus).
Thallus corticolous, brown, cracked, glossy, verrucose, delimited by a black hypothalloidal region at the periphery.
Ascomata lirelline, 0·5–1·5 mm long, simple, straight, sometimes curved, concolorous with the thallus, immersed to slightly raised arising as a swelling which then cracks and gapes, terminally acute, structure of subcontexta-type. Disc narrow, sunken, epruinose. Exciple entire, non-carbonized, present at the base, convergent, orange-brown, often arising as fissures in swollen thalline margins (slightly ‘puffed’), covered by the distinct orange-brown corticiform layer of 10–12 µm, studded with crystals. Hymenium hyaline, clear, 120–150 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple. Periphysoids short. Asci 1–4-spored. Ascospores hyaline, muriform, 70–78 × 20–25 µm, with a thin halo, I−.
Chemistry. Stictic and hypostictic acids present.
Remarks. The new species Fissurina simplex can be distinguished easily from all other species by the presence of stictic acid, its swollen thalline margin, 1–4-spored asci and large muriform ascospores. Fissurina abdita (A. W. Archer) A. W. Archer and F. undulata (Müll. Arg.) M. Nakan. & Kashiw also produce stictic acid, however, they have smaller muriform or submuriform ascospores in the range of 21–35 µm. Fissurina quadrispora Kalb & Hafellner, the only other species known to have 4-spored asci, differs from the new species in having 3-trans-septate ascospores and a different chemistry with psoromic and conpsoromic acids in the thallus. Fissurina inabensis (Vain.) Nakan & Kashiw. differs from the new species in having small, submuriform ascospores, 21–30 × 10–15 µm.
Fissurina simplex has been collected from the evergreen forest of Kerala and from the moist deciduous forest of Karnataka.
Additional specimens examined. India: Karnataka: Anmod Ghat, 8 km from Anmod check post, Anmod-Goa road, 1974, P. G. Patwardhan & C. R. Kulkarni 74.2455, 74.3628. Kerala: Silent Valley, 1981, P. K. Sethy & S. P. Kekre 81.773 (AMH).
Fissurina sporolata B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561860
Fissurina sporolata, a Fissurinae speciebus typo subcontexta similibus ascosporis muriformibus majoribusque differt.
Typus: India, Karnataka, Hebri, 30 December 1980, P. G. Patwardhan & M. B. Nagarkar 80.510 (AMH—holotypus).
Thallus corticolous, brown, finely cracked, glossy, warty, rough.
Ascomata lirelline, 1·5–3·0 mm long, simple to rarely branched, concolorous with the thallus, immersed, terminally acute, structure of subcontexta-type. Disc narrow, epruinose. Exciple entire, non-carbonized, orange-brown, very thin at the base, convergent, often arising as fissures in swollen thalline margins (‘puffed’), covered by the raised thalline margin up to top, with a distinct upper corticiform layer. Hymenium hyaline, clear, 225–250 µm high, I−, KI−. Hypothecium hyaline, 20–25 µm high. Paraphyses simple. Periphysoids short, slightly warty at the tips. Asci 8-spored. Ascospores hyaline, muriform, ellipsoidal, multilocular, 42–57 × 17–25 µm, with a 5–7 µm thick halo, I+ blue tinge.
Chemistry. No lichen substances present.
Remarks. Fissurina marginata Staiger, the only other species with subcontexta-type of ascomatal structure, muriform ascospores and no lichen substances, differs by its smaller ascospores, 22–32 × 13–15 µm. The other species having subcontexta-type of ascomatal structure are Fissurina alboscripta (Coppins & P. James) Staiger, F. canlaonensis (Vain.) Staiger, F. crassilabra Mont. & Bosch., F. rufula (Mont.) Staiger, F. subcontexta (Nyl.) Nyl. and F. triticea (Nyl.) Staiger; these all have trans-septate ascospores.
Fissurina sporolata was found growing on a roadside tree trunk in the moist deciduous forest of Hebri in Karnataka, state of India.
Additional specimen examined. India: Karnataka: Hebri, 1980, P. G. Patwardhan & M. B. Nagarkar 80.634 (AMH).
Fissurina submonospora B. O. Sharma, Khadilkar & Makhija sp. nov
MycoBank No.: MB561861
Similis Fissurinae monosporae sed ascosporis majoribus differt.
Typus: India, Tamil Nadu, Upper Kodayar, 24 January 1983, P. G. Patwardhan & P. K. Sethy 83.187 (AMH—holotypus).
Thallus corticolous, brownish grey to dark brown, cracked, uneven, warty, glossy, delimited by the black hypothalloid region at the periphery.
Ascomata lirelline, short, 0·5–1·0 mm long, simple to branched, immersed, concolorous with the thallus, structure of dumastii-type. Disc slit-like. Exciple entire, non-carbonized, absent or indistinct at the base, orange-brown, convergent, covered by the thalline margin up to the top, with prosoplectenchymatous upper corticiform layer, studded with crystals. Hymenium hyaline, clear, 125–150 µm high. Hypothecium indistinct. Paraphyses simple. Periphysoids short, indistinctly warty at the tips. Asci 1-spored. Ascospores hyaline, muriform, 10–15 transverse and 4–5 longitudinal septa, 70–100 (–130) × 20– 50 µm, with 2·5–10·0 µm thick halo, I+ weakly blue.
Chemistry. No lichen compounds present.
Remarks. Fissurina monospora is morphologically similar but has larger ascospores, 185–200 × 65 µm. Fissurina undulata (Müll. Arg.) M. Nakan. & Kashiw. from Japan, also with 1-spored asci, differs in having smaller ascospores, 30–38 µm. Another species, Fissurina microcarpa (described above), also has monosporate asci and muriform ascospores but that species contains stictic acid.
The species grows in the moist evergreen forests of the Western Ghats of India.
Additional specimens examined. India: Karnataka: Anmod ghat, 3 km from Anmod check post, Anmod-Goa road, 1974, A. V. Prabhu & M. B. Nagarkar 74.2518; Jog falls-Sagar road, 6 km from Sagar, 1974, M. B. Nagarkar & P. G. Patwardhan 74.2844. Kerala: Cardamom hills, Devicolam-Kumily Road, on 98 km, 1976, C. R. Kulkarni 76.816; Silent Valley, 1981, P. G. Patwardhan & U. V. Makhija 81.870, 81.872; ibid., 1982, P. G. Patwardhan & M. B. Nagarkar 82.81. Tamil Nadu: Upper Kodayar, 1983, P. G. Patwardhan & P. K. Sethy 83.248, 83.261, 83.265; Agasthy hills, on Kakachi road, in moist evergreen forest, elev. approx. 1400 ft., 1984, P. G. Patwardhan & P. K. Sethy 84.61 (AMH).
Fissurina subnitidula (Nyl.) Staiger
Biblioth. Lichenol. 85: 153 ( 2002)—Graphis subnitidula Nyl. In Tuckerman, N. Amer. Lich., 2: 123 (1888).
Thallus corticolous, yellowish brown to brown, cracked, verrucose, slightly glossy, delimited by a distinct black hypothallus.
Ascomata lirelline, 0·5–1·0 mm long, concolorous with the thallus, simple, all over the thallus, curved, immersed to semi-emergent, terminally acute to obtuse, structure of dumastii-type. Disc narrow to broad, brown, epruinose. Exciple entire, non-carbonized, brown, convergent, indistinct at the base, covered with the thalline margin up to the top with thin pseudocortex, and studded with crystals. Hymenium hyaline, clear, 50–75 µm high, I−, K−. Hypothecium indistinct. Paraphyses simple with branched tips. Periphysoids absent. Asci 8-spored. Ascospores hyaline, 3-trans-septate, 10–15 ×5–7 µm with a 2·5–5·0 µm thick halo, I−.
Chemistry. No lichen compounds present.
Remarks. Fissurina subnitidula was previously known from Africa and has now been reported for the first time from India. It has been collected from the moist deciduous forest of Karnataka.
Specimens examined. India: Karnataka: Kumtha-Sirsi road, Devimane Ghat, rainforest, 1977, C. R. Kulkarni 77.113; Hosar Ghat, 1981, S. P. Kekre & M. B. Nagarkar 81.658, 81.701, 81.703 (AMH).
Fissurina sp. 1
Fig. 4. Fissurina sp. 1 (83.228, AMH), section of ascocarp. Scale = 50 µm. In colour online.
Thallus yellowish grey, cracked, slightly glossy, rough.
Ascomata lirelline, up to 0·5–1·0 mm long, immersed to slightly raised arising as the swelling which then cracks and gapes, intermingled, branched, creamish, irregularly curved, terminally obtuse, structure of incrustans-type. Disc sunken, creamish, epruinose. Exciple non-carbonized, entire, present at the base, orange-brown, convergent, covered by the thalline margin with thin, hyaline corticiform layer up to the top, studded with crystals. Hymenium hyaline, clear, 70–100 µm high, I−, KI−. Hypothecium indistinct. Paraphyses simple. Periphysoids short, sometimes warty at the tips. Asci 8-spored. Ascospores hyaline, submuriform, transversely 3–4-septate and vertically 1-septate, ellipsoidal, 7–12 × 5–7 µm, I+ blue-violet.
Chemistry. Norstictic acid present.
Remarks. Fissurina sp. 1, having incrustans-type of ascomatal structure, is distinguishable from F. incrustans Fée by the presence of norstictic acid and smaller ascospores; F. incrustans has rather larger ascospores, 15–25 (–28) × 7–10 (–13) µm, and does not produce any lichen compounds.
This material appears to be a new species but we reserve a final decision until we have had the opportunity to study more material.
Specimen examined. India: Tamil Nadu: Upper Kodayar, 1983, P. G. Patwardhan & P. K. Sethy 83.228 (AMH).
The lichen genus Hemithecium Trevis
After the reintroduction of the genus Hemithecium Trevis. by Staiger (Reference Staiger2002), the genus is so far represented by about 41 species worldwide, including 19 species from the Indian subcontinent (Staiger Reference Staiger2002; Nakanishi et al. Reference Nakanishi, Kashiwadani and Moon2003; Adawadkar & Makhija Reference Adawadkar and Makhija2005; Makhija & Adawadkar Reference Makhija and Adawadkar2005; Makhija et al. Reference Makhija, Dube, Adawadkar and Chitale2005; Archer Reference Archer2006, Reference Archer and McCarthy2009; Chitale et al. Reference Chitale, Makhija and Sharma2009; Jagadeesh & Sinha Reference Jagadeesh Ram and Sinha2009). The genus can be distinguished from the other genera of lirelline Graphidaceae by well-developed, convergent, non-carbonized and distinctly striate exciples, 1–8-spored asci, and hyaline, transversely septate or muriform ascospores, turning I+ blue or blue-violet.
As a result of our studies, three additional new species of Hemithecium have been discovered and five new combinations are proposed. A key for the identification of all 26 species of Hemithecium so far known from India is also provided.
Key to Indian Hemithecium species
1 Ascospores >100 µm in length ... 2
Ascospores <100 µm in length ... 8
2(1) Ascospores muriform ... 3
Ascospores trans-septate ... 7
3(2) Lichen substances absent; ascomata strongly raised, simple to rarely branched,1–7 mm long; exciple woody brown, 4–5 striate; asci 1–2-spored; ascospores 150–200 × 60– 75 µm ... H. multistriatum (Müll. Arg.) Chitale & Makhija
Lichen substances present ... 4
4(3) Salazinic acid present ... 5
Salazinic acid absent ... 6
5(4) Ascomata emergent, 1–9 mm long; exciple non-carbonized, 4–5 striate; ascospores 130–200 × 35–60 µm; only salazinic acid present ... ... H. salacinilabiatum (Patw. & C. R. Kulk.) Chitale & Makhija
Ascomata semi-emergent, 2–10 mm long; exciple non-carbonized, 2–3 striate; ascospores 112–145 × 37–52 µm; salazinic and protocetraric acids present ... ... H. longilirellatum B. O. Sharma & Khadilkar
6(4) Ascomata emergent, short, up to 1–10 mm long; exciple non-carbonized, striate; ascospores 112–209 × 30–66 µm; stictic acid present ... ... H. stictilabiatum (Patw. & C. R. Kulk.) Chitale & Makhija
Ascomata semi-emergent, 0·5–5 mm long; exciple non-carbonized, internally striate; ascospores 87–225 × 22–40 µm; norstictic acid present ... ... H. norlabiatum (Patw. & C. R. Kulk.) B. O. Sharma & Khadilkar
7(2) Ascomata 6–15 mm long, semi-emergent to distinctly emergent; simple to branched; non-carbonized exciple, with subverrucose margin, heavily crenate; 10–25-trans-septate ascospores, (45–)52–122 × 8–12 µm; constictic, norstictic and stictic acids present ... ... H. nagalandicum (Kr.P. Singh & G. P. Sinha) Adaw. & Makhija
Ascomata 2–9(–12) mm long, semi-emergent to distinctly emergent; sparsely branched; exciple dark blackish brown or slightly carbonized at the tips, 3-4 striate; 13–21-trans-septate ascospores, 49–112 × 7–14 µm; constictic, norstictic and stictic acids present ... H. amboliense Makhija & Dube
8(1) Ascospores <50 µm long ... 9
Ascospores >50 µm long ... 20
9 (8) Ascospores muriform ... 10
Ascospores trans-septate ... 14
10(9) Lichen substances absent; ascomata immersed, simple, short, 1–2 mm long; exciple non-carbonized, entire; ascospores muriform, 16–20 × 5–7 µm ... ... H. flavoalbum (Makhija et al.) B. O. Sharma & Khadilkar
Lichen substances present ... 11
11(10) Ascospores > 30 µm long ... 12
Ascospores < 30 µm long ... 13
12(11) Ascomata 1–8 mm long, simple to irregularly branched; exciple internally striate; ascospores 30–47 × 10–12 µm; stictic acid present ... ... H. kodayarense B. O. Sharma & Khadilkar
Ascomata 1–20 mm long, simple to irregularly branched, flexuouse; exciple indistinctly striate; ascospores muriform to submuriform, 20–40 × 10–15 µm; norstictic, stictic and constictic acids present ... ... H. flexile (Makhija et al.) B. O. Sharma & Khadilkar
13(11) Ascomata 1·0–7·5 mm long, simple to irregularly branched, flexuous; exciple entire or with 1–2 striae; ascospores 17–21 × 8–14 µm; constictic, stictic, and norstictic acids present ... H. microspermum Chitale et al.
Ascomata up to 2 cm long, simple to rarely branched; exciple woody brown, fan shaped, indistinctly striate, ascospores muriform, 20–25 × 8–9 µm; stictic, norstictic and constictic acids ... H. andamanicum B. O. Sharma & Khadilkar
14(9) Protocetraric acid present; ascomata 0·5–6·0 mm long, simple to branched, immersed; exciple entire; ascospores 9–12-septate, 29–42 × 4–6 µm; protocetraric, stictic and constictic acids present ... H. fulvescens Adaw. & Makhija
Protocetraric acid absent ... 15
15(14) Norstictic acid present ... 16
Norstictic acid absent ... 17
16(15) Ascomata 0·5–1·5 mm long, delicate, simple to radially or irregularly branched; exciple non-striate; ascospores 17–25 × 3–4 µm; stictic, consalazinic and norstictic acids present ... H. pulchellum Makhija & Adaw.
Ascomata 0·2–2·0 mm long, simple, triradiate or irregularly branched; exciple entire; ascospores 33–46 × 6–8 µm; norstictic and stictic acids present ... ... H. balaghatense Adaw. & Makhija
17(15) Stictic acid absent; ascomata 0·5–3·0 mm long, simple to branched, immersed; exciple entire, orange-brown, with internal striae; ascospores 21–29 × 4–5 µm; consalazinic acid, testacein A and testacein B present ... H. staigerae Adaw. & Makhija
Stictic acid present ... 18
18(17) Ascomata >10 mm long; ascomata 1–13 mm long, simple to branched, immersed; exciple entire; ascospores 21–33 × 3–4 µm; constictic and stictic acids present ... H. aphaneomicrosporum Makhija & Adaw.
Ascomata <10 mm long, mostly simple ... 19
19(18) Ascomata 3–7 mm long; exciple non-carbonized, yellowish to woody brown, entire, ascospores 6–7-trans-septate, 16–21 × 4–5 µm; stictic acid present ... ... H. croceum Makhija & Adaw.
Ascomata 1–5 mm long; exciple non-carbonized, brown, entire; ascospores 10–15-trans-septate, 30–42 × 7–10 µm; stictic and constictic acid present ... ... H. verrucosum B. O. Sharma & Khadilkar
20(8) Ascospores muriform ... 21
Ascospores trans-septate ... 22
21(20) Ascomata 2–6 mm long; exciple yellow to brown, simple to sparsely branched, 3–5 striate; ascospores 50–71 × 15–25 µm; stictic and constictic acids present ... ... H. epixanthum (Mont. & Bosch) Chitale & Makhija
Ascomata short, 1–4 mm long; exciple simple to sparsely branched, fan-shaped, woody brown, 5–8 striate; ascospores 42–63 × 8–17 µm; no lichen substances present ... H. flabillatum (Makhija et al.) B. O. Sharma & Khadilkar
22(20) Salazinic acid present; ascomata 1–4 mm long, immersed to slightly raised; exciple entire, woody brown; ascospores 63–88 × 6–8 µm; constictic, salazinic, and stictic acids present ... H. scariosum Makhija & Adaw.
Salazinic acid absent ... 23
23(22) Ascomata immersed in elevated wart-like structures (not in stroma), 0·3–1·5 mm long; exciple entire to striate, non-carbonized; ascospores 21–53 × 6–9 µm; constictic, norstictic and stictic acids present ... H. consociatum Makhija & Dube
Ascomata not in wart-like structures ... 24
24(23) Only norstictic acid present; ascomata 1–9 mm long, concolorous; exciple internally striate, occasionally slightly carbonized at the apex; ascospores 25–56 × 7–8 µm; only norstictic acid present ... H. norsticticum Makhija & Dube
Norstictic with other acids present ... 25
25(24) Ascomata 2–7 mm long, simple to dendroidally branched; exciple entire to indistinctly striate; ascospores 30–40(–60) × 6–9 µm; constictic, norstictic (trace), and stictic acids present ... H. nakanishianum (Patw. & C. R. Kulk.) Makhija & Dube
Ascomata 0·5–10·0 mm long, simple or branched; exciple with 2–4 internal striae; ascospores 25–67(–80) × 11–12 µm; constictic, norstictic and stictic acids present ... H. aphanes (Mont. & Bosch) M. Nakan. & Kashiw.
The Species
Hemithecium andamanicum (Makhija, Adaw. & Patw.) B. O. Sharma & Khadilkar comb. nov
MycoBank No.: MB561862
Graphina andamanica Makhija, Adaw. & Patw., Biovigyanam 18(1): 22 (1992); type: India, Andaman Islands, South Andaman, Chidya Tapu, 15 February 1985, P. K. Sethy & P. G. Patwardhan 85.67 (AMH—holotypus!).
(Fig. 5A)
Fig. 5. Hemithecium species, habitus. A, H. andamanicum (85.67—holotype, AMH); B, H. flabillatum (85.2384—holotype, AMH); C, H. flavoalbum (85.47—holotype, AMH); D, H. flexile (86.364—holotype, AMH); E, H. kodayarense (84.99—holotype, AMH); F, H. longilirillatum (85.2771—holotype, AMH). Scales: A–F = 1 mm. In colour online.
Remarks. Hemithecium andamanicum is characterized by: a greyish to white greyish, smooth, glossy thallus; pale brown, semi-emergent, long, up to 2 cm long, simple to rarely branched, thin ascocarps; proper exciple present below, non-carbonized, woody brown, fan shaped, indistinctly striate, encircled by the thalline margin; hyaline, clear hymenium; 8-spored asci; hyaline, muriform, 20–25 × 8–9 µm ascospores; stictic, norstictic and constictic acids in the thallus.
Specimens examined. India: Andaman Islands: South Andaman, Pongibalu, Kala Pahad, 1985, P. G. Patwardhan & P. K. Sethy 85.1182, 85.1184, 85.1189; Chidya Tapu, 1985, P. K. Sethy & P. G. Patwardhan 85.65 (AMH).
Hemithecium flabillatum (Makhija, Adaw. & Patw.) B. O. Sharma & Khadilkar comb. nov
MycoBank No.: MB561870
Graphina flabillata Makhija, Adaw. & Patw., Biovigyanam 18(1): 24 (1992); type: India, Andaman Islands, Middle Andaman, Betapur Range, Pitcher Nala, in evergreen forest, 26 December 1985, P. K. Sethy & P. G. Patwardhan 85.2384 (AMH—holotypus!).
(Fig. 5B)
Remarks. The species is characterized by: a brown, rough, delimited thallus; simple to rarely branched, short, 1–4 mm long ascocarps; white-pruinose disc; fan shaped, non-carbonized, woody brown, striate exciple; hyaline, clear hymenium; 8-spored asci; muriform, 42–63 × 8–17 µm ascospores; no lichen substances in the thallus. Hemithecium rufopallidum (Vain.) Staiger is closely related to H. flabillatum, but differs in having 2–6-spored asci and large ascospores, 50–86 × 17–30 µm.
Hemithecium flabillatum appears to be endemic to the Andaman Islands of India, where it grows on exposed tree trunks on roadsides.
Specimens examined. India: Andaman Islands: Middle Andaman, Betapur Range, Pitcher Nala, in evergreen forest, 1985, P. G. Patwardhan & P. K. Sethy 85.2384, 85.2385, 85.2401, 85.2411 (AMH).
Hemithecium flavoalbum (Makhija, Adaw. & Patw.) B. O. Sharma & Khadilkar comb. nov
MycoBank No.: MB561867
Graphina flavoalba Makhija, Adaw. & Patw., Biovigyanam 18(1): 24 (1992); type: India, Andaman Islands, South Andaman, Chidya Tapu, 15 February 1985, P. G. Patwardhan 85.47 (AMH—holotypus!).
(Fig. 5C)
Remarks. The species is characterized by: a grey-white, smooth thallus; immersed, simple, short, 1–2 mm long ascocarps; non-carbonized, entire, exciple; hyaline, clear hymenium; 8-spored asci; muriform, 16–20 × 5–7 µm ascospores; no lichen substances in the thallus.
Hemithecium flavoalbum was found growing on tree trunks and is known from the type collection only.
Hemithecium flexile (Makhija, Adaw. & Patw.) B. O. Sharma & Khadilkar comb. nov
MycoBank No.: MB561868
Graphina flexilis Makhija, Adaw. & Patw., Biovigyanam 18(1): 25 (1992); type: India, Andaman Islands, North Andaman, Diglipur forest, Millangram in evergreen forest, 3 January 1986, M. B. Nagarkar & P. G. Patwardhan 86.364 (AMH—holotypus!).
(Fig. 5D)
Remarks. This species is characterized by: a greyish white, cracked, smooth, glossy, delimited thallus; lirellae, short to long and up to 1–20 mm long, simple to irregularly branched, flexuose, concolorous with the thallus, semi-emergent to moderately emergent, terminally acute ascocarps; convergent, non-carbonized, woody brown, indistinctly striate exciple; hyaline, clear hymenium; 8-spored asci; muriform to submuriform, hyaline, ascospores 20–40 × 10–15 µm, and norstictic, stictic and constictic acids in the thallus.
Remarks. The species is found also on the mainland of India in the evergreen forest of the Western Ghats.
Additional specimens examined. India: Andaman Islands: North Andaman, Diglipur Range, kalara, in evergreen forest, 1986, M. B. Nagarkar & P. K. Sethy 86.394, 86.423, 86.443; Tugapur Range, Mayabandar, Pather Tikri, in moist deciduous forest, 1985, M. B. Nagarkar & P. G. Patwardhan 85.2575, 85.2594, 85.2692; ibid. P. G. Patwardhan & P. K. Sethy 85.2587; Middle Andaman, Parlobjig Islands, 1985, M. B. Nagarkar & P. K. Sethy 85.2306; Betapur Range, Dhaninala, 1985, M. B. Nagarkar & P. G. Patwardhan 85.2525; South Andaman, Shol Bay, near Wright Myo, 1986, P. K. Sethy & P. G. Patwardhan 86.692. Karnataka: Jog-Honawar road, 10 km from Jog, 1974, C. R. Kulkarni 74.2779, 74.2780; Sitanadi, Agumbe-Udipi road, 1974, P. G. Patwardhan & M. B. Nagarkar 74.3054, 74.3055; Khushalnagar, Coorg, 1974, M. B. Nagarkar & A. V. Prabhu 74.3284; Londha-Dharwar road, 5 km from Londha, 1974, A. V. Prabhu & M. B. Nagarkar 74.2540, 74.2541; Devimane Ghat, Sirsi-Kumtha road, 1974, P. G. Patwardhan & M. B. Nagarkar 74.2614. Kerala: Anamalai hills, Sholayar, near dam in the ravine, 1976, C. R. Kulkarni, 76.326; Ponmudi, 1973, P. G. Patwardhan & M. B. Nagarkar 73.2630, 73.2643, 73.2673, 73.2658 (AMH).
Hemithecium kodayarense B. O. Sharma & Khadilkar sp. nov
MycoBank No.: MB561863
Similis Hemithecii nagalandici, sed ascoporis minoribus et acido stictico continente differt.
Typus: India, Tamil Nadu, Upper Kodyar, Agasthi Hills, in moist deciduous forest elev. c. 1400 m, 20 February 1984, P. G. Patwardhan & P. K. Sethy 84.99 (AMH—holotypus).
Thallus crustose, corticolous, epiphloeodal, creamish grey, cracked, smooth, dull to slightly rough, delimited by thin black hypothallus.
Ascomata lirelline, 1–8 mm long, simple to irregularly branched, concolorous with the thallus, emergent, terminally round. Exciple convergent, non-carbonized, striate, covered by the raised thalline margin. Hymenium hyaline, clear, 100–125 µm in height. Paraphyses simple. Asci 8-spored. Ascospores muriform to submuriform, hyaline 30–47 × 10–12 µm, weakly I+ violet or almost I−.
Chemistry. Stictic acid present.
Remarks. The new species can be differentiated from the externally somewhat similar Hemithecium nagalandicum (Kr. P. Singh & G. P. Sinha) Adawadkar & Makhija by its muriform, smaller ascospores and the presence of stictic acid only (H. nagalandicum has transversely septate, large ascospores, 52–122 × 8–12 µm and contains norstictic, constictic and stictic acids). Hemithecium radicicola (A. W. Archer) A. W. Archer, a species from Australia having only stictic acid and muriform ascospores, 35–40 µm long, differs from the new species by its initially fissurine lirellae and the absence of a proper exciple.
Additional specimens examined. India: Tamil Nadu: Upper Kodyar, Agasthi Hills, in moist deciduous forest elev. c. 1400 m, 1984, P. G. Patwardhan & P. K. Sethy 84.137, 84.138 (AMH).
Hemithecium longilirellatum B. O. Sharma & Khadilkar sp. nov
MycoBank No.: MB561864
Species nova distinctus inter omnibus Hemithecii speciebus ob praesentia acidarum salazinici et protocetrarici.
Typus: India, Andaman Islands, North Andaman, Mayabandar Range, Kaichi Nala, moist deciduous forest, 30 December 1985, P. G. Patwardhan & P. K. Sethy 85.2771 (AMH—holotypus).
Thallus crustose, corticolous, epiphloeodal, brown, smooth, delimited by a black thin, hypothallus.
Ascomata lirelline, 2–10 mm long, pale brown, simple to branched, irregular, slender, semi-emergent, terminally rounded. Exciple non-carbonized, brown, striate, present below, encircled by the thalline margin with pseudocortex and crystals. Hymenium hyaline, clear, 150–200 µm high. Paraphyses simple. Asci 1-spored. Ascospores hyaline, muriform, 112·5–145·0 × 37·5–52·5 µm, weakly I+ violet.
Chemistry. Salazinic and protocetraric acids present.
Remarks. Hemithecium longilirellatum is characterized by: long, irregularly curved lirellae; striate exciple; 1-sporate asci; muriform, large, ascospores; salazinic and protocetraric acids in the thallus.
The new species can be differentiated from the other species of Hemithecium having more or less similar morphological features and large, muriform ascospores: Hemithecium orizaeforme (Fée) Staiger, H. chlorocarpum (Fée) Trevis., H. salazinilabiatum (Patw. & C. R. Kulk.) Chitale & Makhija, H. stictilabiatum (Patw. & C. R. Kulk.) Chitale & Makhija and H. multistriatum (Müll. Arg.) Chitale & Makhija all have differences in their chemistry. Hemithecium orizaeforme contains norstictic acid, H. chlorocarpum has no lichen substances, H. salazinilabiatum has salazinic acid, H. stictilabiatum contains stictic acid, and H. multistriatum has no lichen acids.
The species has been collected in the moist deciduous forest areas of the Andaman Islands.
Additional specimen examined. India: Andaman Islands: North Andaman, Mayabandar Range, Kaichi Nala, moist deciduous forest, 1985, P. G. Patwardhan & P. K. Sethy 85.2790 (AMH).
Hemithecium norlabiatum (Patw. & C. R. Kulk.) B. O. Sharma & Khadilkar comb. nov
MycoBank No.: MB561866
Graphina norlabiata Patw. & C. R. Kulk., Biovigyanam 5: 6 (1979); type: India, Karnataka, Agumbe, 2 km on Koppa road, in rainforest, 17 December 1974, C. R. Kulkarni 74.3243 (AMH—holotypus!).
(Fig. 6A)
Fig. 6. Hemithecium species, habitus. A, H. norlabiatum (74.3243—holotype, AMH); B, H. verrucosum (04.281—holotype, AMH); Scales A & B = 1mm. In colour online.
Fig. 7. Hemithecium, sections of ascocarps. A, H. kodayarense (84.99—holotype, AMH); B, H. longilirellatum (85.2771—holotype, AMH); C, H. verrucosum (04.281—holotype, AMH). Scales: A–C = 100 µm. In colour online.
Remarks. The species is characterized by: a citrine green to pale olivaceous buff, smooth to uneven, glossy, cracked thallus; 0·5–5·0 mm long, simple, curved, flexuous, semi-emergent, white to dull, pale brownish or concolorous ascocarps; non-carbonized, brown, indistinctly striate exciple; a hyaline, clear, hymenium; 1–2-spored asci; hyaline, muriform, 87–225 × 22–40 µm ascospores; norstictic acid in the thallus.
Additional specimens examined. India: Karnataka: Devimane Ghat, 1974, M. B. Nagarkar & A. V. Prabhu 74.3244; Someshwar, 1982, P. G. Patwardhan 82.711; near Agumbe, 1977, P. G. Patwardhan 77.411, 77.414, 77.415; Anmod Ghat, 1975, C. R. Kulkarni & M. B. Nagarkar 75.564; Hebri, 74.3020, 74.3021, 74.3244. Kerala: Silent Valley, 1976, M. B. Nagarkar 76.221, 76.222, 76.223, 76.233, 76.234; Chalkudi, 40 kms, 1976, A. V. Prabhu & M. B. Nagarkar 76.293; Attapadi, 1976, M. B. Nagarkar 76.212, 76.213 (AMH).
Hemithecium verrucosum B. O. Sharma & Khadilkar sp. nov
MycoBank No.: MB561865
Similis Hemithecii aphaneomicrospori, sed thallo verruculoso et ascosporis majoribus differt.
Typus: India, Karnataka, Dandeli forest, 5 October 2004, U. V. Makhija 04.281(AMH—holotypus).
Thallus crustose, corticolous, epiphloeodal, creamish, cracked, warty, verrucose, shiny, delimited with black hypothallus.
Ascomata lirelline 1–5 mm long, simple, rarely branched, emergent, wavy, curved, raised, terminally acute. Disc reddish brown, broad, epruinose. Exciple brown, entire, non-carbonized, present at the base, covered by the thick thalline margin with pseudocortex. Hymenium hyaline, clear, 100–125 µm high. Paraphyses simple. Asci 8-spored. Ascospores transversely septate, 10–15 septa, 30–42 ×7–10 µm, I+ purple.
Chemistry. Stictic and constictic acids present.
Remarks. The new species is characterized by its verruculose thallus, simple, emergent lirellae, trans-septate ascospores, and stictic and constictic acids in the thallus.
The species can be compared with Hemithecium aphaneomicrosporum Makhija & Adawadkar in having more or less similar exciple characters, transversely septate ascospores and chemistry, but it differs from the new species in having slightly smaller ascospores, 21–33 × 3–4 µm.
Additional specimens examined. India: Karnataka: Dandeli forest, 2004, U. V. Makhija 04.294, 04.265 (AMH).
We are grateful to the Department of Science and Technology (DST), Government of India, New Delhi, for financial support. We also thank two anonymous reviewers for their helpful comments and valuable suggestions.
