INTRODUCTION
The lichenized fungal genus
Fissurina Fée, with mostly slit-like lirellae, belongs in Graphidaceae in the order Ostropales (
Staiger 2002;
Archer 2009;
Lumbsch et al. 2014;
Miadlikowska et al. 2014). The genus was established by Fée (1824) for species with fissurine, lirelline ascomata and ascospores with a halo. The name
Fissurina was used by Vainio (1921) at the subgeneric level or for sections of the genus
Graphis by Redinger (1935) and
Zahlbruckner (1923). With the description of
Fissurina quadrispora Kalb & Hafellner, the genus was again resurrected (
Kalb & Hafellner 1992). Following this,
Fissurina was formally reintroduced by
Staiger & Kalb (1999), and phylogenetic studies (
Staiger et al. 2006;
Mangold et al. 2008;
Rivas Plata et al. 2012,
2013) showed that it was distinct among Graphidaceae, forming a separate subfamily with some related genera.
Fissurina comprises more than 100 species worldwide, with a mainly tropical to subtropical distribution. The genus is characterized by a pale to yellow-brown to olive green (rarely whitish), mostly smooth and glossy thallus, mostly fissurine, simple to branched, immersed to prominent lirellae with uncarbonized or rarely carbonized proper exciple, a mostly clear non-amyloid hymenium, 1-8-spored asci, hyaline, oval or narrowly to broadly ellipsoid, trans-septate to muriform, amyloid to non-amyloid, often thick-walled, mostly halonate ascospores, and a
Trentepohlia photobiont (
Archer 2009;
Rivas Plata et al. 2012;
Sharma et al. 2012;
Joshi et al. 2013).
Previous reports of
Fissurina from China were historically listed under other genus names, such as:
Fissurina micromma (Zahlbr.) Aptroot [as
Phaeographina micromma Zahlbr. by Zahlbruckner (
1933,
1940),
Lamb (1963), Wang-Yang & Lai (1973)],
Fissurina adscribens (Nyl.) Z.F. Jia & Lücking [as
Graphina adscribens (Nyl.) Müll. Arg. by
Thrower (1988)],
Fissurina incrustans Fée [as
Graphina incrustans (Fée) Müll. Arg., by
Thrower (1988)],
Fissurina radiata Mont. [as
Graphis radiata (Mont.) Nyl. by
Aptroot & Sparrius (2003)],
Fissurina tachygrapha (Nyl.) Staiger [as
Graphis tachygrapha Nyl. by
Thrower (1988)], and
Fissurina triticea (Nyl.) Staiger [as
Graphis triticea Nyl. by
Aptroot & Sparrius (2003)]. Later, several species of
Fissurina were added (
Meng et al. 2011;
Kalb & Jia 2014;
Jia & Lücking 2017a,
2017b).
Fissurina canlaonensis (Vain.) Staiger, the name reported by
Meng et al. (2011), is now a synonym of
F. consentanea Nyl., and
Fissurina isidiata Z.F. Jia (
Jia et al. 2012) was found to be a synonym of
Platythecium dimorphodes (Nyl.) Staiger. In the present paper, three new records of
Fissurina from China are given; hence there are 16 species of this genus confirmed for the country.
1 MATERIALS AND METHODS
The specimens are deposited in HMAS-L, LCU, KUN, BGBM, hb.-Kalb and W. Two dissecting microscopes (OLYMPUS SZ51 & SZX12; TECH XTS-30D) and two microscopes (OLYMPUS BX53; NiKON Eclipse-55i) were used for morphological and anatomical studies. Measurements and illustrations were taken from manual cross-sections of lirellae in tap water. Amyloidity of the ascospores was tested using Lugol’s solution. Spot tests with KOH (20%) were performed on the thallus surface and on thin thallus sections. Lichen substances were further analyzed and identified by thin-layer chromatography using solvent systems C & A (
Orange et al. 2001;
Jia & Wei 2016).
Five lirellae types were defined for the characterization of the ascomatal structures:
comparilis-type,
dumastii-type,
globulifica-type,
incrustans-type and
subcontexta-type, based on their transition and formation of fruiting bodies, following
Staiger (2002).
2 THE SPECIES
Fissurina adscribens (Nyl.) Z.F. Jia & Lücking
MycoKeys 25: 15, 22 (2017).
≡ Graphis adscribens Nyl., Bull. Soc. Linn. Normandie, sér. 2 2: 117 (1868). - Graphina adscribens (Nyl.) Müll. Arg., Hedwigia 31: 284 (1892).
= Graphina olivascens Zahlbr., Feddes Repert. Spec. Nov. Regni Veg. 31: 215 (1933).
Thallus corticolous, crustose, surface grey to olive, waxy and smooth. Ascomata lirelliform, elongate, single and rarely branched, 1-4mm long and 0.2-0.35mm wide, structure of dumastii-type; disc closed, slit-shaped; proper margin unconspicuous, concolorous with thallus; proper exciple not carbonized; hymenium colorless, clear, at most 150μm high; paraphyses unbranched; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, muriform, 5-7 transverse and 0-3 longitudinal septa, 20-30×8-10μm, halonate, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina adscribens is an eastern paleotropical species reported from Hunan [as
Graphina olivascens, Zahlbruckner
1930,
1932;
Wei 1991; synonymized by
Jia & Lücking (2017b)]. Unfortunately, the material reported from Hong Kong as
Graphina adscribens (Trower 1988: 92) could not be studied but based on its reported chemistry (stictic and constictic acids) and larger ascospores (30-35×5-10μm) possibly represents
Acanthothecis dialeuca (Kremp.) Staiger & Kalb, a species morphologically similar to
Fissurina.
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 420-450m, 1-4 June 2007, J. Li FJ1016, FJ1045, FJ1216, Q.F. Meng FJ1051-1, FJ1165; Wuyishan City, Tongmu, 690m, 24 July 2016, Z.F. Jia FJ16397; Wuyishan City, Longchuan, 450m, 24 July 2016, Q.D. Wang FJ16312 (LCU). Hainan, Qiongzhong County, Mt. Limushan, 730m, 25 September 2008, J. Li HN081239 (HMAS-L). Hunan, Wugang City, Mt. Yunshan, 1 100m, 9 August 1917, Handel-Mazzetti 11220 (W), Mt. Yunshan, 1 050- 1 140m, 28 April 2018, Z.F. Jia HNX18044, HNX18062 & HNX18063 (LCU); Sangzhi County, Mt. Taipingshan, 1 500m, 19 August 1997, J.B. Chen, D.P. Wang & S.L. Wang 9497 (HMAS-L); Zhangyi County, Mt. Mangshan, Guizizhai, 1 200m, 12 September 2002, Y.M. Jiang & J. Yang M047 (HMAS-L).
Fissurina baishanzuensis Kalb & Z.F. Jia
Phytotaxa 189(1): 148 (2014).
Thallus corticolous, crustose, surface uneven, grey-green to olive. Ascomata lirelliform, flexuose, single, sparsely branched, erumpent to prominent, with lateral thalline margin, 1-5mm long and 0.3-0.6mm wide, structure of incrustans-type; disc concealed; proper margin distinct, visible as thick, white labia; labia entire, with lateral thalline margin that often splits off, light olive-green to grey-green; proper exciple not carbonized; hymenium 100-120µm high, colorless, clear; paraphyses unbranched, apically smooth; periphysoids present along the slit, 10-20µm long, with irregular surface but not spinulose; asci fusiform, 8-spored; ascospores hyaline, broadly ellipsoid to subglobose, muriform, 1-3 transverse and 1-2 longitudinal septa, 10-12(-15)×9-10µm, thick-walled, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina baishanzuensis is similar to
F. instabilis (Nyl.) Nyl., but the latter is easily distinguished by its I+ blue (amyloid) spores. The species is collected on the trunk of rather young trees near the edge of a rather open, mixed mountain rainforest; it is currently considered as an endemic species to China, only reported from Baishanzu National Nature Reserve, Qingyuan County, Zhejiang (
Kalb & Jia 2014).
Specimens examined: China. Zhejiang, Qingyuan County, Mt. Baishanzu, Baishanzu Protection Station, 1 300-1 500m, 11 October 2010, K. Kalb, Z.S. Sun & Z.F. Jia 38639 (HMAS-L), 38609 & 38618 (hb. Kalb).
Fissurina cingalina (Nyl.) Staiger
Biblioth. Lichenol. 85: 128 (2002).
≡ Graphis cingalina Nyl., Acta Soc. Sci. Fenn. 26(10): 21 (1900). - Graphina cingalina (Nyl.) Zahlbr., Cat. Lich. Univers. 2: 401 (1923).
Thallus corticolous, crustose, splitting, surface grey to grey-greenish, smooth. Ascomata lirelliform, elongate, single, often branched, immersed, 1-3.5mm long and 0.2-0.4mm wide, structure of dumastii-type; disc closed, slit-shaped; proper margin unconspicuous, concolorous with thallus; proper exciple not carbonized; hymenium colorless, clear, 100-130μm high; paraphyses simple; periphysoids absent; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, often wider in one end, muriform, 5-9 transverse and 1-3 longitudinal septa, 20-35×10-16μm, halonate, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina cingalina, an eastern paleotropical species, is similar to
F. adscribens, but differs by the splitting cortex and wider ascospores. It is also similar to
F. columbina (Tuck.) Staiger, but the latter is distinguished by the erumpent and irregularly branched lirellae and the I+ violet ascospores. The species grows on the trunk of trees in rather open, mixed mountain rainforest in subtropical areas of China. The previously reported materials from Hunan by
Meng et al. (2011) turned out to represent
Fissurina adscribens.
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 300-600m, 01-03 June 2007, J. Li FJ1002, FJ1106, FJ1154, FJ1184, and Q.F. Meng FJ853, FJ938, FJ1292 (LCU). Yunnan, Gejiu City, Manhao, Malongdi, 1 280m, 23 September 2011, Z.F. Jia 11-480, 11-487 (LCU).
Fissurina columbina (Tuck.) Staiger
Biblioth. Lichenol. 85: 130 (2002).
≡ Graphis columbina Tuck., Syn. N. Amer. Lich. (Boston) 2: 123 (1883). -Graphina columbina (Tuck.) M. Wirth & Hale, Smithson. Contr. Bot. 40: 36 (1978). -Phaeographina columbina (Tuck.) Zahlbr., Cat. Lich. Univers. 2: 436 (1923).
Thallus corticolous, crustose, surface grey-greenish to brownish, smooth. Ascomata lirelliform, elongate, single to irregularly branched, erumpent, 0.5-4mm long and 0.1-0.3mm wide, structure of dumastii-type; disc closed, with thin, gently sloping labia (roof-like); proper margin unconspicuous, concolorous with thallus; proper exciple not carbonized; hymenium colorless, clear, 100-120μm high; paraphyses simple, ±interwoven, with brownish slightly thickened tips, moderately to distinctly conglutinated; periphysoids absent; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, often wider in one end, muriform, 5-9 transverse and 1-3 longitudinal septa, 20-30×10-15μm, halonate, I+ voilet.
Chemistry: Psoromic acid.
Remarks:
Fissurina columbina is similar to
F. adscribens but differs by the erumpent and irregularly branched lirellae, roof-like labia and I+ violet ascospores. The species is a corticolous species known also from North and South America (
Staiger 2002), and it was previosly reported from China (Guizhou, Hainan and Hunan;
Meng et al. 2011.
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 420-610m, 1-3 June 2007, J. Li FJ883, FJ884, FJ919, FJ921, FJ1003, FJ1049, FJ1095, FJ1126, FJ1143, FJ1210, FJ1202, Q.F. Meng FJ1107, FJ1140, FJ1167, FJ1171, FJ1176; Wuyishan City, Tongmu, 690m, 24 July 2016, J. Li FJ16372, FJ16379; Wuyishan City, Longchuan, 330m, 24 July 2016, Z.F. Jia FJ16294 (LCU). Guizhou, Tongren City, Mt. Fanjingshan, 1 570m, 5 September 1963, J.C. Wei 0790; 1 200m, 1 September 1963, J.C. Wei 0610 (HMAS-L). Hainan, Ledong County, Mt. Jianfengling, Mingfenggu, 1 080m, 14 May 2007, Q.F. Meng M131; Changjiang County, Mt. Bawangling, Yajia, 800m, 5 October 2008, J. Li HN081494; Wuzhishan City, Mt. Wuzhishan, 720-730m, M. Liu HN09027, HN09339 (HMAS-L). Hunan, Sangzhi County, Mt. Bagongshan, 1 500m, 20 August 1997, J.B. Chen, D.P. Wang & S.L. Wang 9699 (HMAS-L).
Fissurina consentanea Nyl.
Fig. 1A, B
Fig. 1 A, B: Fissurina consentanea (A: Habit; B: Asci containing ascospores with transverse septa. LCU, Z.F. Jia HN014); C, D: Fissurina globulifica (C: Habit; D: Asci containing ascospores muriform. LCU, J. Li FJ1117-3); E, F: Fissurina insidiosa (E: Habit; F: Asci containing ascospores with transverse septa and I-. LCU, Z. F. Jia FJ16106). Bars: A, C, E=1mm; B, D, F=20μm. |
Full size|PPT slide
Lich. Nov. Zeland. (Paris): 126 (1888).
≡ Graphis consentanea (Nyl.) Nyl., in Hue, Nouv. Arch. Mus. Hist. Nat., Paris, sér. 3, 3: 164 (1891).
= Fissurina canlaonensis (Vain.) Staiger, Biblthca Lichenol. 85: 161 (2002). - Graphis canlaonensis Vain., Ann. Acad. Sci. Fenn., ser. A 15(6): 261 (1921). - Hemithecium canlaonense (Vain.) A.W. Archer [as 'canlaonensis'], Syst. Biodiv. 5(1): 17 (2007).
Thallus corticolous, crustose, surface grey to grey-greenish, uneven to verrucose. Ascomata lirelliform, elongate, single, rarely branched, erumpent to prominent, 1-3mm long and 0.2-0.7mm wide, structure of subcontexta-type; disc concealed, with distinct thick labia (hemithecioid) mostly covered by nearly complete thalline margin, concolorous with thallus; proper exciple uncarbonized; hymenium colorless, clear, 90-120μm high; paraphyses simple, with brownish slightly thickened tips; periphysoids present; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, 3-septate, 15-25×8-12μm, halonate, I-.
Chemistry: Stictic acid.
Remarks:
Fissurina consentanea is similar to
F. triticea (Nyl.) Staiger, but the latter differs in having exposed labia covered by a lateral thalline margin only and by I+ weakly violet-blue ascospores.
Staiger (2002) listed
F. consentanea as a synonym of
F. triticea, but the two species are here accepted separate on account of these differences. An eastern paleotropical taxon (
Staiger 2002;
Galloway 2007), this is a new record for China.
Fissurina canlaonensis (Vain.) Staiger, as new combination based on
Graphis canlaonensis Vain., was reported by
Staiger (2002) and
Galloway (2007); later, the taxon was transferred to the genus
Hemithecium as
H. canlaonense (Vain.) A.W. Archer (
Archer 2007). Based on thallus structure,
subcontexta- type ascomata, and ascospore size [15-25×8-14μm, by
Redinger (1933); 12-16×5-8μm, by
Patwardhan & Kulkarni (1978); 15-26×8-14μm, by
Staiger (2002)] with I-, and the presence of stictic acid, we propose this name as new synonym of
Fissurina consentanea Nyl.
The specimens Wei 0762 & 0763 (deposited in HMAS-L), reported from Guizhou as Fissurina canlaonensis (Meng et al. 2011: 160) and Hemithecium canlanonensis (Vain.) A.W. Archer (Jia & Wei 2016: 145), are likely incorrect identifications; due to the reported absence of lichen compounds, so the materials need to be further studied. The most likely identification is F. baishanzuensis Kalb & Ze Feng Jia, but F. rufula (Mont.) Staiger is also a possibility.
Specimens examined: China. Hainan, Qiongzhong County, Mt. Limushan, 670m, 25 September 2008, Z.F. Jia HN080611 (HMAS-L), Z.F. Jia HN014 (LCU).
Fissurina dumastii Fée
Essai Crypt. Exot. (Paris): 59 (1825) [1824].
≡ Graphis dumastii (Fée) Spreng., Syst. Veg., Edn 16 4(1): 254 (1827).
= Graphis glauca Müll. Arg., Bull. Herb. Boissier 3(1): 58 (1893). - Fissurina glauca (Müll. Arg.) Staiger, Biblioth. Lichenol. 85: 159 (2002).
Thallus corticolous, crustose, surface smooth to uneven, olive-green. Ascomata lirelliform, elongate, short to elongate, erumpent, single to sparsely branched, 0.5-4mm long and 0.2-0.3mm wide, structure of dumastii-type; disc concealed, with thin but distinct labia (slit-like), covered by thalline margin; proper exciple uncarbonized; hymenium colorless, clear, 90-110μm high; paraphyses simple, apically spinulose; periphysoids present; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, 3-septate, 10-20×6-10μm, halonate, I+ weakly violet-blue.
Chemistry: No lichen substances detected.
Remarks:
Fissurina dumastii is distinguished by its olive-green thallus,
dumastii-type lirellae with slit-like labia, apically spinulose paraphyses, 3-septate ascospores and absence of secondary substance. No other species with this combination of characters is known in the genus. The species has a pantropical distribution and was previously reported from Queensland, New South Wales, Indonesia, Solomon Islands, Brazil and Vietnam (
Staiger 2002;
Archer 2006,
2009;
Joshi et al. 2013). In China, it was previously reported from Hong Kong (
Aptroot & Seaward 1999;
Aptroot & Sipman 2001, as
Graphis dumastii), Taiwan (
Aptroot & Sparrius 2003, as
Graphis dumastii), and Fujian and Hainan (
Meng et al. 2011).
Staiger (2002) reported
Fissurina glauca (ascospores 18-20×8-10µm) and
F. dumastii (ascospores 11-18×5-6µm), but suggested that the former could be synonym of the latter, which is here confirmed.
Specimens examined: China. Fujian, Wuyishan City, Sangang, 810m, 27 March 2004, Z.F. Jia FJ305; Jianou City, Mt. Wanmulin, 260m, 29 March 2004, Z.F. Jia FJ369 (LCU). Guangxi, Wuming County, Mt. Damingshan, 1 500m, 21 May 2015, J. Li GX15400 (LCU). Hainan, Ledong County, Mt. Jianfengling, 1 020m, 13 May 2007, Q.F. Meng M103; 1 000m, 14 May 2007, Q.F. Meng M151 (HMAS-L); Ledong County, Jiaxi, 400m, 22 July 2009, J. Li HN238 (HMAS-L). Hong Kong, Pok Fu Lam, Lung Fu shan, 200m, 6 July 2000, Sipman & Aptroot 45049; Lantau North Country Park, Ngong Ping, 400-450m, 8 July 2000, Sipman & Aptroot 45131 (BGBM).
Fissurina elaiocarpa (A.W. Archer) A.W. Archer
Telopea 11: 71 (2005).
≡ Graphina elaiocarpa A.W. Archer, Mycotaxon 77: 167 (2001).
= Fissurina marginata Staiger, Biblioth. Lichenol. 85: 144 (2002).
Thallus corticolous, crustose, surface slightly subtuberculate and shiny, olive-green. Ascomata lirelliform, elongate, single, prominent to sessile, 1-6mm long and 0.4-0.7mm wide, structure of subcontexta-type; disc mostly concealed; labia orange-brown, almost completely covered thalline margin (hemithecioid); proper exciple uncarbonized; hymenium colorless, clear, 120-180μm high; paraphyses simple, warty at the tips; periphysoids absent; asci cylindrical, 8-spored; ascospores hyaline, ovoid, muriform, 3-4 transverse and 2-3 longitudinal septa, 20-30×12-18μm, weakly halonate, I+ violet-blue.
Chemistry: No lichen substances detected.
Remarks:
Fissurina elaiocarpa is similar to
Fissurina subundulata Kalb & Z.F. Jia, but the latter differs in having prominent to sessile lirellae, whitish labia with a lateral to almost complete thalline margin, and I- ascospores.
F. marginata Staiger is a synonym of
F. elaiocarpa (
Archer 2005).
Meng et al. (2011) reported this species from China as
F. marginata, collected in a primitive mountain forest on Mt. Fanjingshan (Guizhou). The species has a worldwide distribution, reported from Africa, Australia, South and Western America, Western Indian Ocean and Vietnam (
Staiger 2002;
Archer 2005,
2006;
Joshi et al. 2013).
Specimens examined: China. Fujian, Wuyishan City, Mt. Longjing, 270m, 22 July 2016, Z.F. Jia FJ16106 (LCU). Zhejiang, Pan’an County, Mt. Dapanshan, 500m, 14 October 2010, Z.F. Jia 10-217 (LCU). Guozhou, Tongren City, Mt. Fanjingshan, 1 500m, 24 August 1963, J.C. Wei 0524 (HMAS-L). Hainan, Ledong County, Jiaxi, Mt. Nanlishan, 240m, 22 July 2009, J. Li HN09124 (HMAS-L).
Fissurina globulifica (Nyl.) Staiger
Fig. 1C, D
Biblioth. Lichenol. 85: 137 (2002).
≡ Graphis globulifica Nyl., Bull. Soc. Linn. Normandie, sér. 2 2: 117 (1868). - Graphina globulifica (Nyl.) Zahlbr., Cat. Lich. Univers. 2: 407 (1923) [1924].
Thallus corticolous, crustose, surface smooth to uneven, olive-green. Ascomata lirelliform, elongate, short and unbranched to sparsely branched, erumpent to prominent, 0.5-3mm long and 0.2-0.3mm wide, structure of globulifica-type; disc concealed, with distinct, white labia (hemithecioid), covered by thalline margin; proper exciple uncarbonized; hymenium colorless, clear, 60-100μm high; hypothecium very thick, up to 50μm; paraphyses simple, with apically slightly thickened tips; periphysoids present; asci fusiform, 8-spored; ascospores hyaline, subglobose to oval, muriform, locular irregular, 1-3 transverse and 2-3 longitudinal septa, 8-14×7-9μm, halonate, I+ violet.
Chemistry: Psoromic acid.
Remarks:
Fissurina globulifica is easily distinguished by its
globulifica-type, small subglobose to oval muriform, I+ violet ascospores and the presence of psoromic acid. It is morphologically similar to
F. instabilis (Nyl.) Nyl., but the latter has larger ascospores (5-7/3-4-locular, 17-24×10-15μm) and lacks lichen substances. The species has a pantropical distribution and was previously reported from Malaysia, Philippines, Australian and New Caledonia (
Staiger 2002; Archer
2006,
2009;
Joshi et al. 2013). This is the first report from China.
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 400m, 1 June 2007, J. Li FJ1117-3, FJ1117-4 (LCU).
Fissurina incrustans Fée
Essai Crypt. Exot. (Paris): 46 (1825) [1824].
≡ Graphina incrustans (Fée) Müll. Arg., Mém. Soc. Phys. Hist. nat. Genève 29(8): 47 (1887). - Graphis incrustans (Fée) Nyl., Mém. Soc. Imp. Sci. Nat. Cherbourg 5: 130 (1857).
Thallus corticolous, crustose, surface smooth to warty, beige-green. Ascomata lirelliform, elongate to short, single or branched, erumpent, 1-4mm long and 0.3-0.4mm wide, structure of incrustans-type; disc exposed, with thin labia (platythecioid), covered by thalline margin; proper exciple uncarbonized; hymenium colorless, clear, 80-100μm high; paraphyses simple, with apically slightly thickened tips; periphysoids absent; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, muriform, 3-5 transverse and 1-2 longitudinal septa, locular irregular, 16-20(-25)×8-10μm, halonate, I+ weakly violet.
Chemistry: No lichen substances detected.
Remarks:
Fissurina incrustans is similar to
F. egena (Nyl.) Nyl. in having similar-sized, muriform, amyloid ascospores and lacking lichen substances, but
F. incrustans has erumpent lirellae similar to those found in
Platythecium (
Staiger 2002;
Lücking et al. 2011). The species has been reported from South America (
Staiger 2002) and Vietnam (
Aptroot & Sparrius 2006;
Joshi et al. 2013). In China, it was previously reported from Hong Kong (
Thrower 1988;
Aptroot & Seaward 1999;
Aptroot & Sipman 2001, as
Graphina incrustans.
Seaward & Aptroot 2005; as
F. incrustans).
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 360-400m, 1 June 2007, J. Li FJ920, FJ1076 (LCU); Yongtai County, Mt. Yunding, 890m, 22 August 2016, Z.F. Jia FJ16452 (LCU). Hainan, Changjiang County, Mt. Bawangling, Yajia, 930m, 5 October 2008, J. Li HN08504 (HMAS-L). Hong Kong, New Territories, Tai Po Kau, near Nature Reserve, 150m, 9 July 2000, Aptroot 48765, Sipman 45157 (BGBM).
Fissurina insidiosa C. Knight & Mitt.
Fig. 1E, F
Trans. Linn. Soc. London 23: 102 (1860).
Thallus corticolous, crustose, surface verrucose, rugose, yellowish-green to olive. Ascomata lirelliform, elongate, single, erumpent, 1-3mm long and 0.2-0.4mm wide, structure of subcontexta-type; disc very narrow, labia almost completely covered by uneven to verrucose thalline margin of same color as thallus (hemithecioid); proper exciple uncarbonized; hymenium colorless, clear, 100-150μm high; paraphyses simple, warty at the tips; periphysoids short, indistinctly warty; asci cylindrical, 8-spored; ascospores hyaline, ovoid, 3-septate, 15-20(-25)× 8-12μm, weakly halonate, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina insidiosa is similar to
F. triticea (Nyl.) Staiger, from which it differs in lacking stictic acid and having less emergent ascomata with a less well developed exciple. The material was collected in moist deciduous forest with many waterfalls in southeast China. The species was previously known from Dominica, Florida, New Zealand and India (
Staiger 2002;
Makhija & Adawadkar 2007).
Specimen examined: China. Fujian, Wuyishan City, Mt. Longjingshan, 270m, 22 July 2016, Z.F. Jia FJ16106 (LCU).
Fissurina micromma (Zahlbr.) Aptroot
Symb. Bot. Upsal. 34(1): 34 (2004).
≡ Phaeographina micromma Zahlbr., Feddes Repert. 31: 219 (1933).
Thallus corticolous, crustose, surface smooth to uneven, olive-green. Ascomata lirelliform, elongate, short to elongate, erumpent, single to sparsely branched, 0.5-3mm long and 0.1-0.5mm wide, structure of dumastii-type; disc more or less fissured, opened in growth later period, with thin but distinct labia (slit-like), covered by thalline margin; proper exciple uncarbonized; hymenium colorless, clear, 70-100μm high; paraphyses simple, apically spinulose; periphysoids present; asci fusiform, 2-spored; ascospores hyaline, ellipsoid, muriform, dense locular, (50-)60-80×17-25μm, without halo, I+ violet-blue.
Chemistry: No lichen substances detected.
Remarks:
Fissurina micromma is easily distinguished from other species of
Fissurina by its larger, muriform ascospores, 1-2-spored asci and lack of secondary substances. The species is otherwise morphologically similar to
F. dumastii, but differs by sporomorph. It is a corticolous taxon previously reported from Taiwan as
Phaeographina micromma (Type locality,
Zahlbruckner 1933,
1940;
Lamb 1963;
Wang-Yang and Lai 1973) and later placed in
Fissurina (
Aptroot 2004), subsequently reported by
Jia & Lücking (2017a).
Specimen examined: China. Taiwan, Changhua, Rengechi, 30 July 1925, Y. Asahina F376 (W).
Fissurina nitidescens (Nyl.) Nyl.
Lich. Japon.: 108 (1890).
≡ Graphis nitidescens Nyl., in Tuckerman, Syn. N. Amer. Lich. (Boston) 2: 123 (1888). - Graphina nitidescens (Nyl.) Riddle, Memoirs of the Brooklyn Botanical Garden 1: 115 (1918).
Thallus corticolous, crustose, surface uneven, grey to light yellow, glossy. Ascomata lirelliform, elongate, single, immersed to erumpent, terminally acute, 1-4mm long and 0.2-0.3mm wide, structure of dumastii-type; disc narrow to broad, brown; labia orange-brown, covered with thin thalline margin; proper exciple appearing black and much darker than the thallus, partially carbonized; hymenium colorless, clear, 80-100μm high; paraphyses simple; periphysoids absent or indistinct; asci cylindrical, 8-spored; ascospores hyaline, ellipsoid, muriform, 3-4 transverse and 0-1 longitudinal septa, 15-20×6-10μm, halo 2.5-5μm, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina nitidescens is similar to
F. adscribens but the latter differs in having uncarbonizied lirellae and longer ascospores (20-30µm long). This species is known from Africa, Australia, USA, Brazil and India (
Sharma et al. 2012), and south China (Meng
et al. 2011).
Specimens examined: China. Fujian, Jianou City, Mt. Wanmulin, 420m, 1 June 2007, J. Li FJ1088; Mt. Wanmulin, 420m, 29 March 2004, ER Zhang wy391 (HMAS-L). Guangxi, Longsheng County, Huapiang, 1 000m, 6 June 2001, J.B. Chen, G.L. Hu & L. Xu 20046, 20093 (HMAS-L). Hainan, Qiongzhon County, 630m, Mt. Limushan, 630m, 25 September 2008, J. Li HN013, HN170 (LCU). Hunan, Sangzhi County, Mt. Tianpingshan, 1 500m, 19 August 1997, J.B. Chen, D.P. Wang & SL. Wang 9510 (HMAS-L).
Fissurina radiata Mont.
Annls Sci. Nat., Bot., sér. 2 18: 280 (1842).
≡ Graphis radiata (Mont.) Nyl., Acta Soc. Sci. Fenn. 7(2): 473 (1863).
Thallus corticolous, crustose, surface uneven, grey-green, glossy. Ascomata radiate, in indistinct pseudostromatic clusters, lirellae elongate, structure of dumastii-type; disc concealed; labia inconspicuous and gaping; proper exciple uncarbonized; hymenium colorless, clear, 90-100μm high; paraphyses simple; periphysoids absent or indistinct; asci cylindrical, 8-spored; ascospores hyaline, ellipsoid, with thickened walls and septa, transverse 3-septa, 12-16×6-9μm, halo thick, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina radiata differs from other
Fissurina species in having aggregated, mostly radiate ascomata nearly forming pseudostromata, with inconspicuous and gaping labia, amyloid, 3-septate ocular ascospores of 15×7µm, and lack of lichen substances (
Lücking et al. 2011).
F. radiata is comparable to material described from Florida as
F. aggregatula Common & Lücking (
Lücking et al. 2011), which differs in having conspicuous labia and a concealed disc, sparsely branched aggregated lirellae, and non-amyloid ascospores. The species is known from America, Malaysia, Singapore, and Vietnam (
Aptroot & Sparrius 2006;
Lücking et al. 2011;
Joshi et al. 2013), and was previously reported from Taiwan as
Graphis radiata (
Aptroot & Sparrius 2003).
Specimens examined: China. Hainan, Ledong County, Mt. Jiangengling, Tianchi, 900-920m, 1 October 2008, Z.F. Jia HN080713; HN080729 (HMAS-L).
Fissurina subundulata Kalb & Z.F. Jia
Phytotaxa 189(1): 149 (2014).
Thallus corticolous, crustose, surface uneven, olive green to dark green. Ascomata lirelliform, flexuose, sparsely branched, erumpent to prominent and becoming sessile when old, with apically thin, complete thalline margin, 3-10mm long and 0.5-1mm wide, structure of subcontexta-type; disc concealed to fissured; proper margin distinct, visible as thick, creme-colored lines along the slit; labia entire, with complete thalline margin except along the slit, light olive-green to grey-green; proper exciple not carbonized; hymenium 120-170µm high, colorless, clear; paraphyses unbranched, apically smooth; periphysoids present along the slit, 10-20µm long, with irregular surface but not spinulose; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, muriform, 5-7 transverse and 1-3 longitudinal septa, 20-30×10-12µm, thin-walled, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina subundulata is distinguished by very large, robust, hemithecioid lirellae and muriform, I- ascospores. Similar species are
F. undulata (Müll. Arg.) M. Nakan. & Kashiw. in
Nakanishi et al. (2003) and
F. baishanzuensis (see above), but the former produces stictic acid and the latter has much smaller ascospores and smaller ascomata. The species is also similar to
F. elaiocarpa, but the latter has prominent lirellae, with mostly concealed, orange-brown labia and almost complete thalline margin. This species was collected on the trunk of old deciduous trees in a dark, mixed mountain rainforest on a steep slope of Mt. Baishanzu along a rivulet at ca. 1 600m; it is thus far only known from the type locality of Zhejiang (
Kalb & Jia 2014).
Specimens examined: China. Zhejiang, Qingyuan County, Mt. Baishanzu, Baishanzu Protection Station, 1 300-1 500m, 12 October 2010, K. Kalb, Z.S. Sun & Z.F. Jia 386589 (HMAS-L), 38684 (herb. Kalb); 1 750-1 800m, 12 October 2010, Z.F. Jia 10-136, 10-140 & 10-206 (LCU).
Fissurina tachygrapha (Nyl.) Staiger
Biblioth. Lichenol. 85: 154 (2002).
≡ Graphis tachygrapha Nyl., Acta Soc. Sci. Fenn. 7(2): 473 (1863).
Thallus endoperidermal, crustose, whitish, smooth. Ascomata lirelliform, elongate, single, erumpent, 0.5-2mm long and 0.2-0.25mm wide, structure of comparilis-type; disc gaping (thallolomoid); labia lacking, covered thalline margin; proper exciple apically carbonized; hymenium colorless, clear, 90-100μm high; paraphyses simple, with brownish thickened tips; periphysoids absent or indistinct; asci cylindrical, 8-spored; ascospores hyaline, ellipsoid, with thickened walls, 3-septate, 20-30×10-13μm, halo thick, I-.
Chemistry: No lichen substances detected.
Remarks:
Fissurina tachygrapha is easily distinguished from other
Fissurina species by its whitish, endoperidermal thallus, its apically carbonized exciple, 3-septate ascospores and absence of lichen substances. It is known from South America (
Staiger 2002), and was previously reported from Hong Kong, China (
Thrower 1988, as
Graphis tachygrapha). The material was collected on the trunk near the edge of a light, mixed mountain rainforest from south China.
Specimen examined: China. Fujian, Jianou City, Mt. Wanmulin, 440m, 3 June 2007, J. Li FJ1192 (LCU).
Fissurina triticea (Nyl.) Staiger
Biblioth. Lichenol. 85: 156 (2002).
≡ Graphis triticea Nyl., Acta Soc. Sci. Fenn. 7(2): 470 (1863).
Thallus corticolous, crustose, surface uneven, yellowish, yellow-greenish to olive, smooth to uneven. Ascomata lirelliform, short, sparsely branched, erumpent to prominent, 1-2mm long and 0.2-0.7mm wide, structure of subcontexta-type; disc exposed, sometimes gaping; labia distinct, thick (hemithecioid), yellowish-white, only laterally covered by thalline margin; proper exciple uncarbonized; hymenium 90-120µm high, colorless, clear; paraphyses simple; periphysoids present along the slit, with irregular surface; asci fusiform, 8-spored; ascospores hyaline, ellipsoid, with thickened walls and septa, 3-septate, 15-25×8-12µm, I+ weakly violet-blue.
Chemistry: Stictic acid.
Remarks:
Fissurina triticea resembles
F. consentanea in having
subcontexta-type lirellae, 3-septate ascospores, and the stictic acid chemistry, but the latter is distinguished in having labia completely covered by thallus and I- ascospores. The species is known from Micronesia, Madeira, Reunion Island, Australia, New Zealand, and Costa Rica (
Staiger 2002), and was previously reported from Taiwan, China (
Aptroot & Sparrius 2003, as
Graphis triticea).
Specimen examined: China. Fujian, Jianou City, Mt. Wanmulin, 460m, 2 June 2007, J. Li FJ1177 (LCU).
Excluded name:
Fissurina isidiata Z.F. Jia, Mycotaxon 121: 75 (2012). Type: China. Hainan, Mt. Wuzhishan, 950m, 30.XI.2010, Z.F. Jia 10-612 (Holotype, HMAS-L 117919!; isotype, LCU!).
= Platythecium dimorphodes (Nyl.) Staiger, Biblioth. Lichenol. 85: 383 (2002); - Graphis dimorphodes Nyl., in Leighton WA., Trans. Linn. Soc. London 27: 176 (1869); - Fissurina dimorphodes (Nyl.) Nyl. Acta Soc. Sci. Fenn. 26 (10): 23 (1900). - Graphina dimorphodes (Nyl.) Zahlbr., Cat. Lich. Univ. 2: 404 (1923).
Thallus corticolous, crustose, thick, pale gray to somewhat olive, waxy and slightly warty, with tiny cream-colored warts (isidia-like, dia. 0.1-0.3mm) situated in the edge. Apothecia lirelliform, elongate, single to branched, 0.5-5(-10)mm long, 0.3-1.2mm wide; discs can be seen, jelly-like, brownish, slightly pruinose, flat to somewhat concave; proper exciple convergent, non-carbonized, composed of yellowish to brownish, swollen and loosely twisted hypha, the basal region thick and darkened while the lateral developed radially into the adjacent thalline margin; hymenium clear, 60-90µm tall; mature ascospores 8/ascus, hyaline, ovoid to elongate, submuriform, (3-)4-5/1-2-locular, 15-20×5.5-7.5μm, I+ violet.
Chemistry: norstictic acid.
Remarks:
Fissurina isidiata was described as new species by
Jia et al. (2012). Examination of the type material revealed that it contained norstictic acid, and its morphology and anatomy are in accordance with
Platythecium dimorphodes. Therefore, we propose
F. isidiata as a new synonym of
P. dimorphodes.
Platythecium dimorphodes was previously reported from Guizhou by
Miao et al. (2007).
Specimen examined: China. Hainan, Wuzhishan City, Mt. Wuzhishan, 950m, 30 November 2010, Z.F. Jia 10-612 (HMAS-L). Guizhou, Tonger City, Mt. Fanjingshan, 1 300m, 5 August 2004, J.C. Wei & T. Zhang G420, G426, G567, G191, G444, and G198 (HMAS).
Key to Fissurina species recorded from China
1a. Ascospores muriform 2
1b. Ascospores transversely, 3-septa 11
2a. Ascospores large, (50-)60-80×17-25µm; no lichen substances present F. micromma
2b. Ascospores smaller, usually not exceeding 30 µm in longth 3
3a. Ascospores 8-15×7-10µm 4
3b. Ascospores usually >15µm long 5
4a. Ascospores 8-14×7-9µm, I+ violet; psoromic acid present F. globulifica
4b. Ascospores 10-12(-15)×9-10µm, I-; no lichen substances present F. baishanzuensis
5a. Psoromic acid present; ascospores 20-30×10-15µm, I+ violet F. columbina
5b. Lichen substances absent; ascospores variable 6
6a. Lirellae immersed to erumpent 7
6b. Lirellae prominent to sessile 10
7a. Disc exposed; ascospores 16-20(-25)×8-10µm, I+ weakly violet F. incrustans
7b. Disc concealed or gaping; ascospores I- 8
8a. Lirellae partially carbonized; ascospores 15-20×6-10µm F. nitidescens
8b. Lirellae uncarbonized; ascospores 20-30(-35)µm long 9
9a. Ascospores 8-10µm wide F. adscribens
9b. Ascospores 10-20µm wide F. cingalina
10a. Lirellae prominent, with orange-brown labia; ascospores 20-30×13-20µm, I+ violet-blue
F. elaiocarpa
10b. Lirellae prominent to sessile, with whitish labia; ascospores 20-30×10-12µm, I- F. subundulata
11a. Sticitic acid present 12
11b. Lichen substances absent 13
12a. Labia concealed, mostly covered by nearly complete thalline margin; disc concealed; ascospores 15-25×8-12µm, I-
F. consentanea
12b. Labia exposed, yellowish-white, only laterally covered by thalline margin; disc sometimes gaping; ascospores 15-25×8-12µm, I+ weakly violet-blue F. triticea
13a. Thallus endoperidermal, whitish; excipulum apically carbonized; ascospores 20-30×10-13µm, I-
F. tachygrapha
13b. Thallus epiperidermal, olive-green to yellow-green; excipulum uncarbonized 14
14a. Thallus uneven to verrucose; lirellae hemithecioid; ascospores 15-20(-25)×8-12µm, I-
F. insidiosa
14b. Thallus smooth to uneven; lirellae fissurinoid; ascospores I+ weakly violet-blue 15
15a. Lirellae radiating; ascospores 12-16×6-9µm F. radiata
15b. Lirellae short to elongate, unbranched to sparsely branched; ascospores 10-20×6-10µm F. dumastii
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Fig. 1 A, B: Fissurina consentanea (A: Habit; B: Asci containing ascospores with transverse septa. LCU, Z.F. Jia HN014); C, D: Fissurina globulifica (C: Habit; D: Asci containing ascospores muriform. LCU, J. Li FJ1117-3); E, F: Fissurina insidiosa (E: Habit; F: Asci containing ascospores with transverse septa and I-. LCU, Z. F. Jia FJ16106). Bars: A, C, E=1mm; B, D, F=20μm.
