Spirographa Zahlbr.  (Figs 25–30)

MycoBank number: MB 5151; Index Fungorum number: IF 5151; Facesoffungi number: FoF

Generic type: Spirographa spiralis (Müll. Arg.) Zahlbr.

= Asteroglobulus Brackel, Herzogia 24(1): 69. 2011., syn.nov.

Type: Asteroglobulus giselae Brackel

= Cornutispora Piroz., Mycologia 65(4): 763. 1973., syn.nov.

Type: Cornutispora limaciformis Piroz.

= Graphinella Zahlbr., Cat. Lich. Univers. 2: 285. 1923.

Type: Graphinella fusisporella (Nyl.) Zahlbr.

= Pleoscutula Vouaux, Bull. Soc. Mycol. Fr. 29: 434. 1913.,syn. nov.

Type: Pleoscutula arsenii Vouaux

= Pleospilis Clem., Gen. Fung. (Minneapolis): 69. 1909.

Type: Pleospilis vermiformis (Leight.) Clem.

= Spilomela (Sacc. & D. Sacc.) Keissl., Beih. Bot. Zbl., Abt. 2 37: 272. 1920. ≡ Melaspilea subgen. Spilomela Sacc. & D. Sacc., Syll. Fung. (Abellini) 18: 179. 1906. Type: Spilo- mela vermifera (Leight.) Keissl.

= Spirographomyces Cif. & Tomas., Atti Ist. bot. Univ. Lab. Crittog. Pavia 10(1): 43, 69. 1953.

Type: Spirographomyces spiralis (Müll. Arg.) Cif. & Tomas.

Description of the sexual state. Ascomata apothecioid or perithecioid, cleistohymenial, arising singly or aggregated in stromata, immersed in the host or sessile. When hymenium exposed at maturity then disc concave, dark-brown, orange-brown or black, usually concolorous with the margin or paler. Exciple honey-brown, dark-brown or black, K–, N–, prominent, without hairs, composed of isodiametric cells. Hymenium hyaline, K/I–, I–. Sub-hymenium indistinct. Epihymenium with honey-brown, orange brown or black, granular pigments, K–, N–. Paraphyses 1.5–4 µm thick, hyaline, simple, septate, sometimes slightly branched in the upper part, usually apically thickened and covered by granular pigment. Asci clavate to cylindrical, functionally unitunicate, wall apically not thickened, K/I–, I– (only endoascus I+ slightly orange), ~ 16–32-spored. Ascospores 1-septate, hyaline, narrowly ellipsoidal to fusiform, usually spirally arranged in the ascus, with rounded- or pointed-ends, sometimes with large cilia developed on both apices, not constricted at the septum, straight to curved or sigmoid, multiguttulate, smooth, without perispore, 7–48 × 1–4 µm.

Description of the asexual state. Conidiomata immersed in the host thallus or hymenia, pycnidial, globose to pyriform, hyaline to yellowish brown or dark brown. Pycnidial wall composed of thin- to thick-walled isodiametric cells, disintegrating irregularly around the top to release a colorless to pale-pink mass of agglutinated conidia. Conidiophores hyaline, septate, thin-walled, arising from the innermost cells of the pycnidial walls, branched. Conidiogenous cells holoblastic, synchronous or sympodial, integrated, terminal to lateral, thin-walled, producing 1–3 conidia from minute loci. Conidia hyaline, truncate, aseptate, either Y-shaped, with a main axis and two diverging arms, or triangular, or tetra- to polyhedral, 3–35 µm diam. For a detailed description see Pirozynski (1973), Hawskworth (1976) and Punithalingam (2003).

Notes. Members of the cosmopolitan genus Spirographa (especially its asexual Cornutispora-like states) were previously treated as generalists and reported from a broad variety of host lichens and also from non-lichenized fungi (e.g., Punithalingam 2003; Santesson et al. 2004; Ihlen & Wedin 2008; Brackel 2014). Our phylogenetic analyses showed that species in the genus are strongly host-specific, as the individuals collected from the same host genus clustered together (Fig. 24B). Future examination of a larger material may reveal additional undescribed species and could also help to understand the species boundaries, distribution, and host range in Spirographa.

Our results clearly show that asexual states of Spirographa with very similar characteristics of conidia may be not related phylogenetically (e.g. S. giselae, S. pyramidalis and S. macropyramidalis) and their sexual states strongly differ in ascospore size (11–20 × 2–3.5 µm, 7–10 × 2–3 µm, and 9–13 × 2.5–3 µm, respectively). Keys for the identification of the majority of described Cornutispora-like anamorphs are available (Punithalingam 2003; Etayo 2017; Diederich et al. 2019).

Ecology and distribution. Spirographa is a cosmopolitan genus known from a variety of lichen hosts and non-lichenized fungi. However, the distribution of the majority of the species is poorly known.

Figure 25. Morphological diversity of ascomata in Spirographa. A – S. aggregata on Polyblastidium corallophorum (J.E. 34-5, holotype); B – S. ar- senii on Heterodermia flabellata (J.E. 30-3); C – S. ascaridiella on Porpidia sp. (H-Nyl-21874, lectotype); D – S. fusisporella on Fissurina sp. (M0086813, lectotype); E – S. giselae on Lichenopeltella cf. ramalinae growing on Ramalina farinacea (J.E. 30112); F – S. giselae on Lichenopeltella cf. communis (white arrow showing catathecium of Lichenopeltella) growing on Parmotrema crinitum (A.F. 26962); G – S. hypotrachynae on Hypotrachyna sp. (A.F. 19896); H – S. maroneae (perithecioid ascomata) in Maronea constans (A.F. 27345, holotype); I – S. ophiurospora on Lobariella cf. pallida (A.F. 28890); J – S. pyramidalis on Remototrachyna costaricensis (J.E. 55-1) ; K – S. triangularis on Pertusaria pertusa (J.E. 30859); L – S. usneae on Usnea sp. (A.F. 8176-2, holotype). Scales: A, D = 500 µm; B–C, E–H, J = 250 µm; I = 100 µm; K = 300 µm; L = 200 µm.

Figure 26. Sections of ascomata in selected species of Spirographa (A–C, E–G, I–K in LPCB; D, L in water; H in Congo Red). A–B – S. aggregata on Polyblastidium corallophorum (J.E. 34-5, holotype); C – S. arsenii on Heterodermia flabellata (J.E. 30-3); D–F – S. fusisporella (perithecioid ascomata) in hymenia of Fissurina sp. (M0086813, lectotype); G – S. giselae on Lichenopeltella cf. communis growing on Parmotrema crinitum (A.F. 26962); H–I – S. maroneae (perithecioid ascomata) in thallus of Maronea constans (A.F. 27345, holotype); J – S. ophiurospora on Lobariella cf. pallida (A.F. 28890); K – S. pyramidalis on Remototrachyna costaricensis (J.E. 55-1); L – S. usneae on Usnea sp. (A.F. 8176-2, holotype). Scales: A–B, D–G, K–L = 50 µm; C, H–J = 25 µm.

Figure 27. Morphological variability of ascospores in Spirographa. A – S. aggregata on Polyblastidium corallophorum (J.E. 34-5, holotype, in Congo Red); B – S. arsenii on Heterodermia flabellata (J.E. 30-3, in water); C – S. ascaridiella on Porpidia sp. (H-Nyl-21874, lectotype, in Congo Red); D – S. fusisporella in hymenia of Fissurina sp. [(M0086813, lectotype, in water (left) and Congo Red (right)]; E – S. giselae on Lichenopeltella cf. communis growing on Parmotrema crinitum [A.F. 26962, in water (up) and Congo Red (down)]; F – S. maroneae (perith- ecioid ascomata) in thallus of Maronea constans (A.F. 27345, holotype, in water); G – S. ophiurospora on Lobariella cf. pallida (A.F. 28890, in Congo Red); H – S. pyramidalis on Remototrachyna costaricensis (J.E. 55-1, in water); I – S. usneae on Usnea sp. (A.F. 8176-2, holotype, in water). Scales: A–I = 10 µm.

Figure 28. Morphological diversity of conidiomata in Spirographa. A – S. arsenii on Heterodermia flabellata (J.E. 29-2); B – S. galligena on Erioderma sp. (A.F. Fla39B, holotype); C – S. giselae on Lichenopeltella cf. communis growing on Parmotrema crinitum (A.F.26959); D – S. li- maciformis on Therrya fuckelii (DAOM138360, holotype); E – S. ophiurospora on Lobariella pallida (A.F. 25162); F – S. parmotrematis on Parmotrema sancti-angeli (A.F. 28887, holotype); G – S. pyramidalis on Remototrachyna costaricensis (A.F. 25129); H – S. usneae on Usnea sp. (A.F. 26545); I – S. vermiformis on Lepra amara (A.F. s.n.). Scales: A, C, E, G–H = 100 µm; B, D, F, I = 250 µm.

Figure 29. Sections of conidiomata in selected species of Spirographa (A, C–I in LPCB; B in water). A – S. arsenii in Heterodermia flabellata (J.E. 29-2); B–C – S. galligena in Erioderma sp. (A.F. Fla39B, holotype); D – S. giselae in Lichenopeltella cf. communis growing on Parmotrema crinitum (A.F.26959); E – S. ophiurospora in Lobariella pallida (A.F. 25162); F – S. parmotrematis in Parmotrema sancti-angeli (A.F. 28887, holotype); G – S. pyramidalis in Remototrachyna costaricensis (A.F. 25129); H – S. usneae in Usnea sp. (A.F. 26545); I – S. vermiformis in Lepra amara (A.F. s.n.). Scales: A–I = 25 µm.

Figure 30. Morphological variability of conidia in Spirographa [all in water (left) and LPCB (right)]. A – S. arsenii in Heterodermia flabellata (J.E. 29-2); B – S. galligena in Erioderma sp. (A.F. Fla39B, holotype); C – S. giselae in Lichenopeltella cf. communis growing on Parmotrema crinitum (A.F.26959); D – S. limaciformis on Therrya fuckelii (DAOM138360, holotype); E – S. ophiurospora in Lobariella pallida (A.F. 25162); F – S. parmotrematis in Parmotrema sancti-angeli (A.F. 28887, holotype); G – S. pyramidalis in Remototrachyna costaricensis (A.F. 25129); H – S. usneae in Usnea sp. (A.F. 26545); I – S. vermiformis in Lepra amara (A.F. s.n.). Scales: A–I = 10 µm.

Species