Strigula Fr., Syst. Mycol. 2(2): 535 (1823) [nom. sanct.];

Fries, Kongl. Vet.-Akad. Handl. 1821: 323 (1821) [nom. nud.].

MycoBank number: MB 5285; Index Fungorum number: IF 5285; Facesoffungi number: FoF 07706; 30 morphologically delimited species (Jiang et al. 2020b); molecular data available for approximately 15 species (Jiang et al. 2016, 2017a, b, 2020b, Ford et al. 2019; Oh et al. 2019; Woo et al. 2020).

= Nemathora Fée, Essai Crypt Écorc. 1: LVIII, XCIV, XCIX (1825); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).

Type species: Nemathora viridissima Fée [= Strigula smaragdula Fr.].

= Craspedon Fée, Essai Crypt Écorc. 1: LIX, XCIV, C (1825); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).

 Type species: Craspedon concretum Fée [≡ Strigula con- creta (Fée) R. Sant.].

= Melanophthalmum Fée, Essai Crypt. Écorc. 1: LX, XCIV, C (1825); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).

Type species: Melanopthalmum antillarum Fée [≡ Strigula antillarum (Fée) Müll. Arg.].

= Haploblastia Trevis., Conspectus Verrucarinarum: 16 (1860); Santesson, Symb. Bot. Upsal. 12(1): 138 (1952).

Type species: Haploblastia nitidula (Mont.) Trevis. [≡ Strig- ula nitidula Mont.].

= Discosiella Syd. in Sydow & Sydow, Leafl. Philipp. Bot. 5: 1546 (1912); Eriksson & Hawksworth, Syst. Asco-

mycetum 11: 56 (1992).

 Type species: Discosiella cylindros- pora Syd. [= Strigula sp.].

= Strigulomyces [as Strigulomyces] Cif. & Tomas., Atti Ist. Bot. Univ. Lab. Crittogam. Pavia, Ser. 5, 10: 61 (1953) [nom. illeg.]; Harris, More Florida Lichens: 152 (1995); Lücking & Hawksworth, Taxon 56: 1274 (2007).

Type species: Strigula elegans (Fée) Müll. Arg. [= Strigula smarag- dula Fr.].

= Catenata Bat. in Batista & Bezerra, Publ. Inst. Micol. Univ. Recife 321: 15 (1961) [nom. inval.]; Lücking et al., Lichenologist 30: 161. (1998).

Original species: Catenata antillarum (Fée) Bat. [≡ Strigula antillarum (Fée) Müll. Arg.].

= Kilikiostroma Bat. & J. L. Bezerra, Publ. Inst. Micol. Univ. Recife 321: 13 (1961); Lücking et al., Lichenologist 30: 144 (1998).

Type species: Kilikiostroma peresii Bat. & J. L. Bezerra [= Strigula prasina Müll. Arg.].

= Pycnociliospora Bat. in Batista et al., Publ. Inst. Micol. Univ. Recife 251: 6 (1962); Lücking et al., Lichenologist 30: 154 (1998).

Type species: Pycnociliospora belluciae Bat. & J. A. Lima [= Strigula antillarum (Fée) Müll. Arg.].

= Porina sect. Sagediastrella Vain., Ann. Acad. Sci. Fenn., Ser. A, 19: 10 (1923); Santesson, Symb. Bot. Upsal. 12(1): 138. 1952.

Type species: Porina nitens Vain. [= Strigula nitidula Mont.].

= Shanoria Anahosur in Harris, More Florida Lichens: 144 (1995) [nom. inval., non Shanoria Subram. & K. Ramakr., non Shanoria sensu Anahosur (1967)].

Original species: Shanoria indica Anahosur [= Strigula cf. nitidula Mont.].

Lichenized on leaves in terrestrial, lowland to mon- tane, tropical to subtropical, and extending into temperate, habitats. Thallus (pseudo-)corticate, grey-green to bright green, subcuticular and sometimes damaging the supporting leaf. Photobiont Cephaleuros. Sexual morph: Ascomata perithecia, erumpent to prominent, black or covered by thallus layer, lens-shaped to wart-shaped or conical, carbonaceous, ostiolate. Involucrellum present, carbonized. Excipulum prosoplectenchymatous, dark brown to brown- black. Hamathecium comprising 0.5–0.7 µm wide paraphyses, hyaline, flexuose, unbranched or very rarely branched and anastomosing. Asci usually 8-spored but sometimes appearing 9–16-spored due to ascospores breaking in halves within the asci, bitunicate, fissitunicate, cylindrical to narrowly obclavate, shortly pedicellate, with narrow ocular chamber, non-amyloid. Ascospores irregularly arranged to uni- or biseriate, fusiform to ellipsoid or bacillar, hyaline, 1-septate, rarely 3-septate, with thin eusepta and rectangular lumina, smooth-walled, often constricted (and sometimes breaking apart) at the septa. Asexual morph: Pycnidia common, immersed to erumpent, visible as black dots, rarely in pseudostromatic clusters, often whole thalli only producing pycnidia. Conidia acrogenous, either macro- or microconidia; macroconidia mostly 1-septate, ellipsoid to bacillar, with gelatinous appendages, hyaline; microconidia usually aseptate, ellipsoid to fusiform, small, hyaline.

Chemistry: No secondary substances known.

Type species: Strigula smaragdula Fr., Linnaea 5: 550 (1830).

Notes: Strigula as here defined includes the bulk of foliicolous species centered around S. smaragdula, after separation of the genera Dichoporis, Phyllocharis, Phylloporis, Puiggariella, Raciborskiella, and Racoplaca, and introduction of the new genera Serusiauxiella and Swinscowia (see below). As shown by Jiang et al. (2020b), a much more narrowly defined Strigula is still heterogeneous: whereas the species with a morphology similar to S. smaragdula form a supported clade, three basal lineages differ in morphology and/or anatomy, including the S. nitidula and S. prasina groups and S. macaronesica. While Strigula sensu stricto forms thickened, usually bright green thalli, those of the S. nitidula group are very thin and more similar to those of Racoplaca, and S. prasina and relatives have hypophyllous thalli with bluish tinge and branched and anastomosing paraphyses. Should these be confirmed as separate clades in a broader sampling of taxa, the names Haploblastia (S. nitidula group) and Kilikiostroma (S. prasina group) would be available. Currently, about 30 exclusively foliicolous species are to be assigned to the genus Strigula sensu stricto as here defined.

Harris (1995) listed the name Shanoria Anahosur as a synonym of Strigula, given that the only species described by that author, S. indica Anahosur, represents the pycnidial stage of Strigula cf. nitidula. While the latter is correct (see also Raj 1981), Harris (1995) misinterpreted its protologue, since Anahosur (1967) did not establish a new genus but placed his new species in the already existing genus, Shanoria Subram. & K. Ramakr., typified by S. bambusarum Subram. & K. Ramakr. (Subramanian and Ramakrishnan 1956). The latter has been established as a synonym of Placonema (Sacc. & D. Sacc.) Petr. (asexual morph; Sutton 1977) and Kiehlia Viégas (sexual morph; Farr 1968), with Placonema having priority. The name Shanoria Anahosur does therefore not exist with that place and date of publication (Anahosur 1967), but was inadvertently, yet effectively created by Harris (1995), who ascribed the name to Anahosur. Unfortunately, this is a rather common lapsus, frequently seen in early mycological literature, which has perpetuated many presumed homonyms that were in fact never established as such by their ascribed authors. The Code (Shenzen Code; Turland et al. 2018) does not include a specific provision for names created through erroneous citation, and so various rules must be consulted to ascertain whether such a name is actually valid (and consequently illegitimate). In the case of Shanoria Anahosur in Harris (1995), Harris (1995) provided a full and direct reference to the Latin description of S. indica and cited the latter as type of the genus, so the provisions of ICN Art. 38.1, 38.13, 39.1 and 40.1 are theoretically fulfilled. Since Harris (1995)

did not use an explicit rank designation, such as ‘gen. nov.’, because he did not intend to describe a new genus, one could cite ICN Art. 37.1 to render the name invalid; however, that article does not include a specific provision how it should be applied to genus names, and the genus rank of the name is clear from context. Further, ICN Art. 36.1(b) may suggest the name as not validly published, since it is merely cited in synonymy. However, that article specifically refers to the intention of the author of the name, not the author of the publication. As Harris (1995) ascribed the name to Anahosur and not to himself [ICN Art. 46.2], his placement of Shanoria as synonym of Strigula appears irrelevant with respect to ICN Art. 36.1(b). ICN Art. 38.5 provides another potential avenue, as for a simultaneously established new genus and species, validation through reference to an earlier description is not possible. That article does not apply, though, as Harris (1995) only established a new genus name, whereas the name Shanoria indica Anahosur had already been validly described earlier (Anahosur 1967). Fortunately, this case finds its solution in ICN Art. 38.11(b), which specifies that “…for a name of a genus or subdivision of a genus, the earlier description or diagnosis must be that of a genus or subdivision of a genus”. Hence, the name Shanoria Anahosur in Harris (1995) is not validated by reference to the Latin description of Shanoria indica Anahosur (1967), and it its therefore invalid. It appears ineffective that one has to fully explore the Code to eventuelly render a name invalid that was inadvertently created through erroneous citation. ICN Art. 48.1 may seemingly reflect such cases, but in reality it does not apply, as it specifically refers to names intentionally adopted by an author but in a sense different from the protologue, i.e. deliberately excluding the type of the name. Harris (1995) did not intentionally adopt the name Shanoria in a different sense, he simply cited it incorrectly, due to a misinterpretation of Anahosur’s (1967) paper. He also did not explicitly exclude its original type but did so by implication, through citing another species erroneously as type of a name that did not actually exist, and through indicating Shanoria Anahosur as illegitimate, which means Harris (1995) was aware of the existence of Shanoria Subram. & K. Ramakr. but did not realize that Anahosur (1967) was using precisely that name. Molecular data show that the type species, Strigula smaragdula, in its current definition is a collective species (Jiang et al. 2016, 2017a, b; Oh et al. 2019; Woo et al. 2020). Given that the type is from Nepal, the name is possibly to be applied to a taxon restricted to eastern Asia, whereas the bulk of tropical species will have to be redispositioned using in part available names currently in synonymy of S. smaragdula. This may include reinstatement of the epithet elegans for a common and possibly widespread tropical taxon, with the type being from the Caribbean region.


  • Strigula smaragdula