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An update of Podospora sect. Podospora, with first descriptions from Italy of P. appendiculata and P. perplexens Francesco DOVERI Summary: A survey of coprophilous fungi from Italy has allowed the author to recognise nineteen Podospora species, three (P. fimiseda, P. appendiculata and P. perplexens) belonging to section Podospora, whose main morphological features are mentioned and also examined in the light of recent molecular studies. P. appen- diculata and P. perplexens are described for the first time from Italy and compared with P. fimiseda, which has frequently been collected and described by the author. The three species are also compared with others in Ascomycete.org, 7 (2) : 45-53. sect. Podospora, and a key to the section is provided. Species currently excluded from sect. Podospora are Avril 2015 also discussed. Mise en ligne le 2/04/2015 Keywords: Cercophora, dung, Lasiosphaeriaceae, phylogeny, taxonomy. Introduction Abbreviations: CLSM = author’s personal herbarium; MCVE = Her- barium of Venetian Civic Museum. My ongoing survey of coprophilous fungi from Italy, started in 1995, has discovered one new Podospora sp. (DOVERI, 2004a) and des- Taxonomy cribed eighteen Podospora spp. new to Italy (DOVERI 2004b, 2008; DOVERI et al., 1998, 2000). Recently I have been lucky enough to iso- Accepted species in section Podospora late, from herbivore dung, Podospora appendiculata and P. per- plexens, two species which exhibit morphological features very close Podospora appendiculata (Auersw.) Niessl, Hedwigia, 22: 156 to P. fimiseda, which I have already described from Italy. The latest (1883). Figs 1–7 finds enable me to compare all three taxa with each other and with Basionym: Sordaria appendiculata Auersw., in Niessl, Ver. Nat. Ver. other species of Podospora Ces. in sect. Podospora s. LUNDQVIST (1972), Brünn., 10 (2): 188 (1872). which previously have not been found in Italy. My study has invol- ≡ Podospora fimiseda var. appendiculata (Auersw.) G. Winter, Ra- ved a review of the world literature on sect. Podospora and resulted benh. Krypt. Fl., 1 (2): 170 (1885). in an increased knowledge of it. The aim of the present work is to ≡ Pleurage appendiculata (Auersw.) Kuntze, Rev. Gen. Plant.: 505 provide an update on this section. (1898). Section Podospora is characterised by a coriaceous, multi-layered, ≡ Sordaria fimiseda var. appendiculata (Auersw.) Massee & E.S. Sal- gelatinous (pseudo-bombardioid) peridium, perithecia fully covered mon, Ann. Bot., 15: 340 (1901). with hairs hyaline at their tips, clavate immature ascospores, and = Sordaria coprophila f. hirtula P. Karst., Fungi Fenn. Exs. no. 284 asci with a thin apical apparatus. The close relationship between this (1865), nom. nud. section and the genus Cercophora Fuckel in Lasiosphaeriaceae = Sordaria winteri P. Karst., Mycol. Fenn., 2: 251 (1873). Nannf. was emphasized by LUNDQVIST (1972) after finding that P. ap- = Podospora winteri (P. Karst.) Niessl, Hedwigia, 22: 156 (1883). pendiculata and P. fimiseda have a layered, gelatinous peridium, like = Sphaeria amphicornis Ellis, Bull. Torrey Bot. Club, 6: 109 (1876); some coprophilous Cercophora spp. [C. albicollis N. Lundq., C. ele- Sordaria amphicornis (Ellis) Sacc., Syll. Fung., 1: 235 (1882); Podospora phantina (Henn.) N. Lundq., C. scortea (Cain) N. Lundq.], asci with an amphicornis (Ellis) Ellis & Everh., N. Amer. Pyren.: 130 (1892); Pleurage occasional subapical globulus and ascospores able to germinate amphicornis (Ellis) Kuntze, Rev. Gen. Plant.: 505 (1898). when still at the hyaline stage, as seen throughout Cercophora as a = Sphaeria eximia Peck, Ann. Rep. N.Y. State Mus., 28: 78 (1876). whole. = Pleurage superior Griffiths, Mem. Torrey Bot. Club, 11: 68 (1901); Phylogenetic or comparative morphological and phylogenetic Sordaria superior (Griffiths) Sacc. & D. Sacc., Syll. Fung., 17: 603 (1905). studies (MILLER & HUHNDORF, 2005; CAI et al., 2006; CHANG et al., 2010, (all synonyms fide LUNDQVIST 1972) KRUYS et al., 2015) confirmed UNDQVISTL ’s (1972) observations, which were based only on morphological affinities. They showed that Perithecia 600–750 × 450–500 μm, ovoidal, dark brown, coria- many genera in the Lasiosphaeriaceae Nannf. were polyphyletic, in- ceous, fully covered with thick, rigid, non-agglutinated hairs, with a cluding Podospora Ces., and Podospora species with a pseudo-bom- scarcely differentiated (subconical), darker neck, about 100 × 100 bardioid peridium occurred in well supported clades, where they μm, glabrous only at its apex. Peridium pseudobombardioid, 4- nest with Cercophora spp. possessing the same kind of peridium. layered: first layer from inside pseudoparenchymatous, a textura an- Further comparative, morphological and phylogenetic studies are gularis of pale, thin-walled, polygonal cells, 12–15 × 10–12 μm; desirable, including the sequencing of the greatest number of Cer- second layer of pale, parallel, cylindric hyphae; third layer a gelati- cophora and Podospora species with a pseudo-bombardioid peri- nous, shapeless material; fourth layer of dark brown, thick-walled, dium, likely to be included in a distinct genus of Lasiosphaeriaceae. irregular cells, 6–12 × 4–8 μm, usually with wavy walls (textura an- gularis to epidermoidea). Hairs numerous, dark brown, hyaline and Materials and methods roundish at the tips, 1–3-septate, straight to curved, thick-walled, 50–80 × 3–5 μm, with a branched or simply curved base. Para- Some collections came from fresh dung in the field, while most physes not seen. Asci not seen. Ascospores one-celled, hyaline, el- were obtained from different types of dung cultured in non-axenic lipsoid-fusiform in the early stages, two-celled later, as subdivided damp chambers. Collections were studied according to RICHARDSON by a transverse septum in a dark upper cell (spore head) and a hya- & WATLING (1997) and RICHARDSON (2001a), slightly modified as descri- line lower cell (pedicel). Spore head (22–) 23–25 (–26) × (11.5–) 12– bed by DOVERI (2004b). 13 μm, ellipsoidal or narrowly ellipsoid (Q= 1.76–2.16; average Q = 45 1 2 34 6 5 7 Plate 1 – Podospora appendiculata 46 89 11 10 12 Plate 2 – Podospora fimiseda 47 Legend of Plate 1 – Podospora appendiculata 1. Apical part of perithecium. 2. Detail of exoperidium. 3-5 Peridial hairs. 6. Base of peridial hairs. 7. Ascospores with hollow (arrows) caudae. Scale bars = 1) 20 μm; 2) 10 μm; 3-6) 20 μm; 7) 20 μm. Legend of Plate 2 – Podospora fimiseda 8. Exoperidium with rigid hairs. 9. Hairy perithecial neck. 10. Detail of exoperidium. 11. Biseriate ascospores. 12. Ascospores. Scale bars = 8) 100 μm; 9) 20 μm; 10) 40 μm; 11) 40 μm; 12) 50 μm. 1.96), equilateral, dark brown, smooth, thick-walled, flattened at the 1100 m, on cattle dung, A. Bizzi, 14.9.2005, CLSM 01995-XV; Trento, base, with a slightly umbonate apex and an apical germ pore. Pedi- Dimaro-Malghetta, 1100 m, on hare dung in culture, F. Doveri, cel obconical, 8–12 × 5–6 μm, soon collapsing. Upper gelatinous 8.9.2007, CLSM 01995-XVI; Catania; Randazzo, Foresta demaniale Ca- cauda lash-shaped, central, hiding the germ pore, 30–60 × 5–6 μm, rone-Bosco del Flascio, 1100 m, on wild pig dung in culture, L. Le- hollow. Lower cauda very similar to the upper cauda, arising from vorato, 29.4.2013, CLSM 01995-XVII. the pedicel apex, 45–60 × 4–5 μm. Material examined: ITALY, Belluno, Santo Stefano-Alto Comelico, Podospora lindquistii García-Zorrón, Bol. Soc. Arg. Bot., 18 (1-2): 1600 m a.s.l., about ten scattered, semi-immersed to superficial, 173 (1977). overmature specimens on hare (Lepus sp.) dung, L. Levorato, 16.8.2010, 030.1-Comelico Superiore, CLSM 002.11. Podospora minipistillata R.S. Khan & J.C. Krug, in Krug & Khan, Can. J. Bot., 67: 1179 (1989). Podospora fimiseda (Ces. & De Not.) Niessl, Hedwigia, 22: 156 (1883). Figs 8–12 Basionym: Sordaria fimiseda Ces. & De Not., Comm. Soc. Critt. Ital., Podospora minor Ellis & Everh., Am. Nat., 31: 341 (1897). 1: 226 (1863). ≡ Sordaria minor (Ellis & Everh.) Sacc. & P. Syd., Syll. Fung., 14: 493 ≡ Cercophora fimiseda (Ces. & De Not.) Fuckel, Jahrb. Nass. Ver. Na- (1899). turk., 23–24: 244 (1870). ≡ Pleurage minor (Ellis & Everh.) Griffiths, Mem. Torrey Bot. Club, 11: ≡ Sphaeria fimiseda (Ces. & De Not.) Cooke & Berk., Int. Sci. Ser., 14: 67 (1901). 134 (1875). ≡ Pleurage fimiseda (Ces. & De Not.) Griffiths, Mem. Torrey Bot. Club, 11: 69 (1901). Podospora perplexens (Cain) Cain, Can. J. Bot., 40: 460 (1962). Figs ≡ Bombardia fimiseda (Ces. & De Not.) Kirschst., Krypt. Fl. Branden- 13–21 burg, 7 (2): 185 (1911). Basionym: Sordaria perplexens Cain, Univ. Toronto Stud.: 50 (1934). = Sphaeria equina Fr., in Montagne, Ann. Sci. Nat., Bot., 2a Sér., 1: ≡ Pleurage perplexens (Cain) C. Moreau, Encyclop. Mycol., 25: 264 337 (1834), nom. illeg.; Hypocopra equina (Fr.) Sacc., Syll. Fung., 1: 247 (1954). (1882); Sordaria equina (Fr.) Cooke, Grevillea, 16: 78 (1887); Pleurage equina (Fr.) Kuntze, Rev. Gen. Plant.: 505 (1898). Perithecium 950 × 400 μm, obpyriform, dark brown, coriaceous, = Podospora fimicola (Corda) Ces., s. Cesati, in Rabenh., Bot. Zeit., fully covered with dense, rigid, hairs, with a scarcely differentiated, 14: 429 (1856). subconical neck, 200 × 180 μm, covered with the same kind of hairs. (all synonyms fide LUNDQVIST 1972) Peridium pseudobombardioid, 4-layered: first layer from inside a textura angularis of pale, thin-walled, polygonal cells, 10–17 × 6– N.B.: for description of Italian collections, see DOVERI et al. (1998) 12 μm; second layer formed of pale, parallel, cylindric hyphae; third and DOVERI (2004b).