AveSkAMp et al. Group R: This group comprises five species that previously were accommodated <strong>in</strong> <strong>the</strong> sections Phoma, Peyronellaea and Phyllostictoides. As <strong>the</strong> name <strong>of</strong> Ph. tropica already suggests, it concerns a <strong>the</strong>rmotolerant species, which is ma<strong>in</strong>ly found <strong>in</strong> European greenhouses on a wide range <strong>of</strong> hosts, but which probably has a tropical orig<strong>in</strong> (Schneider & Boerema 1975), as do most o<strong>the</strong>r species found <strong>in</strong> <strong>the</strong> present clade. The sole host <strong>of</strong> Ph. costarricensis is c<strong>of</strong>fee bean (C<strong>of</strong>fea arabica), while Ph. piperis is associated with Indian Long Pepper (Piper longus), and <strong>the</strong> novel species Ph. m<strong>in</strong>or has been isolated twice from clove (Syzygium aromaticum) <strong>in</strong> Indonesia. In addition, Ph. labilis is a warmth-preferr<strong>in</strong>g plurivorous species that has been isolated <strong>in</strong> European greenhouses and from nature <strong>in</strong> <strong>the</strong> Middle East, Turkey and Indonesia (Boerema et al. 2004). Phoma zantedeschiae is widespread throughout <strong>the</strong> Western Hemisphere, but always <strong>in</strong> association with arum or calla (Zantedeschia sp.), a genus that is <strong>in</strong>digenous <strong>in</strong> sou<strong>the</strong>rn Africa (Boerema & Hamers 1990). Thus far, however, no data <strong>of</strong> temperature-growth studies are available for <strong>the</strong>se species except for Ph. tropica. Several o<strong>the</strong>r <strong>the</strong>rmotolerant species, such as Ph. calidophila, Ph. calorpreferens and Ph. multirostrata, are, however, not accommodated <strong>in</strong> this group. These three species are soil-borne, <strong>in</strong> contrast to Ph. tropica and Ph. costarricensis, which are associated with leaf-spots. Phoma tropica and Ph. costarricensis are both closely related, and colony characters are highly similar. However, <strong>the</strong> stra<strong>in</strong>s available revealed a significant difference <strong>in</strong> conidial and pycnidial sizes, consistent with <strong>the</strong> data obta<strong>in</strong>ed <strong>in</strong> previous studies (Schneider & Boerema 1975, De Gruyter & Noordeloos 1992). Phoma costarricencis Echandi, Rev. Biol. Trop. 5: 83. 1957. Specimens exam<strong>in</strong>ed: Nicaragua, from a twig <strong>of</strong> C<strong>of</strong>fea sp., 1991, CBS 506.91 = PD 91/876 = IMI 215229. Unknown orig<strong>in</strong>, from C<strong>of</strong>fea arabica, 1979, CBS 497.91 = PD 79/209. Notes: Stra<strong>in</strong> CBS 497.91 was <strong>in</strong>itially identified as Ph. tropica. The close phylogenetic association between this species and Ph. costarricensis concurs with <strong>the</strong>ir overlapp<strong>in</strong>g morphological characters (see Schneider & Boerema 1975, De Gruyter & Noordeloos 1992). Phoma labilis Sacc., Michelia 2(7): 341. 1881. Specimens exam<strong>in</strong>ed: Israel, from a stem <strong>of</strong> Rosa sp., 1970, G.H. Boerema, CBS 479.93 = PD 70/93. The Ne<strong>the</strong>rlands, Barendrecht, from a stem <strong>of</strong> Lycopersicon esculentum, 1987, J. de Gruyter, CBS 124.93 = PD 87/269. Phoma m<strong>in</strong>or Aveskamp, Gruyter & Verkley, sp. nov. MycoBank MB515641. Fig. 11. Conidia ellipsoidea, ovoidea vel leniter allantoidea, glabra, hyal<strong>in</strong>a, cont<strong>in</strong>ua, (3–)3.5–4.5(–5) × 1.8–2.5(–3) μm, (0–)1–3(–4) guttulis m<strong>in</strong>utis praedita. Matrix conidiorum alba. Etymology: Epi<strong>the</strong>t derived from <strong>the</strong> small-sized conidia. Conidiomata pycnidial, solitary, (sub-)globose to broadly ellipsoidal, glabrous or with some hyphal outgrows, on <strong>the</strong> agar surface and immersed, (125–)150–280(–330) × (125–)150–220(–245) μm. Ostioles (1–5), slightly papillate or non-papillate. Pycnidial wall pseudoparenchymatous, composed <strong>of</strong> oblong to isodiametric cells, outer cell layer pigmented, 2–4 layers, 8–15 μm thick. Conidiogenous cells phialidic, hyal<strong>in</strong>e, simple, smooth, flaskshaped or somewhat isodiametric, ca. 4–5.5(–6.2) × 3–4.5(–4.7) 42 μm. Conidia ellipsoidal to ovoid or slightly allantoid, th<strong>in</strong>-walled, smooth, hyal<strong>in</strong>e, aseptate (3–)3.5–4.5(–5) × 1.8–2.5(–3) μm, with (0–)1–3(–4) m<strong>in</strong>ute guttules. Conidial matrix white. Culture characteristics: Colonies on OA (44–)45–50(–54) mm diam after 7 d, marg<strong>in</strong> regular. Aerial mycelium flat, grey, but locally well-developed <strong>in</strong> densely floccose white tufts. Immersed mycelium olivaceous with rosy-buff t<strong>in</strong>ges near <strong>the</strong> colony marg<strong>in</strong>; reverse concolourous. Colonies on MEA 46-48 mm diam after 7 d, marg<strong>in</strong> regular. Immersed mycelium hyal<strong>in</strong>e, with abundant semiimmersed pycnidia, but almost completely covered by an aerial mycelial mat. Aerial mycelium pluriform, with a compact white mat and some felty glaucous grey or dull green zones, near colony marg<strong>in</strong> white; reverse black to grey-olivaceous. Colonies on CHA 50–54 mm diam. after 7 d, marg<strong>in</strong> regular. Aerial mycelium similar as on MEA, although <strong>the</strong> felty white and glaucous grey zones are less abundant; reverse slate blue to leaden-black. Application <strong>of</strong> NaOH results <strong>in</strong> a greenish yellow discolouration <strong>of</strong> <strong>the</strong> agar, best to be observed on OA medium. Specimens exam<strong>in</strong>ed: Indonesia, Sumatra, from Syzygium aromaticum, Apr. 1982, R. Kasim, holotype designated here CBS H-20236, ex-holotype culture CBS 325.82; Lampung, from Syzygium aromaticum, Dec. 1982, H. Vermeulen, CBS 315.83. Notes: As for Ph. eucalyptica, this species has been recorded <strong>in</strong> association with clove trees (Syzygium aromaticum, Boerema et al. 2004). Both species, although genetically dist<strong>in</strong>ct, have many characters <strong>in</strong> common, notably <strong>the</strong> colony characters on OA, <strong>the</strong> high variation <strong>in</strong> ostiole number and a similar reaction to application <strong>of</strong> NaOH. Although Phoma m<strong>in</strong>or produces relatively small conidia, <strong>the</strong> conidia <strong>of</strong> Ph. eucalyptica are even smaller, measur<strong>in</strong>g only 2–4 × 1–2 μm (De Gruyter & Noordeloos 1992). Phoma piperis (Tassi) Aa & Boerema, Persoonia 15(3): 398. 1993. Basionym: Phyllosticta piperis Tassi, Boll. Reale Orto Bot. Siena 3(2): 28. 1900. Specimens exam<strong>in</strong>ed: The Ne<strong>the</strong>rlands, Tiel, from a leaf <strong>of</strong> Peperomia pereskiifolia, 1988, J. de Gruyter, CBS 268.93 = CBS 108.93 = PD 88/720; Tiel, from Peperomia sp., 1990, J. de Gruyter, PD 90/2011 Phoma tropica R. Schneid. & Boerema, Phytopathol. Z. 83 (4): 361. 1975. Specimen exam<strong>in</strong>ed: Germany, Horrheim, from Sa<strong>in</strong>tpaulia ionantha, 1973, R. Schneider, isotype CBS H-7629, ex-isotype culture CBS 436.75. Phoma zantedeschiae Dippen., S. African J. Sci. 28: 284. 1931. Specimen exam<strong>in</strong>ed: The Ne<strong>the</strong>rlands, from a bulb <strong>of</strong> Zantedeschiae sp., 1969, G.H. Boerema, CBS 131.93 = PD 69/140. Group S – Stagonosporopsis: This large group (BPP = 1.00, RBS = 55 %) comprises ma<strong>in</strong>ly species with Stagonosporopsis synanamorphs. In <strong>the</strong> Boeremaean classification system, <strong>the</strong>se species were embedded <strong>in</strong> Phoma section Heterospora (Boerema et al. 1997). As with <strong>the</strong> o<strong>the</strong>r sections, this group also appeared to be artificial. Based on LSU and SSU sequences, <strong>the</strong> type species <strong>of</strong> <strong>the</strong> section Heterospora, Ph. heteromorphospora, clusters outside <strong>the</strong> <strong>Didymellaceae</strong> (De Gruyter et al. 2009), as do Ph. samarorum and Ph. dimorphospora.
www.studies<strong>in</strong>mycology.org Phoma And relAted pleoSporAleAn generA Fig. 11. Phoma m<strong>in</strong>or (CBS 325.82). A–C. Fourteen-day-old colonies on OA (A), MEA (B) and CHA (C). D–E. Pycnidia. F. Section <strong>of</strong> <strong>the</strong> pycnidial wall. G. Conidia. Scale bars: D = 200 μm; E = 100 μm; F–G = 10 μm. Three species, Ph. clematid<strong>in</strong>a, Ph. glaucii and Ph. aquilegiicola form a separate clade (Group C) with<strong>in</strong> <strong>the</strong> <strong>Didymellaceae</strong>, and are treated above. Also Ph. nigripycnidia and Ph. narcissi are not accommodated here. In contrast to <strong>the</strong> Heterospora species that are absent <strong>in</strong> this clade, several current Phoma taxa recovered here have been associated with <strong>the</strong> section Phyllostictoides, such as Ph. artemisiicola, Ph. caricae-papayae, Ph. cucurbitacearum, Ph. heliopsidis, Ph. rudbeckiae, and <strong>the</strong> quarant<strong>in</strong>e-organisms Ph. ligulicola var. ligulicola and var. <strong>in</strong>oxydabilis (De Gruyter 2002). These are all <strong>in</strong>cluded <strong>in</strong> subclade S1 (BPP = 0.93, RBS = 73 %). These species do produce a percentage <strong>of</strong> multicellular conidia <strong>in</strong> culture that are <strong>of</strong>ten considerably larger than <strong>the</strong> regular aseptate ones. However, Boerema et al. (1997) decided to exclude <strong>the</strong> Ph. ligulicola varieties and Ph. cucurbitacearum from section Heterospora, as <strong>the</strong> sizes <strong>of</strong> <strong>the</strong> Stagonosporopsis-like conidia do not always exceed that <strong>of</strong> <strong>the</strong> aseptate conidia <strong>in</strong> <strong>the</strong>se species. A sister clade to subclade S1 is S2, which hosts <strong>the</strong> potato pathogens Ph. andigena and Ph. crystall<strong>in</strong>iformis – formerly known as Ph. and<strong>in</strong>a var. crystall<strong>in</strong>iformis. Both species orig<strong>in</strong>ate from <strong>the</strong> Andes region, and are regarded as serious quarant<strong>in</strong>e pathogens <strong>in</strong> large parts <strong>of</strong> <strong>the</strong> world (Smith et al. 1992). In addition, three o<strong>the</strong>r subclades can be recognised <strong>in</strong> this clade. One (S3) comprises <strong>the</strong> species Ph. schneiderae and Ph. subboltshauserii (both <strong>of</strong> <strong>the</strong> section Heterospora) as well as Ph. astragali. This species is known as a pathogen <strong>of</strong> Astragalus spp., and is characterised by a high percentage <strong>of</strong> “distorted” conidia, but thus far, no records have been made <strong>of</strong> a Stagonosporopsislike synanamorph. Whereas records <strong>of</strong> Ph. astragali and Ph. schneiderae are ma<strong>in</strong>ly limited to <strong>the</strong> American cont<strong>in</strong>ent, Ph. 43