Crous et al. system, consisting of a single terminal conidiogenous cell giving rise to several ramoconidia that form secondary ramoconidia, etc., or the branched apparatus is composed of several terminal <strong>and</strong> sometimes lateral conidiogenous cells giving rise to sequences of ramoconidia (conidiogenous cells <strong>and</strong> ramoconidia are often barely distinguishable, with conidiogenous cells disarticulating, becoming ramoconidia). <strong>The</strong> branched apparatus may be loose to dense, metula-like. <strong>The</strong> conidiogenous cells have only few, usually 1–3 (–4), terminal or subterminal subdenticulate loci, <strong>and</strong> ramoconidia are prominent <strong>and</strong> numerous, giving rise to branched chains of secondary conidia with flat-tipped hila. Some species of Penidiella with compact, metula-like branched apices are morphologically close to Metulocladosporiella Crous, Schroers, J.Z. Groenew., U. Braun & K. Schub. (Crous et al. 2006d). This <strong>genus</strong> encompasses two species of banana diseases belonging to Herpotrichiellaceae (Chaetothyriales), characterised by having conidiophore bases with rhizoid hyphal appendages <strong>and</strong> abundant micronematous conidiophores. Penidiella species with less pronounced penicillate apices, e.g. P. strumelloidea (Milko & Dunaev) Crous & U. Braun, are comparable with species of the <strong>genus</strong> Pleurotheciopsis B. Sutton (see Ellis 1976). <strong>The</strong> latter <strong>genus</strong> is distinct in having unbranched, often percurrently proliferating conidiophores, lacking ramoconidia <strong>and</strong> colourless conidia formed in simple chains. <strong>Cladosporium</strong> helicosporum R.F. Castañeda & W.B. Kendr. (Castañeda et al. 1997) is another penidiella-like fungus with terminally branched conidiophores, subdenticulate conidiogenous loci <strong>and</strong> conidia in long acropetal chains, but its affinity to Penidiella has still to be proven. Key to Penidiella species 1. Conidiophores in vivo in well-developed, dense fascicles <strong>and</strong> distinct synnemata arising from a basal stroma; on fallen leaves of Ficus sp., Cuba ............................................................................................................................................................................... P. cubensis 1. Conidiophores solitary, at most loosely aggregated .................................................................................................................................. 2 2. Conidiophores with a terminal conidiogenous cell, often somewhat swollen, giving rise to several ramoconidia (on one level) that form chains of straight to distinctly curved conidia; isolated from leaf of Carex sp., Russia ..................................................... P. strumelloidea 2. Penicillate apex of the conidiophores composed of a system of true branchlets, conidiogenous cells <strong>and</strong> ramoconidia or at least a sequence of ramoconidia on several levels; conidia usually straight ......................................................................................................................... 3 3. Mycelium verruculose; long filiform conidiophores ending with a subdenticulate cell giving rise to sets of penicillate conidiogenous cells or ramoconidia which are barely distinguishable <strong>and</strong> turn into each other; ramoconidia <strong>and</strong> conidia consistently narrow, (1.5–)2(–2.5) µm wide, <strong>and</strong> aseptate, ramoconidia sometimes heterochromous; on living leaves of Nect<strong>and</strong>ra coriacea, Cuba ................... P. nect<strong>and</strong>rae 3. Mycelium more or less smooth; penicillate apex at least partly with true branchlets; conidia wider, 2–5 µm, at least partly septate, uniformly pigmented ................................................................................................................................................................................................. 4 4. Hyphae, conidiophores <strong>and</strong> conidia frequently distinctly constricted at the septa; penicillate apex of the conidiophores sparingly developed, branchlets more or less divergent; isolated from leaf litter of Smilax sp., Cuba .................................................................. P. rigidophora 4. Hyphae <strong>and</strong> conidia without distinct constrictions at the septa; penicillate apex of the conidiophores usually well-developed, with abundant branchings ................................................................................................................................................................................................. 5 5. Conidiophores short, up to 120 × 3–4 µm, frequently with intercalary conidiogenous cell, swollen at the conidiogenous portion just below the upper septum which render the conidiophores subnodulose to distinctly nodulose, apex ± loosely penicillate; conidia (4–)5–7(–8) µm long; occasionally with micronematous conidiophores; isolated from man with tinea nigra, Venezuela .......................... P. venezuelensis 5. Conidiophores much longer, up to 800 µm, 7–9 µm wide at the base, not distinctly nodulose, penicillate apex loose to often more compact, tight, metula-like; conidia longer, 7–25 × 2–5 µm; micronematous conidiophores lacking; isolated from dead leaf of Paepalanthus columbianus, Colombia ........................................................................................................................................................ P. columbiana Penidiella columbiana Crous & U. Braun, sp. nov. MycoBank MB504510. Figs 8–9. Etymology: Named after its country of origin, Colombia. Mycelium ex hyphis ramosis, septatis, levibus, pallide brunneis, 2–3 µm latis compositum. Conidiophora ex hyphis superficialibus oriunda, penicillata, erecta, brunnea, crassitunicata, minute verruculosa, ad 800 µm longa, ad basim 7–9 µm lata, ad apicem pluriramosa, ex ramibus diversibus et cellulis conidiogenis composita, ramibus primariis (–2) subcylindraceis, 1–7-septatis, 50–120 × 4–6 µm; ramibus secundariis (–2) subcylindraceis, 1–5-septatis, 40–60 × 4–6 µm; ramibus tertiariis et subsequentibus 1–4-septatis, 10–30 × 3–5 µm. Cellulae conidiogenae terminales vel laterales, non ramosae, 5–15 × 3–5 µm, modice brunneae, minute verruculosae, apicem versus attenuatae, truncatae vel rotundatae, polyblasticae, sympodiales, cicatrices conidiales incrassatae, sed leviter fuscatae et non refractivae. Ramoconidia 0–1-septata, modice brunnea, levia, ellipsoidea, obclavata vel obovoidea, cum 1–3 hilis terminalibus, 10–20 × 3–5 µm; conidia subcylindrica vel ellipsoidea, 0–1-septata, pallide brunnea, catenata (–10), hila truncata, non incrassata, vix vel leviter fuscata. Mycelium consisting of branched, septate, smooth, pale brown, 2–3 µm wide hyphae. Conidiophores arising from superficial mycelium, terminally penicillate, erect, brown, wall up to 1 µm wide, almost smooth to finely verruculose, up to 800 µm tall, 7–9 µm wide at the base; conidiogenous region consisting of a series of branches composed of true branchlets, conidiogenous cells <strong>and</strong> ramoconidia, branched portion usually rather compact, even metula-like, but also looser, with divergent branches; primary branches (–2), subcylindrical, 1–7-septate, 50–120 × 4–6 µm; secondary branches (–2), subcylindrical, 1–5-septate, 40–60 × 4–6 µm; tertiary <strong>and</strong> additional branches 1–4-septate, 10–30 × 3–5 µm. Conidiogenous cells terminal, intercalary or lateral, unbranched, 5–15 × 3–5 µm, medium brown, finely verruculose, tapering to a flattened or rounded (frequently swollen) apical region, scars thickened, but only somewhat darkened, not refractive. Ramoconidia 0–1-septate, 18
Phylogenetic lineages in the Capnodiales Fig. 8. Penidiella columbiana (type material). A. Conidiophores on pine needle in vitro. B–H. Conidiophores with chains of disarticulating conidia. Scale bars: A = 450, B–C = 10 µm. medium brown, smooth, wall ≤ 1 µm wide, ellipsoidal to obclavate or obovoid, with 1–3 apical hila, 10–25 × 3–5 µm, ramoconidia with broadly truncate base, not or barely attenuated, up to 4 µm wide, or at least somewhat attenuated at the base, hila 1.5–3 µm wide. Conidia subcylindrical to ellipsoid, 0(–1)-septate, pale brown, in chains of up to 10, 7–15 × 2–3 µm, hila truncate, unthickened, barely to somewhat darkened, 1–2 µm wide. Cultural characteristics: Colonies on PDA erumpent, spreading, with moderate aerial mycelium <strong>and</strong> smooth, even, submerged margins; olivaceous-grey in central part, iron-grey in outer region (surface); colonies fertile. Specimen examined: Colombia, Páramo de Guasca, 3400 m alt., isolated from dead leaf of Paepalanthus columbianus (Eriocaulaceae), Aug. 1980, W. Gams, holotype <strong>CBS</strong> H-19937, culture ex-type <strong>CBS</strong> 486.80. Notes: This isolate was originally identified as belonging to the Stenella araguata species complex. <strong>The</strong> latter name has been somewhat confused in the literature, <strong>and</strong> has been incorrectly applied to isolates associated with opportunistic human infections (de Hoog et al. 2000). <strong>The</strong> “araguata” species complex is treated elsewhere in the volume (see Crous et al. 2007a – this volume). www.studiesinmycology.org Penidiella cubensis (R.F. Castañeda) U. Braun, Crous & R.F. Castañeda, comb. nov. MycoBank MB504511. Fig. 10. Basionym: <strong>Cladosporium</strong> cubense R.F. Castañeda, Fungi Cubenses II (La Habana): 4. 1987. In vivo: Colonies on fallen leaves, amphigenous, effuse, pilose, brown. Mycelium usually external, superficial, but also internal, composed of branched, septate, brown, thin-walled, smooth to rough-walled hyphae, 2–3 µm wide. Stromata present, 40–80 µm diam, brown, immersed. Conidiophores densely fasciculate or in distinct synnemata, arising from stromata, erect, synnemata up to about 1000 µm long <strong>and</strong> (10–)20–40(–50) µm wide, individual threads filiform, pluriseptate throughout, brown, thin-walled (≤ 0.5 µm), smooth or almost so to distinctly verruculose, apically penicillate. Conidiogenous cells integrated, terminal <strong>and</strong> intercalary, 10–30 µm long, subcylindrical, terminal conidiogenous cells often slightly enlarged at the tip, with (1–)2–3(–4) terminal or subterminal subdenticulate conidiogenous loci, short conically truncate, 1–2 µm diam, unthickened or almost so, but often slightly refractive or darkened-refractive, intercalary conidiogenous cells usually with a single lateral locus just below the upper septum, conidiogenous cells giving rise to a single set of primary ramoconidia, or a sequence of ramoconidia at different levels. Ramoconidia cylindrical to ellipsoid-fusoid, 8–18(–25) × 2–3 µm, aseptate, pale olivaceous, olivaceous-brown to brown, thin-walled, smooth or almost so to 19
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