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Dipartimento di Biologia Vegetale, Mycotheca, Universitatis Taurinensis, Università di Torino, Viale, Mattioli 25, 10125 Torino, Italia
Vladimír Antonín
Moravian Museum, Department of Botany, Zelny trh 6, CZ-659 37 Brno, Czech Republic
Machiel E. Noordeloos
National Herbarium of the Netherlands, University of Leiden Branch, van Steenis Gebouw, Einsteinweg 2, P.O. Box 9514, 2300 RA Leiden, the Netherlands
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ABSTRACT |
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Crinipellis pedemontana, a new agaric growing on stems of dead grasses in an Italian urban park, is described and illustrated. It fits in sect. Grisentinae on account of the green reaction with KOH and ammonia and its bright colored pileus. The unique combination of purple-violaceous tinges in the pileus and the presence of chlamydospores in the pileipellis characterize this species as new. Its taxonomic position within Crinipellis and the related genus Moniliophthora is discussed.
Key words: Agaricales, chlamydospores, Marasmiaceae, taxonomy
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INTRODUCTION |
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An exhaustive search including monographic treatments and papers by Singer (1942
, 1976), Pegler (1968, 1977, 1986), Dennis (1970), Singer and Clémençon (1972), Redhead and Liu (1982), Paechnatz (1984), Candusso (1986, 1991), Redhead (1986, 1989), Doyle (1987), Halling (1993), Corner (1996), Antonín and Noordeloos (1997), Bon (1999), Takahashi (2000, 2002), Aime and Phillips-Mora (2005) and Arruda et al (2005), confirmed the unique nature of this species: Its characteristics do not match any published species. Therefore an extensive diagnosis, description and illustrations of this previously undescribed Crinipellis are provided.
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MATERIALS AND METHODS |
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. Macroscopic characters were examined from fresh material. Studies are based on dried material with a Leica DM 4500 B and an Olympus BX50 light microscope with magnification up to 1000x. Observations were made on mounts in these reagents: Congo red in 10% ammonia, 3% KOH, cresyl-blue in water and Melzer’s reagent. Measurements are based on observation of 30 basidiospores and do not include the apiculus. These abbreviations were used: E = the quotient of length and width of the spores; Q = the mean value of E values in all collections studied; L = number of entire lamellae; l = number of lamellulae between each pair of entire lamellae.
All examined material has been deposited and preserved in: MUT (Mycotheca Universitatis Taurinensis, Dipartimento di Biologia Vegetale, Università di Torino, Italy), BRNM (Moravian Museum, Brno, Czech Republic), and LIP–Herbarium M. Bon (Laboratoire de Botanique, Faculté des Sciences Pharmaceutiques et Biologiques, Université Lille II, France).
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TAXONOMY |
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–3
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, Les collybiomarasmïoïdes et ressemblants. Doc. mycol. Mém. hors Sér. 5, p. 74; misapplied name). Pileus (5)10–18(24) mm latus, primo hemisphaericus dein plano-convexus, explanatus et obtuse umbonatus, denique aetate depressus, siccus, squamuloso-velutinus, villosus, subzonatus, brunneo-incarnatus, brunneolo-purpureus, disco obscuriori; margo primo incurvus, mox explanatus, villosulus, fimbriatus, non striatus. KOH ope cuticola reactionem virescentem praebet. Lamellae crassiusculae, valde fere distantes, 2–3 mm latae, subadnatae vel prope liberae, sinuato-ventricosae, albo-cremeae, acie concolore, plus minusve intervenatae; lamellulae saepe cum lamellis concrescentes. Stipes 15–32(40) x (1.5) 2–3.5(4) mm, teres, subaequalis, non bulbosus, interdum plano-compressus, superne perspicue dilatatus usque ad 5 mm, fibrosus, solidus dein medullosus, subfistoloso-farctus; siccus, squamuloso-villosus, culmini albidus-cremeus, basi fusco-brunneus, pilis brunneofuscis inductus, maxime inferna parte. Sporata alba. Sporae ellipsoideae-amygdaliformes 7.5–9(10) x (4)4.5–6(6.5) µm, hyalinae, laeves, mono-biguttulatae, non pseudoamyloideae. Basidia claviformia, tetraspora 25–41 x 7–9 µm. Cheilocystidia versiformia, subfusoidea vel lageniformia, 25–42 x 7–11 x 2–4 µm, apice rotundato, interdum bifido vel moniliformi; pleurocystidia absunt. Pili pilei longissimi, 60–400 x 6–9(12) µm, pseudoamyloidei, KOH ope virescentes, flexuosi, crassetunicati, tunica fere 2.5 µm crassa, et apice rotundato. Pili stipiti 50–280(360) x 4–9 µm. Hypotrichium (subpileipellis) ex hyphis 25–40 x 6–12 µm latis, pigmento extracellulari incrustato KOH ope virescente praeditis. Pileipellis plurimis chlamydosporis 10–24 µm, tunica fere 4.5 µm crassa, large pediculatis. Fibulae copiosae.
Holotypus (hic designatus) in loco dicto Parco del Valentino, ad radices vegetalium et culmos graminearum (Phleum pratense) apud Augustam Taurinorum, Piemonte, Italia, a A. Vizzini lectus, 04-X-1992, in herbario Mycothecae Universitatis Taurinensis asservatur, MUT HS122. Isotypi in herbariis BRNM 705399 et LIP–Herb. M. Bon 930316 asservantur.
Etymology. pedemontanus = referring to the collection site, in Piedmont, a region in NW Italy (from 
o
, o
ó, pous, podos [Greek] meaning foot + mons, montis [Latin] meaning mountain, and thus "at the foot of the mountain").
Pileus (FIG. 1
) (5)10–18(24) mm broad, campanulate-convex, expanding to nearly applanate, depressed at center when old, often with an obtuse and distinct umbo, with involute, then straight margin, finally with irregularly undulate marginal zone, sometimes fimbriate, membranaceous to thin-fleshed, not translucently striate; dry, subzonate, not hygrophanous, radially subtomentose-fibrillose to fibrillose, with small adpressed-fibrillose squamules at center, often glabrescent with age, turning dark green with a drop of KOH or NH4OH; ochraceous (Se 174, 201, 202, 248) to pinkish malvaceous lilac (Se 69, 70), with purplish hues (Se 14–15, 165), darker at center (brown, Se 692, 702), and cream-ochraceus (Se 199) under hairy covering. Lamellae (FIG. 1) moderately distant, L = (16)20–26(30), l = (0)1–2–(3), thickish, slightly intervenose, sometimes partially fused with the lamellulae and then forked, emarginate-adnate to adnate, almost free, subventricose, up to 2–3 mm broad, white to cream-pink (Se 200), with entire, concolorous edge. Stipe (FIG. 1) 15–32(40) x (1.5)2–3.5(4) mm, cylindrical, often broadened up to 5 mm at apex, sometimes narrowed toward the base, not bulbous, sometimes compressed, stiff, solid to fistulose, dry, whitish-cream at apex (Se 200), gradually becoming darker toward the base (brown, Se 692), entirely subtomentose-fibrillose (apex) to squamulosestrigose (base). Context thin, tough, elastic, especially in stipe, whitish in stipe, cream ocher (Se 199) in cortex. Odor indistinct, flavor mild. Spore print whitish. Type of development of the basidiome pileostipitocarpic (ss. Reijnders 1963).
Basidiospores (FIG. 2b) 7.5–9.0(10) x (4.0)4.5–6.0(6.5) µm, E = 1.5–2.0, Q = 1.7, (broadly) ellipsoid, ellipsoid-amygdaliform, hyaline, smooth, thin-walled, mono- or biguttulate, nondextrinoid. Basidia 25–41 x 7.0–11 µm, 4-spored, clavate. Basidioles up to 38 x 5.0–10 µm, clavate or cylindrical. Cheilocystidia (FIG. 2a) 25–42 x 7.0–11 x 2–4 µm, lageniform, subfusoid, with obtuse, irregularly curved or branched apex, sometimes moniliform, hyaline, thin-to slightly thick-walled. Pleurocystidia absent. Hymenophoral trama subregular, made up of cylindrical, sometimes slightly inflated, thin-walled, 5.0–12(15) µm wide hyphae with nondextrinoid and in KOH hyaline or yellowish walls. Pileipellis (FIG. 2e) bi-layered, suprapellis a transition between a cutis and a trichoderm, made up of (semi)erect, thick-walled (0.5–2.5 µm), setiform terminal elements (FIGS. 2e, 3b; hairs, setae), 60–400 x 6.0–9.0(12) µm, gradually tapering toward the apex, with a narrow lumen and several secondary septa (pseudosepta); with brown to yellow walls that gradually become hyaline and colorless toward the apex, turning green in alkali and dextrinoid in Melzer’s reagent. Chlamydospores (FIGS. 2d, 3a) present in suprapellis, often in clusters, sphaeropedunculate, about 10–24 µm diam, with long, 5.5–7.0 µm wide pedicel, with 3–4.5(5) µm thick, yellow-brown, dextrinoid wall. Subpellis regular, made up of radially arranged, cylindrical elements, 25–40 x 6.0–12 µm, with brown-incrusted walls that turn green in KOH or ammonia. Stipitipellis similar to pileipellis, consisting of a trichoderm of setiform elements (FIG. 2c), 50–280(360) x 4.0–9.0 µm, with thick, dextrinoid walls. Stipititrama regular, made up of cylindrical, thin-walled, slightly dextrinoid, 4.0–12 µm wide hyphae with hyaline or slightly brownish walls. Clamp connections present in all tissues.
Ecology. – Solitary or gregarious, saprotrophic on herbaceous debris, on basal part of dead grasses (Phleum pratense L., Poaceae).
Material examined. – ITALY, Piedmont, Turin: Parco del Valentino, 4-X-1992, in meadow, near Cedrus atlantica (Endl.) Manetti, leg. A. Vizzini (HOLOTYPE, MUT HS122; Isotypes, BRNM 705399 and LIP–Herb. M. Bon 930316); ibidem 12-X-1994, leg. A. Vizzini, (Paratypus, Topotypus, MUT HS214).
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DISCUSSION |
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, Takahashi 2002), the majority having a pantropical distribution (Singer 1976, 1986).
Most Crinipellis species are saprotrophic on herbaceous or woody plants, except section Iopodinae (Singer) Singer (Singer 1986), whose members are parasitic on various woody plants. This section is characterized morphologically by the presence of purple, violaceous or red pigments, which do not stain green in KOH or ammonia. Recent molecular studies (Aime and Phillips-Mora 2005) have shown that Crinipellis perniciosa (Stahel) Singer, together with some anamorphic fungi parasitic on Cocoa, Theobroma cacao L., form a distinct monophyletic clade, separate from Crinipellis. They accordingly proposed to accommodate these fungi in genus Moniliophthora H.C. Evans, Stalpers, Samson & Benny. Although few data are known from other species of sect. Iopodinae the suggestion was made that all species described from that section eventually should be transferred to genus Moniliophthora. Crinipellis and Moniliophthora form together with Marasmius and Chaetocalathus a monophyletic group that may be considered a separate family Marasmiaceae (clade/marasmioid) within the large assemblage Tricholomataceae sensu lato (Moncalvo et al 2002, Bodensteiner et al 2004, Aime and Phillips-Mora 2005, Wilson and Desjardin 2005).
Crinipellis pedemontana is characterized by rather robust basidiomes, pileus with a pink-purple tinge, nondextrinoid basidiospores, brownish incrusted walls of subpileipellis hyphae turning green in KOH and especially by the presence of chlamydospores in the pileipellis, which may be scattered to common. These chlamydospores may be considered modified setiform hairs that are swollen apically. Therefore C. pedemontana fits well in sect. Grisentinae (Singer) Singer, which contains species with brown, lilac, red or purple tinges and whose pigments turn gray to green in alkaline solutions (KOH or ammonia) (Singer 1976, 1986).
The presence of chlamydospores (chlamydoconidia) is a unique character within Crinipellis. These structures more correctly should be called chlamydocytes, a more general term without functional attribute. Morphologically similar chlamydospores have been observed in the pileipellis and pileitrama (as apical and/or intercalary elements) of some species of Lentinellus (Hericiales, Lentinellaceae; Segedin 1996, Moreau et al 1999, Petersen and Hughes 2004), Panellus (Agaricales, Mycenaceae, called sclerocysts by Pöder 1985), Hemimycena (Agaricales, Mycenaceae, Moreau et al 2005), in basidiomes and/or mycelia of some species of polypores (Stalpers 2000) and on the cleistothecial wall of species of the genus Emericella (the so-called hülle cells, Carvalho et al 2002), a teleomorph of many Aspergillus species (Ascomycota, Eurotiales, Trichocomaceae), where they function as diaspores (Ellis et al 1973). However it is unlikely that these structures are homologous.
Bon (1999), who examined a specimen of the yet undescribed Crinipellis pedemontana, included it in his keys as a form (with a question mark) of Crinipellis alcalivirens Singer. However that species originally described from Colombia does not have pink-purple tinges in the pileus, has a slender stipe, smaller spores (7.8–8.5 x 4.5–5.7 µm), narrower and less pigmented setae, not incrusted subpellis (hypotrichium), abundant pleurocystidia and is found on culms of Bambuseae (Poaceae).
However the only species in Europe that sometimes has a lilac tinge in the pileus (viz. C. tomentosa [Quél.] Singer) differs not only by the lack of chlamydospores but also by having a much deeper brown, brownish gray, grayish-brown to ochraceous-gray pileus, up to 1–1.5(2) mm wide stipe, slightly larger basidiospores, (7.5)8.1–10.8(12.5) x (3.9)4.1–6.5 µm and distinctly shorter setae, only 150–200 µm long; furthermore the pigments of this species of Crinipellis do not stain in alkaline solutions (Antonín and Noordeloos 1997).
C. dusenii (Henn.) Singer, from Africa and South America on fallen leaves of dicotyledonous trees has a purple or bright red pileus, an umber or brown, sometimes also purple or chestnut-colored, slender (0.5–1.5 mm) stipe, narrower spores (3.3–5 µm), smaller basidia (18–27.5 x 6–8 µm), well developed pleurocystidia, and not incrusted hyphae in the supra-and subpileipellis (Pegler 1968, Singer 1976); C. rubida Pat. & R. Heim, found in Venezuela on fallen twigs and branches, has a bright pink pileus drying out to vinaceous, a longer and thinner [(5)40–60 x (0.7)1 µm] stipe concolorous with pileus, smaller (7.3–8 x 4–4.8 µm) basidiospores, which may become thick-walled and septate with age, smaller (20–28 x 6–6.7 µm) cheilocystidia with numerous apical projections, setae that remain hyaline or turn pale vinaceous in NH4OH, and nonincrusted hyphae in the pileipellis (Singer 1942, 1976).
As indicated above all species from sect. Iopodinae (Singer) Singer (the genus Moniliophthora), although their pileus color may be somewhat similar, differ by the absence of a green reaction in alkaline solutions and obviously by lacking chlamydospores.
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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1 Corresponding author. E-mail: alfredo.vizzini{at}unito.it
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LITERATURE CITED |
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