The genus Pleurotus in Argentina

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1 Mycologia, 96(4), 2004, pp by The Mycological Society of America, Lawrence, KS The genus Pleurotus in Argentina Bernardo E. Lechner 1 Jorge E. Wright PRHIDEB-CONICET, Departamento de Ciencias Biológicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires (1428), Buenos Aires, Argentina, and Instituto Fitotécnico de Santa Catalina, Universidad Nacional de La Plata, Facultad de Ciencias Agrarias y Forestales, Llavallol, Buenos Aires, Argentina Edgardo Albertó Instituto de Investigaciones Biotecnológicas IIB- INTECH (CONICET-UNSAM), C. C. 164, (7130) Chascomús, Buenos Aires, Argentina Abstract: Macro- and micromorphological characters of specimens of the genus Pleurotus (Fr.) P. Kumm. in Argentina obtained in the field and from different national herbaria were analyzed. Cultivation techniques were used to obtain basidiomata, allowing for a macro- and micromorphological study of fresh developing fruit bodies. We concluded that in Argentina there are, so far, six species, namely P. albidus, P. cystidiosus, P. ostreatus, P. pulmonarius, P. rickii and P. djamor, the latter with three varieties: var. djamor, var. cyathiformis and var. roseus. Key words: cultivation, Pleurotaceae, South America, taxonomy INTRODUCTION The systematic position of Pleurotus (Fr.) P. Kummer has been much debated. Singer (1951) transferred several species of Pleurotus sensu lato to Hohenbuehelia and placed the former in Polyporaceae, tribus Lentineae and the latter in the Tricholomataceae, tribus Resupinateae. Kühner and Romagnesi (1953) placed several pleurotoid genera, included Pleurotus, within the family Pleurotaceae, irrespective of sporeprint color and other micromorphological characters. Kühner (1980) placed Pleurotus and Hohenbuehelia S. Schulz. in the family Pleurotaceae, with Pleurotus in the tribus Lentineae and Resupinatus S. F. Gray in Resupinateae. Corner (1981) based the segregation of Pleurotus (Fr.) P. Kummer, Lentinus Fr. Accepted for publication January 21, Corresponding author. blechner@bg.fcen.uba.ar and Panus Fr. on the hyphal system and included several species of Singer s Pleurotus in Lentinus. Phylogenetic studies by Hibbett and Vilgalys (1993) and Hibbett and Donoghue (1995) have indicated that there are at least four distinct genera: Pleurotus, Lentinus, Panus Fr. and Neolentinus Redhead & Ginns. Within the genus Neolentinus there are several species that produce brown rot and were included by Pegler (1983) in Lentinus which, like Pleurotus, produces a white rot (Redhead and Ginns 1985). Thorn et al (2000) clarified the systematic position of Pleurotus through phylogenetic analysis based on partial sequences from nuclear 25S rdna that indicated a monophyletic Pleurotaceae with the monophyletic genera Pleurotus and Hohenbuehelia. The capacity of both genera to attack and consume nematodes (Barron and Dierkes 1977, Thorn and Barron 1984, 1986, Barron and Thorn 1987, Hibbett and Thorn 1994) supports the monophyly of this family. The knowledge of the genus Pleurotus in Argentina is scant; Spegazzini ( , 1887, 1898, 1926), Singer and Digilio (1951), Singer (1956, 1960, 1961a), Raithelhuber (1974, 1977, 1987, 1991) and Spinedi (1995) recorded 21 species and only four currently are accepted within Pleurotus (TABLE I). MATERIALS AND METHODS Macro- and micromorphology. Specimens were described macroscopically. The description of species combines cultures and herbarium specimens. Color names are in accordance with Munsell Color Co. (1954). Citation of authors of taxa is according to Kirk and Ansell (1992). Herbarium abbreviations follow Holmgren et al (1990). Specimens are deposited in the BAFC Mycological Herbarium. Microscopic examination of tissues mounted in 5% KOH and 1% aqueous phloxine was undertaken. Culture characters. Cultures were inoculated in 90 mm Petri dishes using Nobles (1948) medium and incubated in the dark at 25 C. Aspect, color, growth rate, odor and microscopic structures of both the aerial and the submerged mycelium were observed weekly for 6 wk according to Nobles (1965). Basidiome production. To obtain fresh basidiomata to complement the study of the macro- and micromorphology, traditional methods for fruiting species of Pleurotus were used (Zadrazil 1974, Stamets 1993). A mixture of sawdust (77%), wheat meal (15%), oatmeal (5%) and CaCO 3 (3%) was in- 845

2 846 MYCOLOGIA TABLE I. Species of the genus Pleurotus, their first citation in Argentina and the present name Species First citation Present name P. applicatus (Batsch. : Fr.) Quél. Spegazzini (1887) Resupinatus applicatus (Batsch. : Fr.) S. F. Gray P. araucariicola Sing. P. aulaxinus (Mont.) Sacc. P. calix (Speg.) Sing. Raithelhuber (1991) Spegazzini ( ) Singer (1956) Pleurotus pulmonarius (Fr. : Fr.) Quél. Favolaschia aulaxina (Mont.) Sing. Lentinus patulus Lév. P. eryngii (DC : Fr.) Quél. Singer & Digilio (1951) According to Singer & Digilio (1951), this species is represented in Argentina by the holotype of Agaricus tucala Speg., but this collection is not extant. P. heteropus Speg. P. hirneolus Speg. P. hirtus (Fr.) Sing. P. laciniatocrenatus Speg. Spegazzini ( ) Spegazzini (1898) Singer (1956) Spegazzini ( ) No holotype extant. No holotype extant. Lentinus scleropus (Pers.) Fr. Pleurotus albidus (Berk.) Pegler. P. lindquistii Singer (1960) Lentinus tigrinus Bull. : Fr. fide Grand et al (2002). P. minusculus (Speg.) Sacc. P. ostreatoroseus Spegazzini (1887) Singer (1961) Panellus longinquus (Berk.) Sing. Pleurotus djamor var. roseus Corner. P. perpusillus Fr. Spegazzini ( ) Pleurotus septicus Fr. fide Pilát 1935 (see under P. septicus). Singer (1952) named P. perpusillus (Speg.) Sacc., but we were unable to find this species cited by Spegazzini. P. perstrictifolius Speg. P. portegnus var. microsporus Sepg. P. pulmonarius (Fr. : Fr.) Quél. P. pusillimus Speg. P. rickii Bres. P. septicus Fr. P. smithii Guzman P. sutherlandii Sing. Spegazzini (1926) Spegazzini ( ) Spegazzini (1898) Spegazzini (1898) Singer & Digilio (1951) Spegazzini ( ) Spinedi (1995) Singer & Digilio (1951) Lentinellus perstrictifolius (Speg.) Sing. Hohenbuehelia grisea (Peck.) Sing. Pleurotus pulmonarius (Fr. : Fr.) Quél. Clitopilus argentinus Sing. Pleurotus rickii Bres. Probably a Crepitotus. Pleurotus smithii Guzman Panellus longinquus (Berk.) Singer. troduced in cm polypropylene bags and autoclaved at 120 C for 2 h. After cooling they were inoculated with spawn of different strains and incubated in the dark at 25 C. After 15 d, bags were kept at C with 9 h light/15 h dark photoperiod to induce basidiome formation. Strains used. P. albidus: ARGENTINA, BUENOS AIRES, La Lucila, in canker of Salix humboldtiana, 5-V-1996, BAFC 2787; La Plata, on stump, III-1996, coll. J. Deschamps, BAFC 809; Pergamino, on Salix sp., 6-IV-1996, coll. E. Albertó, BAFC 695; Tigre, on Populus sp., 10-III-1997, coll. C. Lázzari, BAFC 136; CAPITAL FEDERAL, on Cordyline sp., 10- VI-1989, coll. A. Martinez, BAFC 2545; CÓRDOBA, La Punilla, on Populus sp., 25-II-2001, coll. N. Manero & B.J. Lechner, BAFC 190; MISIONES, Pto. Libertad, on dead tree of Araucaria angustifolia, 10-IV-1985, coll. J. Deschamps, BAFC 215. P. cystidiosus: ARGENTINA, CAPITAL FEDERAL, Barrio La Paternal, on dead zone of Platanus sp., coll. S. Frachia, 18-IV-2000; BAFC 188. P. djamor var. djamor: ARGEN- TINA, MISIONES, El Palmital del cruce, 28-V-2001, coll. E. Albertó, BAFC 821. P. djamor var. roseus: ARGENTINA, MI- SIONES, El Palmital del cruce, 28-V-2001, coll. E. Albertó, BAFC 815; BRAZIL, (as P. ostreatoroseus), on leaf of Agavaceae, MUCL P. ostreatus: ARGENTINA, CAPITAL FEDERAL, 18-IV-1994, coll. P. Pica, BAFC 2034; NEUQUEN, Moquehue, on trunk of Araucaria araucana, III-1993, coll. J. del Vas, BAFC 120; ITALY, (as P. columbinus), isolated from commercial fruit body, isolated and determined by T.H. Quimio, MUCL P. pulmonarius: ARGENTINA, BUENOS AIRES, Ezeiza, on living declining tree of Populus sp., 25-VII-1987, coll. J. Deschamps, BAFC 1003; MISIONES, San Pedro, on branches of Araucaria angustifolia, 27-V- 2001, coll. E. Albertó & O. Popoff, BAFC 76; San Pedro, S, W, on branches of A. angustifolia, 27-V-2001, coll. D. Krueger, BAFC 213; same location, same host, same date, coll. E. Albertó, BAFC 263. TAXONOMY Pleurotus (Fr.) P. Kummer (1871) Champignons du Jura & Vosges p em., nomen conservandum. Spec. Typ.: Pleurotus ostreatus ( Jacq. et Fr.) P. Kummer. Syn.: Agaricus trib. Pleurotus Fr., Syst. Mycol. 1: Crepidotus Nees ex S. F. Gray, Nat. Arr. Brit. Pl. 1: Pleurotus P. Kummer, Führ. Pilzk. p Pleurotus Quél., Enchir., p Dendrosarcus Paulet ex Kunze. Rev. Gen. Pl. 2: nom. nud. Antromycopsis Pat. & Trabut, Bull. Soc. Myc. Fr. 13: (imperfect form of sect. Coremiopleurotus (cfr. Pollack & Miller, Mem. N. Y. Bot. Gard. 28: ).

3 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 847 Lentodiopsis Bubák, Hedwigia 43: (type L. albida Bubák, l. c.). Lentodiellum Murr., Mycologia 7: (type Panus concavus Berk.). Nothopanus Sing., Mycologia 36: Pterophyllus Lév., Ann. Sc. Nat. III 2: (type P. bovei Lév.). Basidiomes usually large, fleshy, solitary to imbricate, flabellate to dimidiate, glabrous to tomentose, white, cream, gray, pink, brown, more rarely blue, yellow or lilac. Stem short, solid, eccentric to lateral, rarely subcentral. Lamellae decurrent, sometimes anastomosing to the stem, light-colored, thin to broad, margin entire. Veil present or absent at margin of pileus or forming an annular zone on the stem. Spore print white, cream, pinkish or lilac. Spores cylindrical to subcylindrical, thin-walled, hyaline, not amyloid or dextrinoid, without germ pore. Cheilocystidia absent or poorly developed, disappearing early, thin-walled, clavate or mucronate. Subhymenium well developed, hymenophoral trama irregular. Pileipellis frequently poorly developed, with parallel radial hyphae, sometimes pigmented. Hyphal system monomitic or dimitic, gelatinous tissue usually absent; clamps present. Lignicolous. Observations. The genus Pleurotus does not have clear distinctive characters as occurs with other genera. The hyphal system may be monomitic or dimitic, without binding hyphae. This character might confuse the boundaries with other related genera, such as Lentinus (Stankovičová, 1973). There exists a clear delimitation in Pleurotus between species with a monomitic hyphal system, thin- or thick-walled generative hyphae, such as P. ostreatus, P. pulmonarius, P. albidus, P. cystidiosus, and dimitic ones, such as P. djamor and P. rickii. All Pleurotus species have thin-walled, smooth, cylindrical to subcylindrical spores. They may be small ( 8 m long) such as P. auriovillosus (Corner 1981), medium (8 12 m) such as P. albidus and large ( 13 m long) such as P. cystidiosus. The pileipellis is generally a cutis. In P. cystidiosus and P. djamor, there may be pileocystidia. It is important to stress the significance of the broad pileipellis to differentiate P. ostreatus from P. pulmonarius, the latter much thinner than the former. One of the distinctive characters of the genus is an irregular hymenophoral trama combined with a well developed subhymenium that, according to Singer (1986), is always more than 7 m thick. Pleurotus does not exhibit chemical reactions. Oleiferous hyphae can be distinguished in different tissues of the basidiocarp that turn yellow with 5% KOH. Spores of all known species of Pleurotus are negative in Melzer s solution. Mycologists who studied this taxon (Singer 1986, Kühner 1980, Pilát 1935, Hilber 1982, Moser 1986, Watling and Gregory 1989) have subdivided it differently. In this paper, we adopt the subdivision proposed by Hilber (1982), who divided the genus Pleurotus in three subgenera, to which we add Subgenus tuberregium. Singer (1961b, 1986) considers this a subgenus of Pleurotus and Pegler (1975, 1983) a species of Lentinus. Thorn et al (2000) have demonstrated that its mycelium has microdroplets. We have not found this species in Argentina. Subgenus Tuberregium Sing.: Pileus white to brown, veil not well developed, Stipe eccentric to subcentral; without coremia; Hyphal system dimitic. A conspicuous sclerotium is produced. Subgenus Pleurotus: Hyphal system in mature specimens monomitic or dimitic, with a distinct stipe; neither coremia nor veil present. Spore print white, cream, lilac or pinkish. i) Species with a monomitic hyphal system: P. ostreatus ( Jacq.: Fr.) P. Kummer (with var. ostreatus and var. columbinus Quél. apud Bres.), P. pulmonarius (Fr.) Quél. and P. albidus (Berk.) Pegler. ii) Species with a dimitic hyphal system: P. djamor (Fr.) Boedijn, with three varieties, according to Corner (1981): var. djamor, var. roseus Corner and var. cyathiformis Corner. Subgenus Coremiopleurotus O. Hilber: coremia present on lamellae, base of stipe or mycelia or on the substrate; pileocystidia present and well developed; with or without veil; stipe eccentric or lateral. Hyphal system monomitic; spore print white, cream, never livid or pinkish. P. cystidiosus O.K. Miller Subgenus Lentodiopsis (Bubák) O. Hilber: Basidiocarps with conspicuous veil; distinct stipe; hyphal system dimitic, without coremia. P. rickii Bres. KEY TO SPECIES OF PLEUROTUS IN ARGENTINA 1. Hyphal system dimitic Hyphal system monomitic Basidiome with a conspicuous veil; stipe well developed... P. rickii 2. Basidiome without a veil Pileus cyathiform, spores m P. djamor var. cyathiformis 3. Pileus not cyathiform Pileus white, stipe lateral or absent, spores m... P. djamor var. djamor 4. Pileus pinkish, spores m......p. djamor var. roseus

4 848 MYCOLOGIA 5. Coremia present on the substrate, pileocystidia present, well developed, stipe eccentric or lateral P. cystidiosus 5. Coremia not present Pileus white to cream, margin often lacerate-crenate... P. albidus 6. Pileus grayish to grayish brown, margin entire, rarely lacerate-crenate Pileipellis larger than 100 m thick. Fruiting from autumn to winter, odor pleasant, not anise-like P. ostreatus 7. Pileipellis not larger than 60 m broad, fruiting in summer to autumn, odor often anise-like P. pulmonarius Pleurotus albidus (Berk.) Pegler FIG. 1. Pleurotus albidus, macromorphology. A. collected basidiocarps (BAFC ); B. young basidiocarps obtained from strain BAFC 136 and C. mature; D. mature basidiocarps obtained from strain BAFC 695; E. Idem D. Scale bar 5 cm for A and C; 4 cm for B, D and E. Kew Bull., Addit. ser. 10:219, Lentinus albidus Berkeley Hooker J. Bot. 2:633. Lentinus calvescens Berkeley & Curtis Hook. J. Bot. 8:143. Pleurotus jacksonii Berkeley & Cooke J. Linn. Soc. London 15:363. Panus laciniato-crenatus Spegazzini An. Soc. Cient. Arg. 9:164. Pleurotus laciniato-crenatus (Speg.) Spegazzini Bol. Acad. Nac. Cienc. Córdoba 23: Basidiocarp cornucopioid, clitocybioid, omphalloid to lentinoid (i.e., L. crinitus), usually gregarious in troops of 2 30 basidiomata. Pileus circular, infundibuliform (FIGS. 1, 11A, B), mm broad mm long, glabrous, hygrophanous, white to cream (C2Pl9, Maerz and Paul 1930), sometimes with brown color next to the attachment of the stipe, margin entire to laciniate-crenate, thin, usually incurved (especially when dry), frequently lemon yellow through drying. Lamellae deeply decurrent, becoming anastomosing ridges on the stipe, subdistant to crowded, 6 8 mm broad, white. Stipe mm long 5 33 mm diam, upward fluted through lamellar bases, tapering downward, usually curved-ascendant, dull white, felty, occasionally with hairs at the base. Spore print white to cream. Odor fungal. Usually on fallen trunks, rarely living trees of Salix sp., Ulmus sp. and Platanus sp. Basidiospores (5.2 )8 9.5( 11.4) m; Q 3.15 long, very variable in length, oblong, smooth, hyaline, thin-walled, IKI- (FIG. 2A). Basidia m subcylindrical to clavate, hyaline, 4- spored, numerous, fusiform to mucronate, nonemergent basidioles measuring like the basidia (FIG. 2C). Cheilocystidia absent. Hyphae of pileus context generative, hyaline, frequently branched, m diam (FIG. 2B). Hyphae of hymenophoral trama m diam, irregular to subparallel, close, occasionally oleiferous hyphae present, conspicuously yellow in KOH (FIG. 2C). Subhymenium not well developed, m thick, with pseudoparenchymatous, isodiametric, hyaline hyphae (FIG. 2C). Hyphae of stipe m diam, hyaline, generative, thinwalled and thick-walled (more than 0.4 m thick), branched (FIG. 2D); on the surface, hairs formed by hyaline, generative, branched, thick-walled ( m thick) hyphae m diam Hyphae of pileipellis m diam, hyaline, thin- to thick-walled; pileocystidia not observed. Culture characters. Mycelium hyaline, with thick strands that give a feathery aspect, whitish, cottony zones near inoculum. Colony growth radial, loose and felty. Margin irregular, festooned, to regular according to the strain studied. Odor like almonds. Mycelial mat covering Petri plates in 3 4 wk. Occasionally, orange exudate droplets produced principally in the center and near the inoculum when all Petri plates were fully grown. Advancing mycelium with hyaline, thin-walled, clamped hyphae m diam. At the center of the colony generative, clamped, broader and less branched hyphae m diam. The number of

5 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 849 FIG. 2. Pleurotus albidus, micromorphology. A. Spores; B. Hyphae of the pileus; C. Hymenophoral trama; D. Hyphae of stem. Scale bar 20 m. hyphal branches increases when the colony becomes old, near the inoculum m diam. Microdroplets m diam. Geographic distribution. Northeastern and central Argentina. Material studied. ARGENTINA, BUENOS AIRES, San Clemente del Tuyu, Parque Municipal, on fallen trunk of Salix sp., 13-VI-1999, leg. E. Albertó, BAFC ; Costa del Este, at the base of trunk of living Platanus sp., 17-III-2002, leg. B.E. Lechner, BEL 79, (BAFC ); Monte Grande, on Populus sp., 25-V-1997, leg. M. Vallejo, BAFC ; Llavallol, Sta. Catalina forest, on fallen trunk, 26-III-1998, leg. E. Albertó, BAFC ; same location, same substrate, 6- V-1999, leg. Albertó & Pire, BEL 10 (BAFC ); same location, 15-VI-1999, leg. Albertó & Pire, BAFC ; same location, 1-V-2000, leg B. Lechner, BEL 40 (BAFC ); same location, 8-V-2000, leg. B. Lechner, BEL 39 (BAFC ); Punta Lara, Boca Cerrada, 20-VII-1971, leg. Gomez & Del Busto, BAFC ; same location, on fallen trunk of Salix humboldtiana, 1-VI-1949, leg. Lindquist & Singer, S 60; Parque Pereyra, on trunk of Ulmus sp., 4-VI-1999, leg. Albertó, Pereyra & Analía, BAFC ; Escobar, in canker of Populus sp., 18-VI-2000, leg. B. Lechner, BEL 80 (BAFC ); same location, 23-VI-2002, BAFC ; Bella Vista, 13-III-1996, leg. P. Pica, BAFC ; Llavallol, fruit bodies obtained from strain BAFC 2787, 13-VI-2000 and 14-VI- 01, leg. B. Lechner, BAFC ; basidiocarps obtained from strain BAFC 136, XII-99 and 13-VI-2000, leg. B. Lechner, BAFC ; fruit bodies obtained from strain BAFC 809, leg. B. Lechner, 13-X-2000, BAFC , 31-II-2000, BAFC and V-1999, BAFC ; fruit bodies obtained from strain BAFC 695, 28-VI-1999, leg. B.E. Lechner, BAFC ; fruit bodies obtained from strain BAFC 190, 19-VI-2001, leg. B.E. Lechner, BAFC ; fruit bodies obtained from strain BAFC 215, leg. B. Lechner, 4-VIII-2000, BAFC , 31-XI-2000 (BAFC ), 6-VI-2000 (BAFC ); Castelar, INTA, leg. E. Albertó, BAFC ; CAP- ITAL FEDERAL, 15-III-1996, leg. J. Deschamps, BAFC ; CÓRDOBA, Villa Giardino, La Punilla, on Populus sp., 17-VII-2000, leg. B. Lechner, BEL 55 (BAFC ); same location, 25-II-2001, leg. N. Manero & B. J. Lechner, BEL 70 (BAFC ); TUCUMÁN, Sierra San Javier, Parque de Aconquija, ad truncum vetustum (Phoebe porphyria), 10-VII-1949, leg. R. Singer, T 599; Departamento de Tafí, Río de los Sosas, on rotten trunk, 26-II-1952, leg. R. Singer, T 1879 (LIL). Remarks. The hyphal system of these basidiocarps is monomitic, similar to several Pleurotus species such as P. ostreatus, P. pulmonarius, P. populinus Hilber & Miller and P. abieticola Petersen & Hughes (Albertó et al 2002). P. albidus is characterized by its fragile nature, white to cream color and its remarkable laciniate-crenate margin. Several specimens exhibited great morphological variability, observed not only in collections from nature but also in fruit bodies obtained in culture. It seems as if certain species form basidiocarps similar to P. cornucopiae, characterized by their anastomosed lamellae on the stipe, such as BAFC 809; whereas others form more plane and shorter basidiocarps, without such characteristics, as BAFC 136 (see FIGS. 1, 11A, B). P. albidus is the species of major distribution in the Argentina, with several collections and strains obtained from diverse species of living trees to stumps and fallen trunks. Pleurotus cystidiosus O.K. Miller Mycologia 61: Nomen anamorphosis: Antromycopsis macrocarpa (Ellis & Everh) Stalpers, Seifert & Samson. Basidiome solitary to imbricate (FIGS. 3A, 11C). Pileus cm, pleurotoid, surface brown to grayish brown (Munsell, 7.5YR, 5/4 10YR, 5/2) with numerous punctiform squamules formed by surface cracking, more numerous toward the margin; margin entire, slightly to much festooned, hardly involute. Stipe brown (Munsell, 10YR, 3/3) to grayish (Munsell, 10YR, 5/1 6/1) almost lateral

6 850 MYCOLOGIA FIG. 3. Pleurotus cystidiosus. A. Basidiocarp; B. Spores; C. Basidia; D. Microscopic characters of anamorph. Scale bar 1 30 m for B D; Scale bar 2 10 cm for A. FIG. 4. Pleurotus cystidiosus. A. Hyphae of hymenophoral trama; B. Hyphae of context; C. Hyphae of stem; D. Pileocystidia. Scale bar 30 m. 3.5 cm, tapered to the base. Context white mm broad, fleshy when fresh, compact, corky when dry. Lamellae white when fresh, yellow (Munsell, 2.5YR, 8/8) when dry, 4 10 mm broad, thinner toward the stipe, decurrent, and forming a pseudoreticulum. Odor agreeable, fungal. Edibility: good. Spores hyaline, cylindrical-oblong, thin-walled, smooth, (11.0 )12 16( 18.0) m Q 3.01 (FIG. 3B). Basidia 2-, 3- and 4-spored, m (FIG. 3C). Subhymenium narrow, inconspicuous, ca 5 m thick. Cheilocystidia absent. Pleurocystidia (?) with granular contents, clavate, m. Pileipellis m broad, formed by clamped, brown, parallel, scanty branched, 3 5 m diam hyphae; among them brown, thick-walled, cystidiform hyphae, m diam (FIG. 4D). Context monomitic, formed by clamped, thin-walled, hyaline hyphae, m diam (FIG. 4B). Hyphae of stipe similar to those of context, with cystidiform ends m, and thick-walled, m diam (FIG. 4C). Hymenophoral trama formed by generative, thin-walled, clamped m diam hyphae, and thick-walled m diam (FIG. 4A). Anamorph. Coremia black, caespitose, formed by a globose head, mm diam, and a stem mm long and mm diam (FIG. 11D), formed by thin-walled clamped hyphae, which slowly disarticulate at the ends into fuliginous, cylindrical conidia m (FIG. 3D). Culture characters. Mycelium hyaline, filamentous, dense, whitish near the inoculum, growth radial, well adhered to the substrate. At first, a large amount of hyaline, aerial mycelium was observed, more whitish later; coremia appearing on the inoculum, but in strain BAFC 188, they later appeared away from it. Different densities in the mycelium of the colonies originate folds. At the fourth week, discontinued growth. Odor disagreeable, putrid. Mycelium not covering the Petri plates in 6 wk. Advancing mycelium with hyaline, clamped, thinwalled, regularly branched hyphae m diam. At the middle zone of the colony hyaline, clamped, regular to very branched, contorted hyphae m diam. Hyphae near inoculum less branched, with granular content m diam. Microdroplets m diam. Geographic distribution. East central Argentina. Material studied. ARGENTINA, BUENOS AIRES, Llavallol, fruit bodies obtained from strain BAFC 188, leg. B.E. Lechner, 24-VII-2001 (BAFC ), 9-X-2001 (BAFC ) and III-2002 (BAFC ); La Plata, Plaza San Martín, XII , leg. H. Spinedi, LPS No ; CAP- ITAL FEDERAL, barrio de La Paternal, on dead zone of Platanus sp., IV , leg. S. Fracchia, BAFC: ; SANTA FE, Rosario, Fisherton, III-2000, leg. M. Dimarco,

7 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 851 BAFC ; UNITED STATES, INDIANA, Brown Country, Hoosier National Forest, fruit bodies obtained from culture of a white, stringy trunk rot of a living Acer rubrum; leg. O.K. Miller, OKM 7122, HOLOTY PE. Remarks. P. cystidiosus is characterized by a squamulose pileus surface with pileocystidia and the presence in culture of an anamorph phase with coremia. The study of collections of P. smithii from LPS shows that the pileus had pileocystidia and its macro- and micromorphological characters point to P. cystidiosus. The basidiocarps were collected in warm seasons (summer and early autumn) and on living, not autochthonous trees. This could explain the difficulties to produce fruit bodies artificially. Pleurotus djamor (Fr.) Boedijn in De Wit, H.C.D., Ed., Rumphius Memorial Vol., p. 292, Agaricus djamor Rumphius apud Fries Syst. Mycol. 1:185. Agaricus arboreus secundus Rumphius Fl. Amboin. 11:125. Agaricus caryophyllus Berkeley J. Linn. Soc. 13:157. Agaricus emerici Berkeley Gard. Chron. 21(2):240. Agaricus ëous Berkeley Hooker, J. Bot. 2:83. Pleurotus ëous (Berk.) Saccardo Syll. Fung. 5:361. Agaricus flabellatus Berkeley & Curtis J. Linn. Soc. 11:528. Agaricus griseo-roseus Montagne Syll. Gen. Spec. Cryptog., p Pleurotus griseo-roseus (Mont.) Saccardo Syll. Fung. 5:386. Agaricus luteoalbus Beeli Bull. Soc. Roy. Bot. Belge 60:163. Agaricus leptogramme Berkeley & Broome J. Linn. Soc. 11:529. Agaricus moselei Berkeley Challenger. No. 37. Agaricus ninguidus Berkeley Hooker s J. Bot. 2:84. Pleurotus ninguidus (Berk.) Saccardo Syll. Fung. 5: 361. Pleurotus ostreatoroseus Singer Publ. Inst. Mic., Univ. Recife 304:10. Agaricus pacificus Berkeley London J. Bot. 1:451. Agaricus placentodes Berkeley Hooker s J. Bot. 4: 104. Agaricus prometheus Berkeley & Curtis Amer. Acad. Arts & Sci. 4: No. 30. Pleurotus salmoneostramineus Vasil eva Agar. & Bol. Primorsk Reg., p. 85. Pleurotus scabellus Saccardo Syll. Fung. 5:374. Agaricus scabriusculus Berkeley J. Linn. Soc. Bot. 13:157. Pleurotus scabriusculus (Berkeley) Saccardo Syll. Fung. 5: 374. Pleurotus djamor (Fr.) Boedijn var. djamor FIG. 5. Pleurotus djamor var. djamor. A. Basidiocarp; B. Hyphae of the hymenophoral trama; C. Hyphae of the pileus; D. Hyphae of the stem. Scale bar 1 5 cm for A; scale bar 2 30 m for B D. in De Wit, H.C.D., Ed., Rumphius Memorial Vol., p. 292, Pileus white mm, spathulate to flabelliform (FIGS. 5A, 11E), brown when dry; glabrous, smooth to touch; margin entire, frequently lobate and imbricate in old specimens. Context mm thick, always less than 1 mm when dry. Lamellae white 2 4 mm broad, mm in dry state, decurrent, crowded, smooth, margin entire. Stipe 5 6 mm long 3 4 mm diam, lateral, smooth, rarely absent. Spore print white. Odor not distinct. Good edibility. Spores (6.0 ) ( 7.80) m Q 2.44, cylindrical-oblong, hyaline, not dextrinoid or amyloid, thin-walled. Basidia club-shaped m. Cheilocystidia mucronate m. Hymenophoral trama dimitic, with sharp-pointed skeletal hyphae m and clamped generative hyphae m diam (FIG. 5B). Pileipellis m broad with hyphae arranged parallel to the pileus but somewhat irregularly entangled. Context with generative hyphae m diam and skeletals m (FIG. 5C). Stem context with narrower generatives, m diam and skeletals m diam (FIG. 5D). Culture characters. Mycelium hyaline, filamentous, radial growth, well adhered to substrate. Margin ir-

8 852 MYCOLOGIA regular, undulate. Against the light, an unequal refraction was observed, causing a relief aspect. Odor not distinct. The mycelium did not cover the Petri plates in 6 wk. Pinkish zones were observed in some strains. Advancing mycelium with hyaline, thin-walled, clamped, scantly branched m diam hyphae. In the middle of the colony with some contorted hyphae m diam. Near the inoculum with regularly branched hyphae m diam. Microdroplets m diam not observed in dikaryotic mycelia, but present in monokaryotic mycelia. Geographic distribution. Northeastern Argentina. Material studied. ARGENTINA, MISIONES, Iguazú National Park, San Martin Island, 3-VI-1998, leg. E. Albertó, BAFC ; El Palmital del Cruce, 28-V-2001, leg. E. Albertó, BAFC ; BOLIVIA, Guayaramerín, Prov. Vaca Diez, Dpto. Beni, 16-III-1956, leg. R. Singer, B 2028 (BAFC ; BAFC ). Remarks. P. djamor var. djamor is a white pantropical species characterized microscopically by having a dimitic hyphal system. Pleurotus djamor var. roseus Corner Beih. Nova Hedw. 69, p Pileus mm, spathulate to flabelliform (FIGS. 6A, 11F), pinkish when fresh, whitish or yellowish upon aging, clear brown when dry. Surface smooth, glabrous. Context 1 6 mm thick, less than 1 mm when dry. Lamellae decurrent, close to crowded, whitish, smooth, margin entire. Stipe 3 7 mm long 3 6 mm diam, whitish when fresh, cream when dry, eccentric, almost lateral. Surface irregular. Spore print white. Spores (6.7 ) ( 9.9) m Q 2.8, cylindrical-oblong, hyaline, smooth, neither dextrinoid nor amyloid, thin-walled (FIG. 6B). Basidia club-shaped m, 4-spored. Numerous basidioles or pleurocystidia. Cheilocystidia m, lageniform to mucronate (FIG. 6C). Hymenophoral trama completely irregular, dimitic, with clamped hyphae m diam and skeletals 3 6 m diam, unbranched (FIG. 6D). Subhymenium m thick, well developed, formed by irregularly shaped cells (FIG. 6D). Pileipellis with repent hyphae densely arranged toward the surface. Context dimitic, with skeletal, sharp-ended, m long hyphae, m diam, and thin-walled hyphae, generative, m diam (FIG. 6E). Context of stem formed by generative hyphae 2 4 m diam and skeletals m (FIG. 6F). Geographic distribution. Northeastern Argentina. Material studied. ARGENTINA, BUENOS AIRES, Llavallol, Sta. Catalina, fruit bodies obtained from strain BAFC 815, 25-IX-2001, leg. B.E. Lechner, BAFC ; fruit bodies FIG. 6. P. djamor var. roseus. A. Basidiocarps; B. Spores; C. Cheilocystidia; D. Hymenophoral trama and hymenium; E. Hyphae of the pileus; F. Hyphae of the stem. Scale bar 1 5 cm for A; scale bar 2 30 m for B F. obtained from strain MUCL , 20-X-2000, leg. B.E. Lechner, BAFC ; MISIONES, Iguazú National Park, San Martin Island, 3-VI-1998, leg. E. Albertó, BAFC ; El Palmital del Cruce, 28-V-2001, leg. E. Albertó, BAFC Remarks. The basidiocarps of this variety are pinkish or salmon-pink when young, discoloring to pale pink, whitish to yellowish with age. Distinction of var. djamor from var. roseus probably has been confused often. Pleurotus djamor var. cyathiformis Corner Beih Nova Hedw 69, p Pileus white mm, cyathiform, surface smooth (FIGS. 7A, 11G), with brown stains that disappear when dry. Context white 1 5 mm broad, less than 1 mm when dry. Lamellae decurrent, close to crowded, smooth, margin entire. Stipe mm long 2 4 mm diam, eccentric. Spore print white. Spores (6.2 ) ( 7.3) m Q 2.58, cylindrical-oblong, hyaline, neither dextrinoid nor amyloid, thin-walled (FIG. 7B). Basidia m, club-shaped, 4-spored (FIG. 7C). Numerous basidioles or pleurocystidia present (FIG. 7C). Cheilocystidia not observed. Hymenophoral trama

9 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 853 FIG. 7. P. djamor var. cyathiformis. A. Basidiocarps; B. Spores; C. Portion of hymenia, subhymenia and hymenophoral trama; D. Hyphae of the stem; E. Hyphae of the hymenophoral trama; F. Hyphae of the pileus flesh. Scale bar 1 3 cm for A; scale bar 2 30 m for B F. FIG. 8. Pleurotus ostreatus. A. Basidiocarp; B. Spores; C. Basidia; D. Hyphae of the trama; E. Hyphae of the stem; F. Hyphae of the pileus. Scale bar 1 10 cm for A; scale bar 2 20 m for B F. completely irregular, monomitic, with thin-walled generative hyphae m diam, some with slightly thickened walls (FIGS. 7C and E); subhymenium m thick, well differentiated, with hyphae more spherical than in other varieties (FIG. 7C). Pileipellis m, with repent hyphae arranged in parallel, although sometimes somewhat entangled. Context dimitic, with generative hyphae m diam and skeletals m diam (FIG. 7F). Stem also dimitic, with generative hyphae m diam and skeletals m diam (FIG. 7D). Geographic distribution. Northeastern Argentina. Material studied. ARGENTINA, MISIONES, Palo Rosa forest (El Palmital), on route 101, 4-VI-1998, leg. E. Albertó, BAFC Pleurotus ostreatus ( Jacq.: Fr.) P. Kummer Führer der Pilzkunde: Agaricus ostreatus Jacq Fl. Austr. 3: pl Agaricus ostreatus Jacq.: Fr Syst. Mycol. 1:182. Agaricus salignus Pers Syn. Meth. Fung: 479. Agaricus salignus Pers.: Fr Syst. Mycol 1:183. Pleurotus salignus (Pers.: Fr.) P. Kummer Führ Pilzk: 105. Pleurotus columbinus Quel in Bres., Fungi tridentini 1:10. Pleurotus floridanus. Singer Pap. Mich. Acad. Sci. 32:134. Pleurotus florida Eger Arch. Mikrobiol. 50:343. nom. nud. Pileus mm when dry; caespitose, smooth (FIGS. 8A, 11H J), margin involute at first, solid, brownish yellow (Munsell, 10YR 6/6). Basidiomata obtained in culture with pileus mm, with similar characters to collections obtained in field but the basidiocarps developing more isolated. Lamellae decurrent, close, broad, ca 5 mm, not very thick, pale yellow (Munsell, 2,5Y 8/4), margin entire. Stipe mm, eccentric-lateral, short, surface smooth. Flesh white, thin, ca 2 mm thick, corky. Odor fungal. Spore print white to cream. Spores (8 ) ( 13) m Q 2.4; cylindrical, hyaline, not amyloid, smooth (FIG. 8B). Basidia m, club-shaped, 4-spored (FIG. 8C). Cheilocystidia not observed. Pleurocystidia

10 854 MYCOLOGIA FIG. 9. Pleurotus pulmonarius. A. Basidiocarp; B. Spores; C. Hyphae of the pileus; D. Hymenium, subhymenium and hymenophoral trama; E. Hyphae of the stem. Scale bar 1 5 cm for A; scale bar 2 20 m for B E. absent or similar to basidioles m. Subhymenium m broad. Hymenophoral trama not completely irregular, with sclerified hyphae in some collections m diam, or thin-walled, m diam (FIG. 8D). Oleiferous hyphae not observed. Hyphae of context branched m (FIG. 8F). Hyphae of stipe thin-walled m diam, scantily branched and constant diameter to sclerified m diam (FIG. 8E). Pileipellis variable, according to the collections, m thick; generally larger than 100 m thick. Monomitic. Culture characters. Mycelium hyaline, growth feathery, with whitish arachnoid zones near the inoculum; increased aerial mycelium and more cottony at the second week. Margin irregular. Bleached zones at reverse of agar. Odor not distinct. The mycelium covered Petri plates between 1 and 2 wk. Primordia of fruit bodies were observed. Advancing mycelium with generative, clamped, hyaline, regularly branched hyphae m diam. In the middle of the colony hyphae are generative, clamped, thin-walled, regularly branched, m diam. Mycelium near inoculum with generative, thin-walled, regularly branched and variable diameter hyphae m diam. Microdroplets m diam. Geographic distribution. East central and southwestern Argentina. Material studied. ARGENTINA, BUENOS AIRES, Llavallol, Sta. Catalina, fruit bodies obtained from strain BAFC 120, leg. B. Lechner & E. Albertó, IV-1999, BAFC , BAFC ; fruit bodies obtained from strain BAFC 2034, 20-III-1999, leg. B. Lechner, BAFC ; fruit bodies obtained from strain MUCL (P. columbinus), leg. B. Lechner & E. Albertó, IV-2000, BEL 64; fruit bodies obtained from strain BAFC 2473 (CBS ), leg. B.E. Lechner, X- 2000, BAFC and ; fruit bodies obtained from strain BAFC 2478 (CBS ), leg. B.E. Lechner, 13-X- 2000, BAFC , and 27-VI-2000 (BAFC ); Ezeiza, 12-III-1972, leg. J. Deschamps, BAFC 1.625; La Plata, obtained from culture, leg. H. Spinedi, 24-VII-1992, LPS No ; Parque Pereyra, on trunk of Populus sp., leg. Singer- Horak-Wright, 20-VI-1962, BAFC ; CATAMARCA, Santa María, on living and dead trunk of Salix sp., T (LIL); NEUQUÉN, Aluminé, Moquehue, on trunk of Araucaria araucana, leg. Jorge del Vas, III-1993 CIEFAP (BAFC ); Lanín National Park, growing on A. araucana, leg. M. Rajchenberg, 20-V-1999, CIEFAP ; ibid., growing on A. araucana, leg. M. Rajchenberg, VI-1999, CIE- FAP ; TUCUMÁN, Estancia Las Pavas, 19-VI-49, leg. Cei, T 573 (LIL); Tapia, on dry trunk, 23-III-1948, leg. Goldbach (LIL); tropical wood, on trunk, 27-I-1951, leg. R. Singer, T (LIL). Remarks. Fruit bodies of different collections showed great variability, characterized by sclerified hyphae or thin-walled hyphae in the stem and hymenophoral trama and by the pileipellis being of variable thickness. This indicates the necessity to make further studies to separate Pleurotus ostreatus and P. pulmonarius, such as mating (Vilgalys et al 1993, Lechner et al 2002) or molecular studies (Thorn et al 2000, Gonzales and Labarère 2000). Pleurotus pulmonarius (Fr.) Quél. Champignons du Jura et des Vosges I; 113, pl. 4. FIG Agaricus pulmonarius Fr. : Fr Syst. Mycol. 1:187. P. pulmonarius (Fr. : Fr.) Quél Mém. Soc. Émull. Montbéliard 2, sér. 5:11. P. ostreatus var. pulmonarius (Fr. : Fr.) Pilát Bull. Trimest. Soc. Myc. Fr. 49:281. P. araucariicola Sing Lilloa 26:141. Pileus mm, flabelliform (FIGS. 9A, 11K, L), whitish to grayish (Munsell, HUE 7/1 7/2 5Y), very pale brown (Munsell, HUE 8/4 10YR) when dry, surface smooth, glabrous, shiny, gelatinous. Lamellae decurrent, crowded, 2 3 mm broad, cream,

11 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 855 FIG. 10. Pleurotus rickii. A. Basidiocarps; B. Spores; C. Basidia; D. Hyphae of the hymenophoral trama; E. Hyphae of the pileus; F. Hyphae of the stem. Scale bar 1 4cm for A; scale bar 2 20 m for B F. yellow (Munsell, 10YR 8/6) when dry, margin entire. Stipe eccentric to lateral, smooth, 2 10 mm long 2 6 mm diam. Flesh 2 5 mm thick, less than 1 mm when dry, corky. Odor agreeable, aromatic, not aniselike. Spores (6.8 ) ( 10.9) m Q 2.8, cylindrical-oblong, hyaline, thin-walled, IKI- (FIG. 9B). Basidia m, club shaped, 4- spored (FIG. 9D). Cheilocystidia absent. Numerous basidioles or pleurocystidia, m (FIG. 9D). Hymenophoral trama completely irregular, with clamped, sclerified hyphae, m diam and thin-walled hyphae m diam; oleiferous hyphae not observed (FIG. 9D). Context of pileus with generative hyphae m diam, thin- and thickwalled (FIG. 9C). Hyphae of stem generative, thickwalled m diam (FIG. 9E). Pileipellis a cutis, m thick, not well differentiated, with hyphae parallel and radial arranged. Culture characters. Mycelium hyaline, whitish and arachnoid near inoculum. Later more white, cottony, with increasing aerial mycelium. Radial growth was generally observed; sometimes feathery. Margin irregular. Reverse of agar unchanged. Odor of bitter almonds. Mycelium covered Petri plates within 1 2 wk. Advancing mycelium with clamped, thin-walled hyphae m diam, more branched in the center of the colony, with hyphae m diam and near inoculum m diam. Microdroplets m diam. Geographic distribution. Northeastern and east central Argentina. Material studied. ARGENTINA, BUENOS AIRES, Ezeiza, on live but diseased tree of Populus nigra, 25-VII-1987, leg. J. Deschamps, BAFC ; CAPITAL FEDERAL, Barrio Palermo, on Salix humboldtiana, VI and VII-1881, leg. C. Spegazzini, LPS ; MISIONES, San Pedro, on branches of Araucaria angustifolia, 27-V-2001, leg. E. Albertó & O. Popoff, BAFC ; San Pedro, S, W, on branches of Araucaria angustifolia, 27-V-2001, leg. D. Krueger, BAFC ; same location, same host, same date, leg. E. Albertó, BAFC ; BRAZIL, Serra, Taimbesino, 5-XI- 1951, leg. R. Singer B 89 (BAFC ), HOLOTY PE of P. araucariicola Sing. Remarks. Differentiation between P. pulmonarius and P. ostreatus is difficult. As we pointed out above, mating and molecular studies are necessary to separate them, however, the pileipellis in P. pulmonarius is thinner (20 40 m), resulting in a more light-colored basidiocarp, whereas P. ostreatus has a thicker pileipellis ( m) and darker color. According to Hilber (1982) they may be distinguished by an anise odor in P. pulmonarius, but we could not perceive this feature. Instead, we could distinguish a sweet aromatic odor, different to the fungal one of P. ostreatus. Based on observations of fruit bodies obtained from culture, we conclude that P. ostreatus specimens were stouter, with a more eccentric stem, against a lateral, rarely eccentric and less developed stem of the cultured P. pulmonarius specimens studied. Regarding P. araucariicola, morphological studies of the holotype, as well as mating studies, revealed that it is clearly a synonym of P. pulmonarius (Lechner et al 2003). Pleurotus rickii Bresadola Ann. Mycologici 18, p Description of the collections in dry state. Pileus mm, brown to grayish brown, glabrous, smooth, rather infundibuliform (FIG. 10A). Lamellae clear brown, mm broad, deeply decurrent. Stipe cream, mm long and 2 6 mm diam, slightly sulcate, solid, eccentric to subcentral, with dark strands formed by skeletal hyphae arranged along it. Spores (8.8 ) ( 12.4) 2.7 (3.3) 3.6 m,

12 856 MYCOLOGIA FIG. 11. Basidiomata of different species of Pleurotus from Argentina. A. P. albidus from nature and B. from culture; C. P. cystidiosus; D. coremia from P. cystidiosus; E G. P. djamor from nature. E. var. djamor, F. var. roseus, G. var. cyathiformis; H J. P. ostreatus, H. from nature, I and J. from culture; K L. P. pulmonarius, K. from nature, L. from culture. Scale bar 30 cm for A, 8 cm for B, 4 cm for C and E, 6 mm for D, 5.5 cm for F G, 8 cm for H, 14 cm for I, 16 cm for J L.

13 LECHNER ET AL: THE GENUS PLEUROTUS IN ARGENTINA 857 Q 3.18, cylindrical to cylindrical-oblong, hyaline, thin-walled, neither amyloid nor dextrinoid (FIG. 10B). Basidia m (FIG. 10C). Hymenophoral trama irregular, with generative hyphae m diam (FIG. 10D). Pileus with generative hyphae, m diam, and skeletals m diam (FIG. 10E). Stem with generative hyphae, thinwalled, m diam and pointed skeletals m long and m diam, with a thick wall m thick (FIG. 10F). On the surface of the stem, dark strands were observed formed by skeletals hyphae m diam, with a thick wall, up to 2.5 m thick (FIG. 10F). Geographic distribution. Northwestern Argentina. Material studied. ARGENTINA, TUCUMÁN, Parque Aconquija, on leafy trunk in tropical wood, 14-IV-1951, leg. R. Singer, T 1472 (LIL); Tafí, San Javier, 6-V-1945, leg. H. Descale, T 2221 (LIL). Remarks. Singer (1951) described the species in the fresh state. The pileus was recorded as brown ( chukker brown, Maerz and Paul P15C8) mixed with pale tones with small close squamules ( cowboy, Maerz and Paul P15E11) and marginal, white, early disappearing veil; the rest smooth and glabrous; lamellae were described as white, yellow through drying; stipe soapstone color (Maerz and Paul P15A7), with white veil, soft, very notable when young. Spore print opal mauve (Maerz and Paul P51A2). It will be important to obtain cultures of this species, so far nonexistent, to observe the presence or absence of microdroplets to confirm its place in Pleurotus because the specimens, only morphologically studied, appear to be on the boundary between Pleurotus and Lentinus. ACKNOWLEDGMENTS This research was supported by PRHIDEB-CONICET/Argentina. We wish to thank the curators of BAFC, LIL and LPS herbaria for making available collections of species of Pleurotus for this study, as well as the curators of BAFC, CBS and MUCL culture collections for making available strains. We also thank Mario Rajchenberg (Esquel, Argentina), who provided the collection and strain of P. ostreatus from Patagonia, Argentina, and Dr R.H. Petersen (Knoxville, Tennessee) for supplying valuable information on Pleurotus species. This is paper No. 158 of the PRHIDEB-CONICET series. LITERATURE CITED Albertó E, Petersen RH, Hughes KW, Lechner BE Miscellaneous notes on Pleurotus. Persoonia 18(1): Barron GL, Dierkes Y Nematophagous fungi: Hohenbuehelia, the perfect state of Nematoctonus. Can J Bot 55: , Thorn RG Destruction of nematodes by species of Pleurotus. Can J Bot 64: Corner EJH The agaric genera Lentinus, Panus, and Pleurotus with particular reference to Malaysian species. Beih Nova Hedw 69: Gonzales P, Labarère J Phylogenetic relationships of Pleurotus species according to the sequence and secondary structure of the mitochondrial small-subunit rrna V4, V6 and V9 domains. Microbiology 146: Grand EA, Petersen, RH, Hughes KW Biogeography and species concepts Lentinus lindquistii is Lentinus tigrinus, a circum global species. Inoculum 53(3):30. Hibbett DS, Thorn RG Nematode trapping in Pleurotus tuberregium. Mycologia 86: , Donoghue MJ Progress toward a phylogenetic classification of the Polyporaceae through parsimony analysis of mitochondrial ribosomal DNA sequences. Can J Bot 73(Suppl 1):S853 S860., Vilgalys R Phylogenetic relationships of the Basidiomycete genus Lentinus inferred from molecular and morphological characters. Syst Bot 18: Hilber O Die Gattung Pleurotus. Biblioth Mycol 87: Holmgren PK, Holmgren NH, Barnett LC Index Herbariorum. New York Botanical Garden. New York, 693 p. Kirk PM, Ansell AE Authors of fungal names. Index of Fungi, Suppl., 95 p. Kühner R Les Hyménomycètes agaricoïdes, étude genérale et classifications. Numéro special. Bull Soc Linn Lyon 49: , Romagnesi H Flore analytique des champignons supérieurs. Masson et Cie (Eds.). Paris. 554 p. Lechner BE, Albertó E, Petersen, R Pleurotus araucariicola is Pleurotus pulmonarius. Mycotaxon 86: , Petersen R, Rajchenberg M, Albertó E Presence of Pleurotus ostreatus in Patagonia, Argentina. Rev Iberoam Micol 19: Maerz A, Paul M Dictionary of color. New York: Mc- Graw-Hill Book Company, Inc. 207 p. Moser, M Guida alla determinazione dei funghi. Vol. 1. Trento: Arti Grafiche Saturnia. 565 p. Munsell, Color Co. Inc Determination of soil color. Baltimore: U.S. Dept. Agriculture Handbook. 16 p., 7 pl. Nobles MK Studies in forest pathology VI. Identification of cultures of wood-rotting fungi. Can J Res 26: Identification of cultures of wood-inhabiting Hymenomycetes. Can J Bot 43: Pegler DN The classification of the genus Lentinus Fr. (Basidiomycota). Kavaka 3: The genus Lentinus. Kew Bull Add Ser 10: Pilát A Pleurotus Fries. In: Kavina C, Pilát A, eds. Atlas

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