e ALBERT K. MANN LIDKARY AT CORNELL UNIVERSITY CORNELL UNIVERSITY LIBRARY wii 1924 090 172 3 4 ay o i at ‘A , ee ie 7 of ne a ~ \y FN eg aa ee 8 Se TR RR EF ead en aay ao A Slane st RSE NY ot GED ae Le ree et iy ae dF ag OE RL ee iS alas ut Soe he Cues 3 # i he on) a ‘ey — Seer nes 4%. AN INTERNATIONAL JOURNAL FOR RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS Volume LXXXI January-March 2002 What is Cytiaria disciformis? Irma J. Gamundi & Cecilia A. M. Brion ] A new species of Aphanoascus (Ascomycota) with a Malbranchea anamorph from marine bottom deposits Michael V. Pivkin & Yulia V. Khudyakova 7 Claviceps purpurea vat. spartinae var. nov.: an ergot adapted to the aquatic environment Russell A. Duncan, Jr., Raymond Sullivan, Stephen C, Alderman, Joseph W. Spatafora, & James F. White, Jr. 11 Some new species and new records of discomycetes in China. IX Wen- Ying Zhuang, Zhi-He Yu, Yan-Hui Zhang, & Ming Ye 27 Re-examinations of Helotium and Hymenoscyphus (Helotiales, Helotiaceae): specimens on deposit in HMAS Yan-Hui Zhang & Wen-Ying Zhuang 35 Two new dermateaceous genera with lanceolate paraphyses Ain Raitviir 45 The Pleurotus eryngii species-complex in Israel: distribution and morphological description of a new taxon D. Lewinsohn, S. P. Wasser, S. V. Reshetnikov, Y. Hadar, & E.Nevo 51 Pleurotus eryngii var. thapsiae var. nov. from Sicily G. Venturella, G. Zervakis, &A. Saitta 69 Materials for a hypogeous mycoflora of the Great Basin and adjacent cordilleras of the western United States. VII: A new truffle-like Peziza (Ascomycota, Pezizales) Robert Fogel & Jack States 75 Materials for a hypogeous mycoflora of the Great Basin and adjacent cordilleras of the western United States. VII: Pachyphloeus lateritius sp. nov. and Cazia quericola sp. nov. (Ascomycota, Pezizales) Robert Fogel & Jack States 83 A new Inonoius (Basidiomycota, Hymenochaetaceae) from Hawat'1 Robert L. Gilbertson & Leif Ryvarden 91 Phaeocollybia in western North America 3: Two new species with notes on the Phaeocollybia festiva complex Lorelei L. Norvell 95 Preliminary checklist of Agaricales from Croatia. I. Families Pleurotaceae and Tricholomataceae Zdenko Tkaléec & Armin MeSi¢é 113 Thaxterogaster is a taxonomic synonym of Cortinarius: new combinations and new names | Ursula Peintner, Meinhard Moser, & Rytas Vilgalys 177 Tremella spicifera sp. nov., a parasite of Massarina arundinacea G. Van Ryckegem, K. Van de Put, & P. Roberts 185 The mating system in two species of Crepidotus M. Catherine Aime & Jessica Ball 191 {Content continues inside front cover] ISSN 0093-4666 MYXNAE 81:1-540 (2002) For subscription details, availability in microform, and availability of articles as photocopies or tear sheets, see back cover. [Contents continued from front cover] Nomenclatural revisions in the sequestrate russuloid genera James M. Trappe, Teresa Lebel, & Michael A. Castellano Corticioid and polyporoid fungi (Basidiomycota) that decay Austrocedrus chilensis in Patagonia, Argentina Mario Rajchenberg Molecular identification of the teleomorph of Beauveria bassiana Bo Huang, Chun-ru Li, Zhen-gang Li, Mei-zhen Fan, & Zeng-zhi Li Plectania carranzae sp. nov. (Ascomycotina), from Costa Rica F. D. Calonge & M. Mata A natural mutant of Ascobolus michaudii (Ascomycotina, Pezizales) Diana A. Dokmetzian & Maria E. Ranalli A new species and further new records in the lichen family Parmeliaceae (Ascomycotina) from the Philippines John A. Elix, Paulina A. Bawingan, & Yolanda G. Flores The lichen family Parmeliaceae (Ascomycotina) on Gough Island, South Atlantic Ocean John A. Elix & Niek J. M. Gremmen Lichenicolous fungi on Jonaspis lacustris Alan Orange Glomus lamellosum (Glomales, Zygomycotina), an arbuscular mycorrhizal fungal species new for Poland and Europe Janusz Blaszkowski, Mariusz Tadych, Iwona Adamska, & Tadeusz Madej Notes on Hyphomycetes. LXXXV. Junewangia, a genus in which to classify four Acrodictys species and a new taxon William A. Baker, E. Christopher Partridge, & Gareth Morgan-Jones Phylogeny of Amphisphaeriaceae (sensu stricto) and related taxa revisited based on nrDNA sequences Ji-chuan Kang, Richard Y. C. Kong, & Kevin D. Hyde Morphological responses of Canadian and U.S.A. isolates of Phoma sclerotioides to different growth media, temperatures and light C. R. Hollingsworth, F. A. Gray, R. W. Groose, & C. W. Mims A new species of Phacellium on Alangium Hyeon-Dong Shin & Jeong-Dong Kim Corticolous myxomycetes associated with four tree species in Mexico Mercedes Rodriguez-Palma, Adalid Varela Garcia, & Carlos Lado Mycological characterization of an Hapludand soil series under three management practices Eduardo Valenzuela F., Dante Pinochet T., & Perla Carias M. Taxonomical studies on Ustilaginales. XXII Kalman Vanky Two new species of Urocystis and a Urocystis (Ustilaginomycetes) new to China Lin Guo Puccinia ariorum sp. nov. and other noteworthy graminicolous rust fungi (Uredinales) from Iran M. Abbasi, Gh. A. Hedjaroude, H. B. Gjaerum, & M. Scholler Conidial aquatic fungi of NainiTal, Kumaun Himilaya, India S. C. Sati, N. Tiwari, & M. Belwal Aquatic hyphomycetes from cerrado regions in the State of Sad Paulo, Brazil Iracema Helena Schoenlein-Crusius Muscodor roseus anam. sp. nov., an endophyte from Grevillea pteridifolia Jeerapun Worapong, Gary Strobel, Bryn Daisy, Uvidelio F. Castillo, Gary Baird, & W. M. Hess Mycoaciella, a synonym of Phlebia Karen K. Nakasone Index of fungi described from the Musaceae W. Photita, K. D. Hyde, E. H. C. McKenzie, & S. Lumyong [Content continues inside back cover] 195 215 229 PRY) 243 24 27 265 281 205 S21 ae 341 345 2131) 367 431 435 445 457 463 477 49] MYCOTAXON AN INTERNATIONAL JOURNAL OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS VOLUME LXXXI, 2002 COMPLETE IN ONE VOLUME CONSISTING OF iv + 540 PAGES INCLUDING FIGURES EDITOR-IN-CHIEF Pavel Lizon Department of Cryptogams, Institute of Botany Dubravska 14 SK-842 23 Bratislava, Slovakia ASSOCIATE EDITORS BOOK REVIEW EDITOR David L. Hawksworth MycoNova, Calle Aguila 12 Colonia La Maliciosa, Mataelpino ES-28411 Madrid, Spain FRENCH LANGUAGE EDITOR Grégoire L. Hennebert 32 Rue de l'Elevage B-1340 Ottignies - LLN, Belgium INDEX EDITOR Karen D. Gettelman 7044 Amherst Avenue St. Louis, MO 63130, USA EDITORIAL ADVISORY BOARD CAROL H. SHEARER, Urbana, Illinois, USA (1998-2003) Chair GREGORY M. MUELLER, Chicago, Illinois, USA (1998-2002) Past Chair RANDOLPH 8S. CURRAH, Edmonton, Alberta, Canada (1999-2004) SEPPO HUHTINEN, Turku, Finland (2000-2005) GARY J. SAMUELS, Beltsville, Maryland, USA (1997-2006) DONALD H. PFISTER, Cambridge, Massachusetts, USA (1997-2007) Published by MYCOTAXON, LTD., P. O. BOX 264 ITHACA, NY 14851-0264, USA www.mycotaxon.com Printed in the United States of America © Mycotaxon, Ltd., 2002 TABLE OF CONTENTS, VOLUME EIGHTY-ONE What is Cyttaria disciformis? Irma J. Gamundi & Cecilia A. M. Brion A new species of Aphanoascus (Ascomycota) with a Malbranchea anamorph from marine bottom deposits Michael V. Pivkin & Yulia V. Khudyakova Claviceps purpurea vat. spartinae var. nov.: an ergot adapted to the aquatic environment Russell A. Duncan, Jr., Raymond Sullivan, Stephen C. Alderman, Joseph W. Spatafora, & James F. White, Jr. Some new species and new records of discomycetes in China. IX Wen-Ying Zhuang, Zhi-He Yu, Yan-Hui Zhang, & Ming Ye Re-examinations of He/otium and Hymenoscyphus (Helotiales, Helotiaceae): specimens on deposit in HMAS Yan-Hui Zhang & Wen-Ying Zhuang Two new dermateaceous genera with lanceolate paraphyses Ain Raitviir The Pleurotus eryngii species-complex in Israel: distribution and morphological description of a new taxon D. Lewinsohn, S. P. Wasser, S. V. Reshetnikov, Y. Hadar, & E. Nevo Pleurotus eryngii var. thapsiae var. nov. from Sicily G. Venturella, G. Zervakis, & A. Saitta Materials for a hypogeous mycoflora of the Great Basin and adjacent cordilleras of the western United States. VII: A new truffle-like Peziza (Ascomycota, Pezizales) Robert Fogel & Jack States Materials for a hypogeous mycoflora of the Great Basin and adjacent cordilleras of the western United States. VIII: Pachyphloeus lateritius sp. nov. and Cazia quericola sp. nov. (Ascomycota, Pezizales) Robert Fogel & Jack States A new Jnonotus (Basidiomycota, Hymenochaetaceae) from Hawai'i Robert L. Gilbertson & Leif Ryvarden Phaeocollybia in western North America 3: Two new species with notes on the Phaeocollybia festiva complex Lorelei L. Norvell Preliminary checklist of Agaricales from Croatia. I. Families Pleurotaceae and Tricholomataceae Zdenko Tkaléec & Armin Mesi¢ Thaxterogaster is a taxonomic synonym of Cortinarius: new combinations and new names Ursula Peintner, Meinhard Moser, & Rytas Vilgalys Tremella spicifera sp. nov., a parasite of Massarina arundinacea G. Van Ryckegem, K. Van de Put, & P. Roberts The mating system in two species of Crepidotus M. Catherine Aime & Jessica Ball Nomenclatural revisions in the sequestrate russuloid genera James M. Trappe, Teresa Lebel, & Michael A. Castellano Corticioid and polyporoid fungi (Basidiomycota) that decay Austrocedrus chilensis in Patagonia, Argentina Mario Rajchenberg Molecular identification of the teleomorph of Beauveria bassiana Bo Huang, Chun-ru Li, Zhen-gang Li, Mei-zhen Fan, & Zeng-zhi Li Plectania carranzae sp. nov. (Ascomycotina), from Costa Rica F. D. Calonge & M. Mata A natural mutant of Ascobolus michaudii (Ascomycotina, Pezizales) Diana A. Dokmetzian & Maria E. Ranalli A new species and further new records in the lichen family Parmeliaceae (Ascomycotina) from the Philippines John A. Elix, Paulina A. Bawingan, & Yolanda G. Flores ill 69 75 83 | oD 13 Li 185 19] LS OAS 229 PEM | 243 oll iv The lichen family Parmeliaceae (Ascomycotina) on Gough Island, South Atlantic Ocean John A. Elix & Niek J. M. Gremmen Lichenicolous fungi on Jonaspis lacustris Alan Orange Glomus lamellosum (Glomales, Zygomycotina), an arbuscular mycorrhizal fungal species new for Poland and Europe Janusz Blaszkowski, Mariusz Tadych, Iwona Adamska, & Tadeusz Made} Notes on Hyphomycetes. LXXXV. Junewangia, a genus in which to classify four Acrodictys species and a new taxon William A. Baker, E. Christopher Partridge, & Gareth Morgan-Jones Phylogeny of Amphisphaeriaceae (sensu stricto) and related taxa revisited based onnrDNA sequences Ji-chuan Kang, Richard Y. C. Kong, & Kevin D. Hyde Morphological responses of Canadian and U.S.A. isolates of Phoma sclerotioides to different growth media, temperatures and light C. R. Hollingsworth, F. A. Gray, R. W. Groose, & C. W. Mims A new species of Phacellium on Alangium Hyeon-Dong Shin & Jeong-Dong Kim Corticolous myxomycetes associated with four tree species in Mexico Mercedes Rodriguez-Palma, Adalid Varela Garcia, & Carlos Lado Mycological characterization of an Hapludand soil series under three management practices Eduardo Valenzuela F., Dante Pinochet T., & Perla Carias M. Taxonomical studies on Ustilaginales. XXII Kalman Vanky Two new species of Urocystis and a Urocystis (Ustilaginomycetes) new to China Lin Guo Puccinia ariorum sp. nov. and other noteworthy graminicolous rust fungi (Uredinales) from Iran M. Abbasi, Gh. A. Hedjaroude, H. B. Gjaerum, & M. Scholler Conidial aquatic fungi of NainiTal, Kumaun Himilaya, India S. C. Sati, N. Tiwari, & M. Belwal Aquatic hyphomycetes from cerrado regions in the State of Sa6 Paulo, Brazil Iracema Helena Schoenlein-Crusius Muscodor roseus anam. sp. nov., an endophyte from Grevillea pteridifolia Jeerapun Worapong, Gary Strobel, Bryn Daisy, Uvidelio F. Castillo, Gary Baird, & W. M. Hess Mycoaciella, a synonym of Phlebia Karen K. Nakasone Index of fungi described from the Musaceae W. Photita, K. D. Hyde, E. H. C. McKenzie, & S. Lumyong Nomenclatural novelties proposed in volume 81 Author index Index to fungous and lichen taxa Errata Reviewers Publication date for volume 80 257 265 281 293 y32)1 331 341 345 B57 367 431 435 445 457 463 477 491 505 509 53 538 539 eee -MYCOTAXON Volume LXXXI, pp. 1-6 _ January-March 2002 WHAT IS CYTTARIA DISCIFORMIS? Irma J. Gamundi (*) & Cecilia A. M. Brion (**> Consejo Nacional de Investigaciones Cientificas y Técnicas, Argentina and Centro Regional Universitario Bariloche, Universidad Nacional del Comahue, Quintral 1250, 8400 S. C. de Bariloche, Rio Negro, Argentina. E-mail: igamundi@crub.uncoma.edu.ar ** Centro Regional Universitario Bariloche, Universidad Nacional del Comahue, Argentina. E-mail: cbrion@crub.uncoma.edu.ar Keywords: Ascomycota, Basidiomycota, Dacrymycetaceae Abstract After examination of the type specimen of Cyttaria disciformis Lév. it was concluded that the species is not an Ascomycota but a Basidiomycota belonging to the order Dacrymycetales, family Dacrymycetaceae. Introduction Léveillé (1846:254) described Cyttaria disciformis under species number 283 as: “Receptaculo orbiculari discoideo emortuarum, tremelloideo flavo stipite brevissimo suffulto, foveolis minutissimis sparsis. Hab.: Chili ad cortices arborum. Cl. Gay, (herb. Mus. Paris)”. Under observations it is said that :“‘le réceptacle de cette petite éspece est orbiculaire, convexe, large de 3 a 6 millimétres, supporté par un court pédicule, qui lui donne quelque ressemblance avec un bouton; la face supérieure présente de trés petites fossettes eloignées les unes des autres dans lequelles je nai vu que des longs filaments libres, continus et rameux, avec une couche de cellules allongées, courtes, trés pressées et terminées par un renflement presque globuleux renferment un corps opaque et irrégulier. Sous l’influence de la teinture d’iode, les filaments ainsi que les cellules se colorent en jaune brun, tandis que le reste du tissu, composeé des petites cellules polygones, conserve sa couleur blanche”. Fischer (1888:815-842) who revised the type, stated that the fungus in question does not shows identifiable apothecia or primordia of apothecia and that it could be any fungus but Cyttaria. Perhaps for that reason, several mycologists who treated taxonomically the genus (Espinosa, 1926, Marchionatto, 1940, Santesson, 1945, Rawlings, 1956, Kobayasi, 1964, 1966, Butin, 1969) excluded C. disciformis Lév. from their papers. This fact drew our attention to the name and lead us to investigate the type specimen. Materials and Methods For microscopical observations pieces of fruitbodies were soaked in distilled water and 5% KOH. They were sectioned and squashed. As mounting media Melzer’s and lactic blue were used. Results During the stay at the Paris Museum in October 1964, one of the autors (1. J. G.) examined the putative type specimen of Cyttaria disciformis Lév. The label read: “Herbarium Mycologicum Berkeleyanum No. 7937, Chili, C. Gay leg., ad truncus”. The exssicata contained one fructification of 6 mm diam., characterised by a cartilaginous consistency, cinnamon reddish colour, with an acute, undulate margin. When rehydrated it was gelatinous. When microscopically examined, its upper surface showed a palisade of clavate cells arising from clamps and below a hyphose structure formed by thin, profusely branched hyphae. No spores were seen. The clavate structures were interpreted as basidioles. The conclusion was that the fungus examined, although immature, was a basidiomycete. These observations confirmed Fisher’s view and, for that reason, C. disciformis Lév. was not included in papers by Gamundi (1971, 1987) and Crisci, Gamundi & Cabello (1988) on Cyttariales. Recently, we examined the type specimen kept in PC. The label read: “Cyttaria disciformnis Lév., Chili, Gay” (Fig. 1). It was impossible to locate the collecting site, but the wood and bark of the host material seems to be of a dicotyledon. Description Macroscopic features The type specimen consists of six pieces of host tissue, which we numbered from 1 to 6 (Fig.l, numbers 1-6). Number 1 shows three discoid fructifications (Fig. 7,1); number 2, two cylindrical fruitbodies (Fig.7.2); number 3, one discoid, substipitate fructification (Fig.7.3); number 4, no fructification (Fig. 7.4); number 5, one discoid— turbinate fruitbody (Fig. 7.5); number 6, one flabelliform, stipitate fruitbody (Fig. 7.6). Macroscopic features common to all of them is the firm, highly cartilaginous, translucent, caramel coloured carpophore when dried, and gelatinous, yellowish fruitbody when soaked in water, arising from the wood. The discoid-turbinate fructifications (numbers 1,3,5, figs. 9-10) are 2-5 mm diameter, the cylindrical ones (number 2, fig. 8) 3-4 mm long x 0.8-1.2 mm diam.; flabelliform (number 6, fig. 11) 7.5 mm long x 4 mm diam.(upper portion), all measures taken on dried material. Microscopic features In section the upper surface of the discoid-turbinate basidiomata (numbers 3-5) comprises the palisade layer of the hymenium (Figs. 2-4); the lower and lateral surfaces comprise an ill-differentiated layer of more o less parallel hyphae perpendicular to the surface; the inner portion is a mass of intricate, branched hyphae (Fig. 5), all the structures are embedded in a highly gelatinous matrix. In the flabelliform fructification (number 6) the hymeniun was unilateral and supported by a stipe. The cylindrical fruitbodies (number 2) have the upper end enlarged and convex. As no hymenium was detected, we presume they are immature. MUSEUM NATIONAL D'HISTOIRE NATURELLE (PARIS) HERBIER CRYPTOGAMIQUE Ve “* Luft d o [fain den stl coma te 1 | os Figs. 1-6. Cyttaria disciformis Lév. Fig. 1. Type specimen, pieces numbered 1-6. Fig. 2. Hymenium showing basidioles and probasidium (arrows). Fig. 3. Hymenium showing bifurcate basidium (arrow). Fig. 4. Hymenium showing septate basidium (arrow). Fig. 5. Cortex (upper arrow) and context (lower arrow). Fig. 6. Context showing clamp connections (arrows). Scale bars: Fig. 1 = 1 em; Figs. 2-5 = 10 um; Fig. 6 = 20 um. 4 Hymenium superior or lateral, with clavate basidioles 27-34 x 2-3 um diam. at the widest part, cytoplasm intensely stained with lactic blue, dextrinoid (Fig. 2); probasidia cylindrical. then bifurcate (Fig. 3), produced in clusters, rare metabasidia apically bifurcate forming two long sterigmata and a transverse septum, arising from a clamp, 27-45 x 3.4 4.8 um ( Fig. 4); protoplasm granulose to guttulate, cytoplasm staining deep with lactic blue; dikaryophyses cylindrical, rare. Cortex an ill-defined palisade of hyphae embedded in a gelatinous matnx, ending irregularly, cytoplasm granulose, deeply stained with lactic blue, dextrinoid (sample numbers 1, 3). In sample number 6 the ends are wider (up to 2.4 pm), fusiform, clavate to trapezoidal, with incrusted walls and light brown cytoplasm. (Fig. 5 ). Context a mass of hyphae profusely branched (2-3-4 divaricate) embedded in a gelatinous matrix, with clamp connections either simple or forming a loop, (Fig. 6) hyaline, thick walled (walls as thick as lumen), intermixed with brownish, wider hyphae with incrusted walls. (Fig. 5). Discussion Léveillé*s description is in agreement with our observation regarding the fruitbody, except that we have not seen small cavities in any of the specimens examined, a feature that perhaps led Léveillé to classify the collection as a Cyttaria. The filaments and elongated cells which lined “the cavities” described by Léveillé are probably a young hymenium , but no reference was made to asci. Léveillé made no reference to host substrate. This would have provided another clue to the identity of the genus, as Cyttaria 1s parasitic only on Nothofagus. There is enough evidence from our observations of microscopic features (clamp connections, bifurcate metabasidia) that Cyttaria disciformis Lév. is a Basidiomycota, Dacrymycetales, Dacrymycetaceae (Hawksworth et al., 1995) and not an Ascomycota, Cyttanales, Cyttariaceae. The bifurcate basidia are characteristic of Dacrymycetaceae (Kennedy, 1958, Martin, 1952) formerly placed in the Tremellales, but later (McNabb & Talbot, 1973, Reid, 1974, Hawksworth et al., 1995) in the Dacrymycetales. Unfortunately, we could not see spores, but the structures of the basidioma suggest one of two genera, owing to the upper or lateral position of the hymenium and differentiation of a cortex and a context: Dacrymyces Fr. (McNabb, 1973) or Femsjonia Fr. (McNabb, 1965). Acknowledgements . We thank Dr. Jean Mouchacca (Museum d’Histoire Naturelle, Paris, Cryptogamie) for kindly attending our request of the type specimen; Dr. Gernot Vobis, for his aid on photomicrography and Dr. Javier Puntieri for scanning the type specimen. Fig. 9. Fruitbody of number 3. Fig. 10. Fruitbody of number 5. Fig. 11. Fruitbody of number 6. Scale bars: Fig. 7 = 1em; Figs. 8-11 = 2 mm. References Butin, H. 1969. Uber die pilzgattung Cyttaria Berk. in Chile. Z. Pilzk. 35: 85-91. Crisci, J. V., Gamundi, I. J. & Cabello, M. N. 1988. A cladistic analysis of the Genus Cyttaria Fungi, Ascomycotina). Cladistics 4: 279-290. Espinosa, M. B. 1926. Los hongos chilenos del Género Cyttana Berk. Reista. Chilena Hist. Nat. 30: 206-256. Fischer, E. 1888. Zur Kenntniss der Pilzgattung Cyttaria. Bot. Zeitung 46 (51): 813- 832; 842-847; Taf. XII. Gamundi, I. J. 1971. Las Cytariales sudamencanas. Darwiniana 16(2/3): 461-510. Gamundi, I. J. 1987 [1986’]. Fungi, Ascomycetes. Cyttanales, Helotiales: Geoglossaceae, Dermateaceae. In Flora Criptogamica de Tierra del Fuego, 10, fasc. 4. (S. A. Guarrera, I. Gamundi de Amos. & D. Rabinovich de Halperin, eds.): 1-126. CONICET, Buenos Aires. Hawksworth, D. L., Kirk, P. M., Sutton, B. C. & Pegler, D. N. 1995. Ainsworth & Bisby’s Dictionary of the Fungi. 8" ed. International Mycological Institute, CABI, Egham. Kennedy, L. L. 1958. The genera of Dacrymycetaceae. Mycologia 50(6): 874-915. Kobayasi, Y. 1964. On the Genus Cyttaria (1). J. Jap. Bot. 29: 216-224. Kobayasi, Y. 1966. On the genus Cyttaria (2) .Trans. Mycol. Soc. Japan 7 (2/3): 118- 132; Leveillé, J. H. 1846. Description des Champignons de I’herbier du Museum de Paris. Ann. Sci. Nat. Sér. Ill, 5: 111-167; 249-304. 6 Marchionatto, J. B. 1940. Las especies de Cyttaria y Cyttanella en la Argentina. Darwiniana 4: 9-32. Martin, G. W. 1952. Revision of North Central Tremellales. State Univ. Iowa Stud. Nat. Hist. No. 423, 19 (3): 1-122. | McNabb, R. F. R. 1965. Taxonomic studies in the Dacrymycetaceae. VI. Femsjonia. New Zealand J. Bot. 3: 223-228. McNabb, R. F. R. 1973. Taxonomic studies in the Dacrymycetaceae. VII. Dacrymyces Nees ex Fries. New Zealand J. Bot. 11: 461-524. McNabb, R. F. R. & Talbot, P. H. B. 1973. Holobasidiomycetidae: Exobasidiales, Brachybasidiales, Dacrymycetales. In The Fungi, An Advanced Treatise 4B, Chapter 18 (G. C. Ainsworth, F. K. Sparrow & A. S. Sussman, eds.): 317-325. Reid, D. A. 1974. A monograph of the British Dacrymycetales. Trans. Brit. Mycol. Soc. 62(3): 433-494. | Rawlings, G. B. 1956. Australasian Cyttariaceae. Trans. Roy. Soc. New Zealand 84: 19-28. Santesson, R. 1945. Cyttaria, a genus of inoperculate Discomycetes. Svensk Bot. Tidskr. 39: 319-345. See Errata MYCOTAXON Volume LXXXI, pp. 7-10 January-March 2002 A NEW SPECIES OF APHANOASCUS (ASCOMYCOTA) WITH A MALBRANCHEA ANAMORPH FROM MARINE BOTTOM DEPOSITS Michael V. Pivkin, Yulia V. Khudyakova Pacific Institute of Bioorganic Chemistry, Far East Branch of the Russian Academy of Sciences, 690022, Vladivostok-22, Russia; e-mail: piboc@eastnet.febras.ru ABSLRAGT Aphanoascus aciculatus sp. nov. from marine bottom deposits of the Sea of Japan 1s described. Key words: Marine fungi, taxonomy. As part of a programme investigating fungi from marine environments, a collection of hyphomycetes from marine bottom deposits was analysed. The growing of these fungi under various conditions has allowed us to reveal several strains, which form holomorphs of the genus Aphanoascus Zukal (Plectomycetes (Benny & Kimbrough, 1980). Up to now, only few species of Plectomycetes have been found solely in the sea: two species of the genus Gywmascella — G. hitoralis (Orr) Gurrah from marine animals and algae near the west coast of North America (Gurrah, 1985; Kohlmeyer & Volkmann- Kohlmeyer, 1991) and G. marnsmortui Buchalo from water of the Dead Sea (Buchalo ef a/. 1998) as well as Microasous senegalensis Arx from mangrove mud in Senegal and Mexico (Arx, 1975; Kohlmeyer, 1986). Members of Aphanoascus are keratophiles widely distributed in soils, dung and birds’ nests, but have not been found in the sea (Cano & Guarro, 1990; Cano, Guarro & Zaror, 1990; Udagawa & Lichiyama, 1999). Therefore the species described below ts of particular interest. Aphanoascus aciculatus Pivkin et Khudyakova sp. nov. (Figs. 1-2) Etym.: aciculatus = needle shaped, conidia are needle-like under the light microscope. Coloniae in agaro ad 25° C densae, lente crescunt, cumulae, circa 5-6 mm elevatae, concentrice plicatae, velutinae vel pulveraceae, albae, in centro luteolae vel citreae; reversum incoloratum vel flavum. Mycelio vegetative ex hyphis hyalinis, ramosis, tenuis, levis, septatis, 1.54 um diam. composito. Ascomata in medium "oryza cum aquata marina" superficialia, sphaerica, non- ostiolata, levia, 400-900 [im diam., brunnea; peridio 4-6 Um crasso, et 34 stratis cellularcum complanatarum composito, 3-6 x 5-8 um constant. Asci numerosi, octospori, subglobosi, evanescentes, 6.5—7.5 x 4.5-7 um. Ascosporae subglobosae vel oblatae, pallide brunneae, unicellulares, 2.2—4 um diam., 1.5—2.2 um crassae. Fig. 1 Aphanoascus aciculatus (LE! 211060). A, peridium; B, ascospores; C, asci; D, conidia (bars = 10 um). Anamorphosis: Malbranchea sp. Hyphae fertiles irregulariter dichotomae,; hyphae primariae 1.5—3 um diam., ramis fertilibus arthroconidis. Arthroconidia hyalina, levia, cylindracea vel leniter curvula, 4—9 (—20) x 1.5—2.2 um. Holotypus: LE 211060, ex sedimentis marinus, Russia, regio Primorskiensis, mare Japonicae, sinus Amurensus, 07.08.1998: Colonies on YpSs and PYE with natural seawater at 25° C are dense, slow- growing (18-22 mm in diameter after 6 weeks), concentrically rugate, velvety at first, becoming powdery, raising in the centre up to S—6 mm, white, but become yellowish or lemony in the centre, reverse colourless or pale yellow. A lemon yellow pigment diffuses into the agar. The vegetative hyphae are narrow, 1.5—4 um diameter. Teleomorph forms on rice medium with seawater after the growing in the flasks during two months. Ascomata are superficial, globose, non-ostiolate, brown, 400-900 um diameter, glabrous; cleistoperidium of “textura angularis” 4—6 um thick, 3-4 layers "LE — Herbarium of V. L. Komarov Botanical Institute RAS, St. Petersburg. Fig. 2 Aphanoascus aciculatus (LE 211060). A, ascospores; B, conidia; C, asci; D, E ascospores (bars A-C = 10 um, D=5 um, E= 1 um); A-C (LM*); D, E (SEM). deep; cells are flattened, 3-6 x 5-8 um. Asci are numerous, globose to subglobose, 6.5— 7.5 x 4.5-7 um, 8-spored, evanescent. Ascospores are unicellular, oblate with polar depressions, smooth under LM to slightly rugose under SEM, light brown, 2.2—4 x 1.5- 2.2 um. Anamorph: Malbranchea sp. Fertile hyphae arise as long side branches from aerial hyphae or as terminal branches of aerial hyphae, often form an irregular dichotomic branching structure. Arthroconidia are hyaline, smooth-walled, cylindrical or slightly bent, 4—9 (—20) x 1.5—2.2 um. Holotype: LE 211060, from sea bottom deposits, Russia, the Sea of Japan, Amursky Bay, 07.08.1998. , Material examined: KMM” 4688 from sea bottom deposits, the Sea of Japan, Trinity *LM - light microscopy, SEM — scanning electron microscopy. "KMM -— Collection of Marine Microorganisms, Pacific Institute of Bioorganic Chemistry, the Far East Branch of the Russian Academy of Sciences, Vladivostok. In “World Directory of Collections of Culture of Microorganisms. Bacteria, Fungi, and Yeasts” 4" ed. WFCC World Data Center on Microorganisms, 1993. 10 Bay, 42°34'00" N, 130°57'29" E, at the depths of 17 m, 27.10.1996; KMM 4692 from sea bottom deposits, the Sea of Japan, Amursky Bay, 44°06'52" N, 131°45'40" E, at the depths of 15 m, 07.08.1998; KMM 4698 from sea bottom deposits, the Sea of Japan, Amursky Bay, 44° 06'50" N, 131° 45'38" E, at the depths of 12 m, 07.08.1998. A, aciculatus belongs to the group of Aphanoascus characterized by ascospores with a prominent equatorial rim. Almost all members of this group, including A. durus (Zucal) Cano et Guerro, A. clathratus Cano et Guerro, A. hispanicus Cano et Guerro, A. ferreus (Randhawa et Sundhu) Apinis (Cano & Guerro, 1990), and A. punsolae Cano et Guarro . (Cano, Guerro & Zaror, 1990) have Crysosporium anamorphs. A. boninensis Udagawa et Lichiyama (Udagawa & Lichiyama 1999) have a Chrysosporium-like anamorph. In this group A. acaculatus and A. multiporus Cano et Guarro (Cano & Guerro, 1990) alone have Malbranchea anamorph. However A. multiporus has pitted large ascospores (5-6 x 3.54 Um), whereas ascospores of A. aciculatus are smaller and smooth. Under the light microscope ascospores of A. aciculatus are allied to A. durus, but they are smaller and smooth. _ Ascospores of A. durus are foveate-reticulate, smooth in the poles, 44.5 x 2.7-3 um. Among Aphanoascus species, only A. punsolae has smooth ascospores that are also larger (4— 4.5 x 2—2.5 tum) than those of A. acaculatus. ACKNOWLEDGEMENTS This work was financially supported in part by grant # 99-04-49363 of the Russian Foundation for Basic Research. Our special thanks to Mrs. N. Shepetova for correcting the English text. REP ERENGES Arx, von J.A. (1975). Revision of Mécroascus with the description of a new species. Persoonta, 8: 191-197. Buchalo, A. S., E. Nevo, S. P. Wasser, A. Oren, & H. P. Molitoris (1998) Fungal life in the extremely hypersaline water of the Dead Sea: first records. Proceedings of the Royal Society of London, B, 265: 1461-1465. Kohlmeyer, J. & B. Volkmann-Kohlmeyer (1991) Illustrated Key to the Filamentous Higher Marine Fungi. Botanica Marina, 34: 1-61 Kohlmeyer, J. (1986) Taxonomic studies of the marine Ascomycotina. In “The biology of marine fungi.” Ed. S. T. Moss, Cambridge University press. Cambridge, London, New York, New Rochelle, Melbourne, Sydney: 199-210. Gurrah, R. S. (1985) Taxonomy of the Onygenales: Artrodermataceae, Gymnoascaceae, Myxotrichaceae and Onygenaceae. Mycotaxon, 29: 1-216. Benny, G. L. & J. W. Kimbrough (1980) A synopsis of the orders and families of Plectomycetes with keys to genera. Mycotaxon, 12: 1-91. Cano, J. & J. Guarro (1990) The genus Aphanoascus. Mycological Research, 94: 355-377. Cano, J., J. Guarro & L. Zaror (1990) Two new species of Aphanoascus (Ascomycotina). Mycotaxon, 38: 161-166. Udagawa, S. & S. Lichiyama (1999) Taxonomic studies on new or critical fungi of non- pathogenic Onygenales 1. Mycoscience, 40: 277-290 MYCOTAXON Volume LXXXI, pp. 11-25 January-March 2002 CLAVICEPS PURPUREA VAR. SPARTINAE VAR NOV.: AN ERGOT ADAPTED TO THE AQUATIC ENVIRONMENT RUSSELL A. DUNCAN, JR.', RAYMOND SULLIVAN', STEPHEN C. ALDERMAN’, JOSEPH W. SPATAFORA’, JAMES F. WHITE, JR.' 'Department of Plant Biology and Pathology, 59 Dudley Rd., Cook College-Rutgers University, New Brunswick, New Jersey 08901; *USDA/ARS National Forage Seed Production Center, Corvallis, Oregon 97331; ‘Department of Botany & Plant Pathology, Oregon State University, Corvallis, Oregon OSL ABSTRACT We investigated systematic relationships and morphological and ecological features of an ergot that commonly infects species of the maritime grass Spartina. Phylogenetic analysis of ribosomal DNA Internal Transcribed Spacer 1 sequences demonstrated that the Spartina ergot is placed in a clade with Claviceps purpurea. Structural differences of sclerotia distinguish it from the Claviceps purpurea var. purpurea as seen on rye and other grasses. Sclerotia from Spartina are larger and contain large intercellular spaces and have thicker rinds than sclerotia of C. p. purpurea. Sclerotia of the Spartina Claviceps float in 1% saline and fresh water while those of C. p. purpurea sink. In addition, the Spartina Claviceps has a unique alkaloid profile. In recognition of the differences between C. purpurea and the Spartina ergot a new variety, C. p. spartinae var. nov., is proposed. KEY WORDS: Ascomycetes, Clavicipitaceae, Spartina, sclerotia, salt marsh INTRODUCTION The genus Claviceps was established by Tulasne (1853) for Sphaeria purpurea Fries from rye (Secale cereale L.) (Saccardo, 1883). To date, over fifty species and varieties of Claviceps have been described, based on details of the ascoma, asci, - ascospores, sclerotia, conidia, and host range. Species of Claviceps infect flowers of grasses, transforming the ovary into a sphacelium and then to a sclerotium. The sclerotium and the disease caused by Claviceps are commonly referred to as ergot (Bove, 1970). Claviceps species are widely distributed and occur in temperate, arid, subtropical and tropical environments (Brady, 1962). Most species have a narrow host 12 range of one to several host genera. Only one species, C. purpurea (Fr.) Tul., is known to have a wide host range, reported from over 100 genera of Pooideae (Brady, 1962; Farr et al., 1989). Claviceps purpurea has been suggested to be comprised of multiple cryptic species that may be distinguished on the basis of conidial morphology and host affiliation (Loveless, 1971). The ergot on rye is considered the type of C. purpurea since early descriptions specifically mentioned this host (Saccardo, 1883). Claviceps purpurea has been reported as the causal agent of ergot of Spartina (Boyle, 1976; Gessner, 1977, 1978). Samuelson and Gjerstad (1966) were the first to suggest that the alkaloid profile of Claviceps from Spartina differed from those of C. purpurea. Eleuterius and Meyers (1977) documented the unique alkaloid profile of this ergot, which they denominated the “feather ergot”. Sclerotia from the rye ergot contained alpha and beta forms of ergocryptine while the Spartina ergot contained only the beta form. Loveless (1971) and Pazoutova (2000) suggested larger conidial size as a characteristic of Claviceps on Spartina anglica Hubbard. To further evaluate the relationship of the Spartina ergot to C. purpurea, data were obtained on morphology, ecology and ITS1 sequences. MATERIALS AND METHODS Herbarium collections were obtained from the National Fungus Collection (BPI) in Beltsville, Maryland (Table 1). Sclerotia were collected from Spartina alterniflora Loisel. plants located on the banks of the South River in South River, New Jersey. Germination of sclerotia was facilitated by surface sterilization using 50% Clorox® for 5 min followed by 3 rinses with sterile water. Sclerotia were then placed on the surface of sterile sand in petri dishes. The sand was kept moist using distilled water until sclerotia germinated. The dishes containing these sclerotia were placed on a lab bench where they remained throughout their germination. The temperature in the lab was about 25C°. Ascospores were ejected onto glass slides for measurement and onto potato dextrose agar (PDA, Difco) plates for isolation into pure culture. Ascomatal and sclerotial tissues were fixed in 3% glutaraldehyde, postfixed in 1% osmium tetroxide and embedded in Quetol 651® (Electron Microscope Sciences, Ft. Washington, Pennsylvania). Thin sections of ascomata were cut with glass knives, mounted on glass slides, stained with aqueous aniline blue (0.1%), and photographed using a compound microscope. For rind measurements 10 sclerotia of each host, Spartina alterniflora, Festuca rubra, Secale cereal, Leymus chiniensis and Lolium perenne, were sectioned by hand using a metal razor, transferred to glass slides, and viewed and measured on a videoscope screen. Conidial measurements were taken by removing conidia from culture plates (PDA) and measuring under a compound microscope with an ocular micrometer. 13 Flotation experiments were conducted by placing sclerotia in a beaker of water. Density of fresh Spartina sclerotia was estimated by measuring the density of an alcohol — water mixture that had been adjusted to the point of equilibrium between the sclerotia and the solution. Sclerotia that were observed to sink were placed in a sodium chloride — water mixture adjusted to the point of equilibrium. Density of each mixture was measured by use of a density hydrometer. A long-term flotation experiment was conducted by agitating sclerotia in a tray of 1% saline water for 30 days. Sclerotia from Secale cereale L., Leymus chinensis (Trin.) Tzvelev, Lolium perenne L., and Festuca rubra L. were also included in this test. Table 1: Collections of C. p. spartinae examined Location Bodie Island, N. Carolina Lander, Maryland Atlantic City, New Jerse Terry’s Creek, Georgia Costa Martina, Urugua Howard Co., Nebraska Bay Co., Florida Richmond, Virginia Belgrade, Montana Valley Lee, Maryland Seaport, Maine 634402 |OE White | Oct. 1909 _| Brookings, South Dakota Spartina sp 634400 | JT Rogers Nov. 1923 | Charleston, South Carolina 634432 |OE White | Dec. 1919 | Searsport, Maine 634431 Dec. 1887 | Flatlands, Louisiana 634429 Nov. 1935. | Hyannis, Massachusetts 634430 Sept. 1965 | Elkhorn, Canada 14 DNA EXTRACTION AND SEQUENCING Total genomic DNA from mycelium of C. purpurea from S. alterniflora was extracted from cultures grown on PDA overlaid with cellophane. Tissue was ground to fine powder in liquid N2. DNA extraction was performed using the procedures of Lee and Taylor (1990). Successful PCR amplification resulted in a single DNA band corresponding to ~600 bp (ITS1, 5.8S ITS2 rDNA). Cycle sequencing reactions were performed using an Amplitaq FS Dye Deoxy Terminator Cycle Sequencing kit (Applied Biosystems Inc., Foster City, California). Reactions were run on a Sho) s\ DNA sequencer (Applied Biosystems Inc., Foster City, California). Other DNA sequences were obtained from GenBank, including C. citrina Pazoutova, Fucik, Leyva-Mir & Flieger from Distichlis spicata (L.) Greene (GenBank AJ000068), C. africana Freder., Mantel & De Milliano from Sorghum sp. Moench (GenBank AJ01784), C. sorghicola from Sorghum sp. (GenBank AJO11591), C. purpurea (Fr.: F) Tul. from Lolium sp. (L.) (GenBank AJ000069), C. purpurea from Phalaris sp. (L.) (GenBank AJ000070), C. purpurea from Dactylis glomerata (L.) (GenBank U57669), Balansia claviceps Speg. from Panicum ghiesbrechtii Fourn. (GenBank U89366), and B. asclerotiaca Henn. from Orthoclada laxa (GenBank U89368). PHYLOGENETIC ANALYSIS Alignment of sequences was accomplished using GCG (Genetics Computer Group, Madison, Wisconsin) program with default program options. Visual adjustments of aligned sequences were made to refine the alignment. To evaluate groupings, sequences were analyzed using PAUP 4.0 Beta version (Smithsonian Institution, Washington D.C.) on a Power Macintosh 9500/132. Maximum parsimony analysis of the aligned sequences was performed with alignment gaps treated as missing data. Heuristic searches were performed with the following options in effect: tree-bisection- reconnection, branch-swapping algorithm, random sequence additions option set to 100, collapse zero length branches to yield polytomies and save all minimal length trees (MULPARS) (Swofford, 1991). To measure relative support for the clades, 500 bootstrap replicates were performed. Bootstrap values were mapped onto the tree (Fig. 1) to show support for groupings. Balansia claviceps and B. asclerotiaca were included as outgroup. C. citrina Distichlis sp. C. purpurea Lolium sp. 100 C. purpurea Phalaris_ sp. 100 C. p. spartinae Spartina sp. 68 C. purpurea Dactylis sp. C. africana Sorghum sp. 100 10 changes C. sorghicola Sorghum sp. Balansia claviceps Outgroup Balansia asclerotiaca Fig. 1. Tree resulting from maximum parsimony search of rDNA ITS1 sequences. (Length 201 steps; CI = 0.97; HI = 0.04; RI = 0.95). Bootstrap values (heuristic with 500 replicates) are mapped onto tree. 16 RESULTS ASCOMATAL DEVELOPMENT Sclerotia from Spartina alterniflora germinated after 30 days in moist sterile sand at room temperature (Figs. 2-6). Stipe elongation and perithecia formation were observed 2-3 days after germination (Figs. 4-6). Stroma formation begins with the emergence of a purplish-white cottony cushion on the surface of sclerotia. The purplish stipe then forms beneath the cushion and elongates as the cushion differentiates to form a roughly spherical cap or ascomatal head. The red ostiolar necks of perithecia are emergent from beneath the surface of the ascomatal head. Perithecia are obpyriform (Fig. 7) with ostiolar necks containing lateral periphyse (Fig. 8). Mature ascospores were discharged 4-5 days after germination of the sclerotium. Table 2: Sclerotial characteristics Rind thickness| Density _ 17.2 + 3.92 X 1.70 +0.56 mm 0.90 g/cc 07 glec Leymus chiniensis _ |5.86 + 1.25 X 1.61+0.39 mm |no [46.8 + 10.5 um |1.05 g/cc 1.07 gee 1.10 g/cc Bib |n ln Psi CHARACTERISTICS OF SCLEROTIA The rind of the sclerotia from Spartina was significantly (P > 0.05) thicker than that of the rye ergot (Table 2, Fig. 9, 12F). The rind contained an inner cellular melanized component and an outer region composed of inflated, often collapsed, cells that appeared to lack cytoplasmic content (Figs. 9, 12F). The rinds of sclerotia from the Festuca rubra ergot were seen to contain an acellular melanized component; an outer layer composed of inflated cells was not evident (Figsi steed. Gy Figs. 2-6. Sclerotia and ascomatal stromata. 2. Sclerotia from Spartina showing subulate shape and lighter basal pigmentation, X 2. 3. Clavate sclerotia of the C. purpurea from Festuca rubra (arrowheads) shown with a single larger subulate sclerotium from Spartina, X 2. 4, Ascomatal stroma of C. purpurea from Spartina, X 8. 5. Ascomatal head showing numerous perithecia, X 27. 6. Ascomatal stroma of the C. purpurea from Spartina showing bifurcation to form two ascomatal heads, X 17. U7 18 Sclerotia of the Spartina ergot contain a large number of intercellular spaces (Fig. 10). Fewer intercellular spaces were observed in the sclerotia of the Festuca ergot (Fig. 11). In a flotation experiment, the sclerotia from S. alterniflora remained afloat during a 30 day long trial in agitated salt water. Removal of the rind did not effect the floatation of the sclerotia of the Spartina ergot. Sclerotia from Secale cereale L., Leymus chinensis (Trin.) Tzvelev, Lolium perenne L., and Festuca rubra L. sank in water (Table 2). TAXONOMIC ANALYSIS Analysis of the rDNA ITS1 sequences of several collections of Claviceps spp. demonstrated that the Spartina ergot grouped within the C. purpurea clade (100% bootstrap support). Pazoutova and Parberry (1999) demonstrated the utility of analysis of the rDNA ITS1 sequence to separate several species of Claviceps. It seems appropriate to weigh heavily the placement of the Spartina ergot within the phylogenetic species C. purpurea. However, several features of the Spartina ergot distinguish it from C. p. purpurea as exemplified by ergots from Secale cereale L., Leymus chinensis (Trin.) Tzvelev, and Lolium perenne L. (see Tanda, 1978; Table 2). Among the distinguishing features of the ergot from Spartina are: 1) thicker sclerotial rind, 2) numerous large intercellular spaces, 3) capacity to float, 4) larger conidia, 5) host range restricted to Spartina, 6) cold conditioning not required, and 7) unique alkaloid profile. It is the opinion of the authors that the many unique features of the ergot from Spartina and its adaptation to a maritime habitat are sufficient to consider this ergot as a unique variety of C. purpurea. DESCRIPTION Claviceps purpurea var. spartinae R. Duncan & J. F. White, var. nov. Sclerotiis 17.2 + 3.9 mm longis, 1.7 + 0.5mm latis, purpureo-brunneis vel atro- brunneis, intus albidis, subcurvatis, subulatatis; stromatibus simplex ad tetrafurcatus; stipite 4.6 + 3.5 mm longo, 1.3 + 1.7 mm lato, erecto vel subcurvato, brunneo-violaceo; Figs. 7-11. Anatomy of ascomata and sclerotia of C. purpurea. 7. Vertical section of perithecium of the C. purpurea from Spartina, X 385. 8. Section through ostiole of perithecium of C. purpurea from Spartina showing periphyses, X 1000. 9. Section of sclerotium of C. purpurea from Spartina showing rind, X 560. 10. Section through sclerotium of C. purpurea from Spartina showing large intercellular spaces (black dots), X 560. 11. Section through sclerotium of C. purpurea from Festuca rubra showing thin rind (arrowheads) and small intercellular spaces (black dot), X 560. 19 20 capitulum 1.4 + 0.5 mm diametro, globosum vel subglobosum, albidum, punctulatum con ostioli rubelli; peritheciis immersis, ovoideis, 144.2 + 23.9 jum longis, 91.8 + 11.2 im crassis; ascis hyalinis, cylindraceis, flexuosis, in basi attenuatis, in apice rotundatis, _ octosporis 62.5 + 1.0 um longis, 2.6 + 0 pm crassis; ascosporis hyalinis, filiformibus, 4 - 5 septata, 84.4 + 4.5 um longis, 0.8 + 0 um crassis. Anamorphus Sphacelia segentum Leva Habitatum: in ovariis vivis Spartina alterniflora Loisel. Type: Claviceps purpurea var. spartinae on Spartina alterniflora, on the banks of South River, South River, NJ, USA. R. A. Duncan, Nov. 1997. Rutgers University Mycological Herbarium RUTPP3419. Sclerotia 17.2 + 3.92 X 1.7 + 0.5 mm, purple brown or dark brown, interior white, curved, often subulate, showing the capacity to float in fresh and salt water; stromata simple to four-forked; stipe 4.6 + 3.5 X 1.25 + 1.73 mm, straight or slightly curved, purple-brown; ascomatal head 1.3 + 0.4 mm in diameter, globose to subglobose, white to slightly tan, punctate with reddish ostioles; perithecia immersed to exposed, ovoid, 144.2 + 23.9 K 91.8 + 11.2 um; asci hyaline, cylindrical, flexuose, at base attenuate, at apex rounded with refractive tip, containing eight filamentous ascospores, 62.5 + 0.9 X 2.6 + 0 um; ascospores hyaline, filiform, 4 - 5 septa, 84.4445 X8+0 um. Anamorph: Sphacelia segentum Lévy. Conidia, enteroblastic, cylindrical, uninucleate, 8.06 + 1.8 X 4.12 + 1.3 um, occasionally septate in larger conidia; conidiogenous cells, phialidic, ampuliform, 2.9 + 0.1 X 4.2 + 0.2 um, budding conidia apically, with thickened walls bordering conidiogenous locus. DISCUSSION Historically Claviceps species were defined based on host range and details of the stromata (Langdon, 1954). Loveless (1964) suggested size and shape of conidia may also be of value in species separation. More recently molecular phylogenetic analyses have played an important role in defining relations among taxa (Pazoutova and Parberry, 1999). In the case of Claviceps from Spartina, a suite of characteristics including physical, physiological, and molecular genetic differences can be applied to separate it Fig. 12A-F C. purpurea. A, B. Sclerotia of Claviceps purpurea from Festuca rubra, X 4. C. Section through sclerotium of C. purpurea from Festuca rubra showing thin rind, X 100. D. Florets of Spartina alterniflora infected by C. p. spartinae, X4._ E. Germinating sclerotium of C. purpurea from Spartina showing the rind composed of multiple layers, X 100. 21 22 as a variety from C. p. purpurea. The physical properties of the sclerotia including the large intercellular spaces, which permit flotation, adapt it to the aquatic environment. The alkaloid profile, which differs from C. p. purpurea, suggests a physiology that differs from that variety. The lack of cold conditioning period indicates a further - physiological separation from C. purpurea and also an adaptation to southern coastal waters where the cold conditioning period would likely not be met. Although the host range of C. p. spartinae has not been fully resolved it is likely restricted to Spartina. Althought C. purpurea has been assumed to infect Spartina, infection studies indicate otherwise (Mantel, 1969). In terms of host range, it is not unexpected that C. p. purpurea cannot infect Spartina. Claviceps purpurea var. purpurea is largely restricted to Pooid grasses; Spartina is a member of the Chloridoid group. Tanda (1978, 1979) separated Claviceps purpurea into four varieties, including C. p. purpurea, C. p. alopecuri Tanda, C. p. phalaridis Tanda and C. p. sasae Tanda. Our use of anatomical features of sclerotia along with capacity for flotation to establish C. p. spartinae follows the precedent set by Tanda. Pazoutova (2000) separated C. purpurea into races based on alkaloid profiles and ecological specialization. The establishment of varieties to identify distinctive types that bear obvious close affinity to C. purpurea seems warranted and consistent with the development of a phylogenetic species concept. Spartina spp. are important components of estuarine ecosystems. In many salt marshes, S. alterniflora is frequently the dominant plant species (Eleuterius and Meyers, 1974; Raybould et al., 1998). Eleuterius (1970) reported that 96% of the flowering culms of S. alterniflora bore florets infected with C.p. spartinae. This ergot thus is a very common parasite of S. alterniflora and has been reported to also infect S. patens (Ait.) Muhl. and S. cynosuroides (L.) Roth in North America, and S$. maritima (M.A. Curtis) Fern in Argentina (Eleuterius and Meyers, 1977) and S. anglica in Great Britain (Raybould et al., 1998). We have further found this ergot on S. brasiliensis Rad. from Uruguay, and S. foliosa Trin., S. michauxiana A.S. Hitchc., S. stricta (Ait.) Roth, and S. pectinata Link from North America (Table 1). The structure of sclerotia of C. p. spartinae appears to be a result of adaptation to its semi-aquatic grass hosts. Eleuterius and Meyers (1974) reported that sclerotia were often dislodged by wave action and transported with the tide before being deposited along the periphery of the marsh. Sclerotia of C. p. spartinae contain large intercellular spaces (Fig. 10). In contrast, sections of non-floating sclerotia from C. p. purpurea (Festuca rubra) (Fig. 11) revealed that such spaces are rare. The flotation capacity of sclerotia of C. p. spartinae may stem from gasses that may accumulate in intercellular Figs.13-16. Conidia and conidiogenous cells of Claviceps purpurea from Spartina. 13. Conidiogenous cell showing ampuliform structure (Bar = 2um). 14. Conidia (Bar = 10pm) 15 - 16. Conidia with conidiogenous cell indicated by arrow (Bar = 10um). 755} 7 mn Ww i i . a i < Hi Lh Heit i Hy . : 24 spaces, and thus the tissues may in effect be aerenchyma. The larger size of sclerotia of C. p. spartinae may enhance their capacity for dispersal in water. The thicker rind of sclerotia of C. p. spartinae may provide them with added protection from salts that concentrate in the mud along the margins of estuaries where the sclerotia eventually lodge. The widespread distribution of C. p. spartinae in North America, South America, and Europe may be due to the long distance dispersal of sclerotia in ocean currents. In one flotation experiment, sclerotia of C. p. spartinae showed no evidence of losing the capacity to float even after 30 days of agitation in 1% salt water. Dispersal and survival in the marine environment must have been a significant component of the selective pressures that molded the divergence of C. p. spartinae from C. p. purpurea. ACKNOWLEDGEMENTS The authors would like to thank Dr. Sylvie Pazoutova (Institute of Microbiology, Czech Republic) for her input on this paper and for general discussion of Claviceps systematics. We are grateful to Dr. David Farr (U.S.D.A., Beltsville, MD) for his review of the manuscript. We would also like to thank the National Science Foundation (DEB 96-96041) and the Rutgers Center for Turfgrass Science for their financial support of this research. LITERATURE CITED BOVE, F. J. 1970. The Story of Ergot. Basel (Switzerland): S. Karger. BOYLE, P. J. 1976. Ergot Epiphytic on Spartina spp. in Ireland. Irish Journal of Agricultural Science 15:419-424. ELEUTERIUS, L.N. 1970. Observations on Claviceps purpurea on Spartina alterniflora. Gulf Res. Rep. 3:105-109. Mycologia 66:978-986. ELEUTERIUS, L.N. and S.P. MEYER. 1974. Claviceps purpurea on Spartina in coastal marshes. Mycologia 66:978-986. ELEUTERIUS, L.N. and S.P. MEYER. 1977. Alkaloids of Claviceps from Spartina. Mycologia 69:838-840. FARR, D.F., G.F. BILLS, G.P. CHAMURIS, and A.Y. ROSSMAN. 1989. Fungi on plants and plant products in the United States. APS Press, St. Paul, Minnesota. KIRCHOFF, H. 1929. Beitrage zur Biologie und Physiologie des Mutterkornpilzes. Centralbl. Fiir Bakt. 77:310-369. LEE, S. and J. TAYLOR. 1990. Isolation of DNA from fungal mycelia and single spores.Pp. 282-287. In PCR protocols. Eds, M.S. Innis, D.H. Gelfand, J.J. Sninsky and T.J. White. Academic Press, New York. LOVELESS, A.R. 1964. Use of the honeydew state in the identification of ergot species. Trans. Brit. Mycol. Soc. 47:205-213. LOVELESS, A.R. 1971. Conidial evidence for host restriction in Claviceps purpurea. Trans. Br. Mycol. Soc. 56:419-434. MANTLE, P.G. 1969. Development of alkaloid production in vitro by a strain of Claviceps purpurea from Spartina townsendii. Trans. Brit. Mycol. Soc. 52:38- 392. MANTLE, P.G., S. SHAW, and D.A. DOLING. 1977. Role of weed grasses in the etiology of ergot disease in wheat. Ann. Appl. Biol. 86:339-351. MITCHELL, D.T. and R.C. COOKE. 1968. Some effects of temperature on the germination and longevity of sclerotia in Claviceps purpurea. Trans. Brit. Mycol. Soc. 51: 721-729. 25 PAZOUTOVA, S. and D.P. PARBERRY. 1999. The taxonomy and phylogeny of Claviceps. Pp. 57-77 In Ergot: the genus Claviceps. Eds. V. Kren and L. Cvak Harwood Academic Publishers, The Netherlands. PAZOUTOVA, S., J. OLSOVSKA, M. LINKA, R. KOLINSKA, and M. FLIEGER. 2000. Chemoraces and habitat specialization of Claviceps purpurea populations. Applied and Enviromental Microbiology 66:5419-5425. RAYBOULD, A.F., A.J. GRAY, and R.T. CLARKE. 1998. The long-term epidemic of Claviceps purpurea on Spartina anglica in Poole Harbour: pattern of infection, effects on seed production and the role of Fusarium heterosporum. New Phytologist 138:497-505. SACCARDO, P.A. 1883. Sylloge fungorum omnium hucusque cognitorum. Vol. **. Sumptibus auctoris, Patavil, Italy. SAMUELSON, R.J. and G. GJERSTAD. 1966. Intermediary metabolism of ergot XIII: Feather ergot, a new, promising Claviceps species. Meddel. Norsk Farm. Selsk. 28:229-237. SWOFFORD, D.L. 1991. When are phylogeny estimates from molecular and morphological data incongruent? Pp. 295-333. In: Phylogenetic analysis of DNA sequences. Eds. Miyamoto and Cracraft. Oxford University Press, New York. TANDA, S. 1978. Mycological studies on ergot in Japan IV. Two varieties of Claviceps purpurea Tul. parasitic on timothy, Phleum pratense L. Journal of Ag Science. Tokyo. Nogyo Daigaku 23:141-148. TANDA, S. 1979. Mycological studies of ergot in Japan. IX. Distinct variety of C. purpurea Tul. on Phalaris arundinacea L. and P. arundinacea var. picta L. Journal of Agricultural Science, Tokyo. Nogyo Diagaku 24:79-95. TULSANE, L.R. 1853. Memoire sur l’ergot des glumacés. Annales des Sciences Naturelles Botanique 20:5-56. rs = J ae ts +7; ee Miss Ma ‘ in ‘sue hone matt | iy ' i. j Ee RE eA a ea Holttvn ge i in ia ba a Natta Ap Olas Wg a i a ‘e reas area Tea EY, tailings te ah vr hignent Suh Na | 5% | eh bia ie nee eet Shatin AN aM cab i a ae = Me aie sna’ Pai ee Mer ely, i a on * pa ae . , FRR (i i & a a oe ie a \s rh ih ; i PX ae i] : 4 vi s ees 3) — fo = i 4 a ii iy aw on . 7 Lone i py, wy) ; ; ' iy i a | ¥ MJ J Ne ty re te Deca a ; + WAL : P f : ; PS) aes / RRL ‘ x# a ass Wi : ole ae Ned ha hee , itis ¢. ! ‘7 rn, py oe a ' ; - ey oh A Gs a y; eee , ; i i Li bed a TS Vs wheel oe 4 Vt has Lt) was: j AV ola ad | as! x wie 7 ; Fit Wie, TAY ry ’ t ty ; f Bas ‘f vey [ «i ve uf cy MP j ay \ t Ste hy } } : r Vout oe gt ioe it A Reh oh ns as ry — Hits) ‘ thee cd ae ne ae te Ls a Mey te 4 hl rr) 7 : = a <e zt if. 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Ia 6) (fees “d vk ait ; i nti ey We ony oy 28 re aoe Dh a Neal ae i 7 Al ep \ ‘i VRS GA ve MYCOTAXON Volume LXXXI, pp. 27-34 January-March 2002 SOME NEW SPECIES AND NEW RECORDS OF DISCOMYCETES IN CHINA. IX" WEN-YING ZHUANG, ZHI-HE YU, YAN-HUI ZHANG, MING YE Systematic Mycology & Lichenology Laboratory Institute of Microbiology, Chinese Academy of Sciences, Beijing 100080, China ABSTRACT: Psilachnum hainanense is described as a new species. Four taxa in the genera Cistella, Perrotia, Proliferodiscus, and Vibrissea are reported for the first time from China. Infraspecific variations among collections of six other discomycetes in tropical and subtropical regions are discussed. KEY WORDS: Taxonomy, Tropical China, Psilachnum hainanense In our studies on the Chinese discomycete flora, collections from tropical and subtropical areas of China were examined. A new species, Psilachnum hainanense, was encountered and is described. Four new records for China are reported, of which two possible new taxa are involved. Morphological diversities have been found in several previously recorded discomycetes from different localities and at different elevations, which are treated as infraspecific variations. Measurements were carried out in cotton blue lactophenol and ascus iodine reaction was observed in the Melzer’s reagent without KOH pretreatment except as otherwise mentioned. TAXONOMY Cistella cf. hungarica (Rehm) Raitv., Scripta Mycol. 8: 151, 1978. (BFR) a) 7A /)\ 5 ik Al Apothecia discoid, 0.6-1 mm diam, hymenium yellow, receptacle whitish, surface downy or pubescent; hairs subcylindrical at lower portion, slightly swollen at apex, granulate at upper portion and smooth-walled below, hyaline, 1-3-septate, up to 70 x 2.5-3.5 wm; ectal excipulum of textura prismatica, cells hyaline, thin-walled; medullary excipulum of textura intricata, hyphae hyaline; hymenium 45-50 um thick; asci clavate to subcylindrical, 8-spored, with crozier at base, J+ in Melzer’s reagent, 35-47 x 4.2-5 um; ascospores rod-shaped, cylindrical-fusoid to clavate, unicellular, * Supported by the National Natural Science Foundation of China and Foundation of the Knowledge Innovation Program of the Chinese Academy of Sciences. E-mail: zhuangwy@sun.im.ac.cn 28 mostly with 2 small polar guttules, biseriate, 5-6.5(-7.5) x 1.4-2 ym; paraphyses subcylindrical, 1.8-2 um wide. | Specimen examined: China. Hainan: Ledong, Jianfengling, alt. 1000 m, on a small twig, Z.H. Yu, W.Y. Zhuang & Y.H. Zhang 3788, 10 XII 2000, HMAS 78607. Notes: The apothecial gross morphology of our fungus resembles, in some way, that of Proliferodiscus J. H. Haines & Dumont, but the distribution of granules on hair surface does not fit. Hairs of the Hainan collection possess a granulate upper portion and a smooth-walled basal part which are typical in Ciste//a Quél. Compared with existing species of the genus, the shape and size of asci and of ascospores and hair septation in our collection are similar to those of C. hungarica (Raitviir, 1970, 1978); the description of the latter differs in the shorter hairs 15-40 x 3-4 um, shorter and wider asci 27-40 x 4-6 um, and smaller apothecia 0.15-0.5 mm diam. HMAS 78607 might represent an undescribed taxon. Unfortunately, it contains very few apothecia, mostly immature, and is too scanty to be a type. It is tentatively treated as Cistella cf. hungarica. Perrotia atrocitrina (Berk. & Broome) Dennis, Kew Bull. 17: 326, 1963. a Fide fia PA Apothecia discoid, sessile, 0.5-1 mm diam, hymenium olivaceous, olive-brown to dark brown, receptacle dirty yellow to pale brown, surface furfuraceous or hairy; hairs subcylindrical, pale yellow, closely septate, hair surface roughened to nearly smooth at and near margin, more or less granulate at apothecial base, 100-150 x 3.7-4.5 um, walls 0.7-1.5 jm thick; ectal excipulum of textura prismatica at margin and upper flank, ca 25-45 um thick, cells pale yellow, thick-walled, 10-17 x 3-5 um; medullary excipulum of textura intricata, ca 30-80 um thick, hyphae subhyaline, thick-walled, somewhat gelatinous, 2-4 um wide; hymenium 183-200 um thick, with gelatinous matrix; asci clavate with a narrow base and truncate apex, 8-spored, J- in Melzer’s reagent, (137-) 152-190 x 13-15 wm; ascospores vermiform, with 6-15 septa, 2-3- seriate, (24-) 28-45.5 x 4.8-6.5 um; paraphyses filiform, slightly enlarged at apex, 1.2- 1.7 um wide. Specimens examined: China. Hainan: Tongshi, Wuzhishan, alt. 1500-1700 m, on rotten bare wood, Y.H. Zhang, Z.H. Yu & W.Y. Zhuang 3873, 3874, 3879, 16 XII 2000, HMAS 81333, 81334, 81335; ibid. Z.H. Yu, Y.H. Zhang & W.Y. Zhuang 3883, HMAS 81336. Notes: Compared with the type material from Sri Lanka (as Ceylon) on bare wood (Dennis, 1963), the Chinese collections have longer hairs (100-150 um vs. 90 uum long) and wider ascospores (4.8-6.5 um vs. 4-5 um wide). These are certainly infraspecific variations. Proliferodiscus dingleyi Spooner, Biblioth. Mycol. 116: 635, 1987. J RKEWAE Apothecia discoid, a few proliferating, dried material 0.5-0.8(-1.2) mm in diam, stipitate, hymenium yellow, receptacle whitish to grayish, surface hairy, stipe base dark; hairs subcylindrical, undulate, 30-100 x 2.3-3 um, walls slightly thickened, up to 0.8 um thick, surface warted, warts 0.7-1.2 um diam and 0.2-0.4 um high; ectal 29 excipulum of textura angularis to textura prismatica, 18-30 um thick, cells subhyaline, 5-12 x 3-5.5 pm; medullary excipulum of textura intricata, ca 10-50 um thick, hyphae hyaline, 1-2.5 um wide; hymenium mostly 53-58 ym thick; asci cylindrical-clavate, J- in Melzer’s reagent, 42-51 x 4.5-5.5 um; ascospores elongate-ellipsoid, hyaline, non- septate, 6-7.8 x 1.4-1.8 um; paraphyses subcylindrical to narrowly lanceolate, not exceeding the asci. Specimen examined: China. Sichuan: Guan County, Qingchengshan, on stems of ?Rubus sp., Q.M. Ma et al. 106, 29 VI 1960, HMAS 30884. Notes: Proliferodiscus dingleyi was originally recorded from New Zealand on bark of Nothofagus and rotten wood (Spooner, 1987). The Sichuan collection extends distribution of the fungus to southwestern China. The Chinese material has slightly wider asci, longer ascospores, and wider paraphyses and is on a different substrate. Psilachnum hainanense W. Y. Zhuang, sp. nov. ##MkG2aE (Figs. 1-3) Ab Psilachno woodwardio apotheciis 0.2-1 mm diam; pilis apicaliter inflatis; ascis 45-70 x 4.6-7 «um; ascosporis fusoideis, uniseptatis, 4-guttulatis, irregulariter uniseriatis, 8.5-12 x 1.5-2.8 uum; paraphysibus subcylindricis differt. Apothecia discoid, 0.2-1 mm in diam, sessile, hymenium yellow, light yellow, greenish yellow to brownish, receptacle concolorous or paler than hymenium, surface downy; hairs as short hyphal protrusions with a swollen apex, non-septate, hyaline, smooth- and thick-walled, up to 4 um wide at apex and 2 um below, 7-13 um long, occasionally with a few delicate ones up to 26 x 1.3 um; ectal excipulum of textura prismatica, 15-20 ym thick; medullary excipulum of textura intricata, 15-35 um thick; Figs. 1-3. Psilachnum hainanense: 1. Short hyphal protrusions on receptacle surface. 2. Ectal excipular structure. 3. Ascospores. 2 from HMAS 81609; 1, 3 from HMAS 81341. Scale bars = 10 um. 30 hymenium 70-85 um thick; asci clavate, 8-spored, 45-70 x 4.6-7 um; ascospores fusoid, sometimes slightly curved, 1-septate, hyaline, usually with 4 small guttules and 2 in each cell, irregularly uniseriate, 8.5-12 x 1.5-2.8 bm; paraphyses subcylindrical, 1.5-2 um wide. Holotype: China. Hainan: Qiongzhong, Limushan, alt. 700 m, on twig, Y.H. Zhang, W.Y. Zhuang & Z.H. Yu 3958, 18 XII 2000, HMAS 81341. Paratypes: China. Hainan: Ledong, Jianfengling, alt. 1000 m, on rotten twig, W.Y. Zhuang, Y.H. Zhang & Z.H. Yu 3762, 10 XII 2000, HMAS 81608; Qiongzhong, Limushan, alt. 630 m, on herbaceous stems, Z.H. Yu, W.Y. Zhuang & Y.H. Zhang 4008, 19 XII 2000, HMAS 81609. Etymology: The specific epithet refers to the locality of the fungus. Notes: Psilachnum Hohn. was reviewed by Raitviir (1970) and its species were recorded by Dennis (1962, 1978), Graddon (1977, 1990), Sharma (1988), and Svrcek (1992) from different regions of the world. Among existing species, P. woodwardiae from India is the closest to our fungus, especially the shape, size, and septation of ascospores. The Indian fungus differs in smaller apothecia up to 375 pm in diam, smaller asci 30-50(-54) x 3.6-5.5 um, eguttulate ascospores, wide paraphyses up to 4.5 um wide, cylindrical hairs with 1-2 septa, and occurring on ferns (Sharma, 1988). Vibrissea cf. sporogyra (Ingold) Sanchez, Mycologia 58: 734, 1966. (28) hefal 7K ik Bal Apothecia on white subiculum, pulvinate, 0.7-0.8 mm in diam, sessile and broadly attached, hymenium dirty orange, receptacle lighter than hymenium; ectal excipulum of textura angularis, tissues gelatinous, 15-40 um thick; medullary excipulum of textura intricata, hyphae hyaline, 1-1.5 um wide; hymenium ca 160 um thick; asci subcylindrical, 8-spored, strongly thickened at apex, ca. 150-160 x 12-13 uum; ascospores cylindrical at upper portion, narrow at the lower end, with 20-32 or more septa, hyaline, 110-130 um long, 3.5-3.7 um wide in the upper portion and ca. 1.5 um at lower end; paraphyses filiform, branched and deformed apically, irregularly undulate, 1-1.5 um wide. Specimen examined: China. Hainan: Ledong, Jianfengling, alt 1100 m, on a living leaf of an unidentified dicotyledon, W.Y. Zhuang, Y.H. Zhang & Z.H. Yu 3716, 9 XII 2000, HMAS 81343. 3 Notes: The apothecia of this fungus are pulvinate and seated on a white subiculum on a living leaf. Compared with the description of Vibrissea sporogyra Sanchez (1967), the Chinese collection differs in the somewhat larger fruitbodies, asci J- instead of J+ in Melzer’s reagent, and shorter ascospores not twisted spirally within ascus. It might represent an undescribed taxon but the material contains only 4 apothecia and is too scanty to be a type. ; INFRASPECIFIC VARIATIONS SHOWN BY A FEW DISCOMYCETES FROM TROPICAL AND SUBTROPICAL CHINA Upon establishment, descriptions of the following discomycetes were based either on a single collection or on specimens from the same locality. Recently, SL specimens from different localities were examined, which gave better understanding of species concepts and revealed what we consider to be infraspecific variations. Updated information is provided for the taxa involved. Albotricha guangxiensis W. Y. Zhuang, Mycotaxon 69: 360, 1998. Specimens examined: China. Yunnan: Malipo, Laojunshan, alt. 1200 m, on culm of a large grass, W.Y. Zhuang & Z.H. Yu 3374, 10 XI 1999, HMAS 78140; Guangdong: Maoming, Dawuling, alt. 1400 m, on rotten culm of a large grass, W.Y. Zhuang & Z.H. Yu 2791, 22 X 1998, HMAS 78159; Hainan: Tongshi, Wuzhishan, alt. 1500-1800 m, on monocotyledon and bamboo, Z.H. Yu, W.Y. Zhuang & Y.H. Zhang 3869, 3890, 16 XII 2000, HMAS 76731, 81594; Jiangxi: Jinggangshan, alt. 800 m, on leaf sheath and culm of a large grass, W.Y. Zhuang & Z. Wang 1057, 23 X 1996, HMAS 81358. Notes: The fungus was originally discovered from Guangxi (Zhuang, 1998). Recent collections on lager graminaceous plants were obtained from Dawuling Mountains of Guangdong, Hainan Island, Laojunshan Mountains of Yunnan, and Jinggangshan Mountains of Jiangxi at elevations between 800 m and 1400 m. Careful examinations indicated that these collections are very similar in shape of apothecia, anatomical structure, and shape of asci and of ascospores, while dimensions varied. The ascospore length seems to be the most variable character. For example, they measured 43-54 x 1.8-2.2 um (Guangxi, holotype), 28-34 x 1.5-1.8 um and 30-45 x 1.5-2.2 um (Hainan, HMAS 76731, 81594), 28-43 x 1.5-1.9 um (Guangdong, HMAS 78159), and 26-37 x 1.8-2 um (Jiangxi, HMAS 81358). Diagnostic features: Apothecia 0.4-1.3 mm diam, hymenium white, cream to light yellow, receptacle white, surface hairy; hairs needle-like, 40-150(-230) um long, asci broadly clavate, pore J+ in Melzer’s reagent, 64-89 x 7-10 wm, ascospores subcylindrical with the basal end slightly narrower, 0-3-septate, 26-54 x 1.5-2.2 um, paraphyses narrowly lanceolate, 2-4 um at the widest, exceeding the asci by 6-25 um. Dicephalospora pinglongshanica W. Y. Zhuang, Fung. Divers. 3: 190, 1999. Specimen examined: China. Hainan: Lingshui, Diaoluoshan, 1050 m, on twigs, Y.H. Zhang, W.Y. Zhuang & Z.H. Yu 3817, 14 XII 2000, HMAS 81344. Notes: One additional collection (HMAS 81344) was found. Its apothecia are much larger than those in the type and up to 1.2 mm in diam; excipular tissues are highly gelatinous; asci are J- in Melzer’s reagent as shown in the holotype but longer and mostly 130-135 x 13-15 ym; and ascospores are larger and 24-32 x 5-7 um; which broaden the ranges displayed in the holotype from Guangxi at a lower elevation (Zhuang, 1999). Eleven Dicephalospora specimens were collected in our recent field work in the Hainan Island. Ten of them belong to D. rufocornea (Berk. & Broome) Spooner and one is D. pinglongshanica. The former is common throughout the tropical areas of China, where as the latter species is obviously rare but quite possibly occurs in other tropical regions as well. a2 Lambertella subrenispora Korf & W. Y. Zhuang, Mycotaxon 24: 367, 1985. Specimen examined: China. Hainan: Tongshi, Wuzhishan, alt 1600 m, on petiole of a rotten leaf, YH. Zhang, W.Y. Zhuang & Z.H. Yu 3882, 16 XII 2000, HMAS | 81357. Notes: The fungus was first recorded as Lambertella taxonomic sp. from India by Arendholz & Sharma (1984), then named and described based on Japanese material by Korf & Zhuang (1985). A recent collection (HMAS 81357) extends the known distribution to tropical China. Information about apothecial gross morphology in fresh condition and ascus iodine reaction of the species can now be updated. The apothecia are discoid, up to 2 mm in diam, and short- to long-stalked; when fresh the hymenium is light pink to reddish orange and receptacle is concolorous and with surface downy to finely scaly. The ascus pores are either obviously blue in Melzer’s reagent without KOH pretreatment as shown in the Indian and Chinese materials, or J- without KOH pretreatment and J+ after KOH treatment in the Japanese collection. Spore size seems stable in collections from different geographical regions, 11-12.5 x 4.5 um (India), 11-13 x 4.8-6.4 um (Japan), and 10-12 x 5 wm (China). Lasiobelonium guangxiense W. Y. Zhuang, Mycotaxon 69: 370, 1998. Specimens examined: China. Yunnan: Simao, alt. 1350 m, on small rotten twig, W.Y. Zhuang & Z.H. Yu 2899, 12 X 1999, HMAS 78058; Guangdong: Fengkai, Heishiding, alt. 300 m, on small twig, W.Y. Zhuang & Z.H. Yu.2850, 27 XII 1998, HMAS 81371. Notes: HMAS 78058, a recent collection on a small twig from Simao, Yunnan, has smaller apothecia 0.3-0.6 mm in diam, wider asci 105-115 x 17-19 um, and shorter and broader ascospores 20-24 x 6.5-8 um than those in the holotype from Damingshan Mountains, Guangxi (Zhuang, 1998). The apothecia of HMAS 81371 from Mt. Heishiding, Fengkai, Guangdong possess a greenish yellow hymenium, vinaceous receptacle, much longer asci 125-142 x 16.5-18 um, and longer ascospores 25-33 x 5.8-7.3 um. The dimensions of the species should be re-written as apothecia 0.3-0.9 mm in diam, asci 98-142 x 13-19 uum, and ascospores 20-33 x 5.5-8 um. The light brown, 4-celled ascospores constricted at septa is probably the most diagnostic feature of the fungus. Parachnopeziza guangxiensis W. Y. Zhuang & Korf, Mycotaxon 69:371, 1998. Specimen examined: China. Guangdong: Maoming, Dawuling, alt. 1400 m, on head of a graminaceous plant, W.Y. Zhuang & Z.H. Yu 2808, 22 XK 1998, HMAS 75554. Notes: The asci and ascospores of HMAS 75554 from Guangdong Province are much longer than those of the holotype (Zhuang, 1998), 74-97 x 9-11.5 um vs. 62-74 x 8.5-11 um and 30-42 x 2.6-3 um vs. 30-35 x 2.7-3 um, respectively. Rutstroemia dabaensis W. Y. Zhuang, Mycotaxon 61: 10, 1997. Specimen examined: China. Yunnan: Jizushan, on a tree, C.C. Chou 170, 18 IX 1938, HMAS 17083. 33 Notes: HMAS 17083 was collected sixty years ago in Yunnan Province and filed under Helotium sp. Its anatomic structure is typical Rutstroemia P. Karst. The small ascospore size recalls those of R. dabaensis known previously only from type locality, Wuxi, Sichuan Province (Zhuang, 1997). The Yunnan collection has larger apothecia, a thicker hymenium, and J- asci in Melzer’s reagent which are longer than those in the holotype and paratypes. The fungus might be common in southwestern China. The updated diagnosis of the fungus is as follows: Apothecia 1-5 mm diam, hymenium orange to yellow when fresh, stipitate, stipe base very dark; ectal excipulum, 25-50 um thick, tissues highly gelatinous, cells thick-walled and refractive; medullary excipulum 50-350 um thick; hymenium 83-100 um thick; asci either J+ or J- in Melzer’s reagent, 71-95 x 4-6.5 um; ascospores 5-8.5 x 2.5-3.5(-3.7) uum; on woody substrates. ACKNOWLEDGEMENTS The authors would like to express their deep thanks to Prof. R. P. Korf of Cornell University, USA for his critical review of the manuscript and valuable suggestions; and to all specimen collectors for making this study possible. REFERENCES Arendholz, W. R. & R. Sharma (1984) Observations on some Eastern Himalayan Helotiales. Mycotaxon 20: 633-680. Dennis, R. W. G. (1962) A reassessment of Belonidium Mont. & Dur. Persoonia 2: 171-191. Dennis, R. W. G. (1963) A _ redisposition of some fungi ascribed to the Hyaloscyphaceae. Kew Bull. 17: 319-379. Dennis, R. W. G. (1978) British Ascomycetes. ed. 2. J. Cramer. Vaduz. Dissing, H. & A. Raitviir (1974) Discomycetes of Middle Asia. III. Otideaceae, Helvellaceae, Morchellaceae and Sarcoscyphaceae from the Tien-shan Mountains. Eesti NSV Tead. Akad. Toim. Biol. 23(2): 104-110. Graddon, W. D. (1977) Some new discomycete species: 4. Trans. Brit. Mycol. Soc. 69: 255-273. Graddon, W. D. (1990) Some new discomycete species: 8. Mycol. Res. 94: 231-236. Korf, R. P. & W.-Y. Zhuang (1985) A synoptic key to the species of Lambertella (Sclerotiniaceae), with comments on a version prepared for Taxadat, Anderegg’s computer program. Mycotaxon 24: 361-386. Raitviir, A. (1970) Synopsis of the Hyaloscyphaceae. Scripta Mycol. 1: 1-115. Raitviir, A. (1978) The genus Lasiobelonium. Scripta Mycol. 8: 99-131. Sanchez, A. (1967) The sections Apostemium and Microstemium of the genus Vibrissea (Fungi). J. Agric. Univ. Puerto Rico 51(1): 79-93. Sharma, M. P. (1988) A new species of Psilachnum Héhn. from India. Nova Hedwigia 46: 369-372. Spooner, B. M. (1987) Helotiales of Australasia: Geoglossaceae, Orbiliaceae, Sclerotiniaceae, Hyaloscyphaceae. Biblioth. Mycol. 116: 1-711. 34 Svréek, M. (1992) New or less known Discomycetes. XXI. Ceska Mykol. 45: 134- 143. Zhuang, W.-Y. (1997) Fungal flora of the Daba Mountains: discomycetes. Mycotaxon 61: 3-12. Zhuang, W.-Y. (1998) Discomycetes of tropical China. III. Hyaloscyphaceous fungi from tropical Guangxi. Mycotaxon 69: 359-376. Zhuang, W.-Y. (1999) Discomycetes of tropical China. VI. Additional species from Guangxi. Fung. Divers. 3: 187-196. MY COTAXON Volume LXXXI, pp. 35-43 January-March 2002 RE-EXAMINATIONS OF HELOTIUM AND HYMENOSCYPHUS (HELOTIALES, HELOTIACEAE): SPECIMENS ON DEPOSIT IN HMAS * YAN-HUI ZHANG and WEN-YING ZHUANG Systematic Mycology and Lichenology Laboratory, Institute of Microbiology Chinese Academy of Sciences, Beijing 100080, China Abstract: Specimens deposited in HMAS under the generic names Helotium and Hymenoscyphus were re-examined. Twelve taxa of Hymenoscyphus were identified according to modern species concepts. Other discomycetes observed are members of Bisporella, Calycellina, Calycina, Ciboria, Crocicreas, Dicephalospora, and Lanzia. Hymenoscyphus sinicus is described as a new species. Three new records for China were reported. Key words: China, Discomycetes, Hymenoscyphus sinicus, Taxonomy As Donk pointed out in 1962, Helotium Fr. 1849 is a later homonym of Helotium Tode 1790. Dennis (1964) transferred a fair number of species assigned to Helotium Fr. to Hymenoscyphus Gray, now commonly accepted by mycologists around the world. Research of Hymenoscyphus in China began in the 1930’s when the generic name Helotium Fr. was used. Teng (1934) reported the first 5 species. Ou (1936) added 5 more taxa. Sixteen species of Helotium were recorded up to the early 1980’s (Teng, 2 1963.0) lal 9192. Zane.. 1983). Inedater studies, the generic name Hymenoscyphus has been accepted. Zhuang (1998) listed 15 taxa previously studied by Korf & Zhuang (1985), Zhuang & Korf (1989), Bi et al. (1990), Zhuang (1995), Wu et al. (1996), and Zhuang & Wang (1998). Recently, six species were found from the Changbai Mountains (Yu ef a/., 2000). This study is based on the dried herbarium specimens of Helotium and Hymenoscyphus on deposit in the Herbarium Mycologicum Instituti Microbiologici Academiae Sinicae (HMAS). Longitudinal sections were made with freezing microtome. Microscopic slides were mounted in cotton blue lacto-phenol and Melzer’s reagent. New records for China are marked with an asterisk (*) in this paper. " Author for correspondence. zhuangwy@sun.im.ac.cn 36 Species Accepted in Hymenoscyphus Hymenoscyphus caudatus (P. Karst.) Dennis, Persoonia 3: 76, 1964. Specimens examined: on leaf veins, Liuba, Shaanxi, alt. 950 m, 21 IX 1991, W.Y. Zhuang (852), HMAS 61875; on herbaceous plant part, Tianhuashan, Foping, Shaanxi, alt. 1300 m, 26 IX 1991, W.Y. Zhuang (887), HMAS 61876; on culms of monocotyledon, Bailang, Inner Mongolia, alt. 1350 m, 18 VIII 1991, W.Y. Zhuang (678), HMAS 69618. Notes: The Chinese collections fit the morphology of material from Europe and South America reported by White (1943), Dennis (1956), and Dumont (1982). The tropical American collections have larger asci [(90-)105-140(-150) x 8-12(-15) um] (Dumont, 1982). HMAS 69618 is different from the other two Chinese collections in slightly thickened (up to 2.5 um thick) and refractive cell walls in the ectal excipulum and 2-6-guttulate ascospores. * Hymenoscyphus cf. consobrinus (Boud.) Hengstm., Persoonia 12: 489, 1985. (& FA) FAH AR bal : Apothecia discoid, 2-4.5 mm in diam, long stipitate, stipe up to 8 mm long when dry, hymenium yellow to orange-red when fresh, drying orange-red to yellow, receptacle yellow; stipe pale to light yellow, with whitish hyphae at the base. Ectal excipulum of textura prismatica, 35-63 um thick, cells 6-20 x 5-13 um; covering layer of one to several cell layers. Medullary excipulum of textura intricata, hyphae 1-4 um wide. Asci cylindric-clavate, 8-spored, arising from croziers, J+ in Melzer’s reagent, as two blue lines, 98-120 x 8-10 um. Ascospores ellipsoid, very slightly narrower at one end, with 2-3 large guttules, non-septate, irregularly uniseriate to biseriate, 12-18 x 4-5(-6) um. Paraphyses filiform, septate, 1.5-2.5 um thick at apex. Specimens examined: on stem base of Rhododendron sp., Xiangcheng, Sichuan, alt. 4500 m, 30 VI 1998, Z. Wang (0002), HMAS 75931; on dead wood, Daocheng, Sichuan, alt. 4000 m, 2 VII 1998, Z. Wang (0015), HMAS 75934. Notes: The two collections differ from Hymenoscyphus consobrinus (Lizon, 1992) by the larger asci [98-120 x 8-10 um vs. 85-95 x 6.5-7.5(-9) um] and broader, non-septate, and ellipsoid instead of fusoid ascospores. Hymenoscyphus epiphyllus (Pers.: Fr.) Rehm in Kauffman, Pap. Michigan. Acad. Sci. Dale 1928) 1929. Specimen examined: on fallen leaves of Betula sp., Arxan, Inner Mongolia, alt. 1210 m, 14 VIII 1991, W.Y. Zhuang (622), HMAS 61894. Hymenoscyphus fructigenus (Bull.: Fr.) Gray, Nat. Arrang. Brit. Pl. 1: 673, 1821. Specimens examined: on rotten seeds of Quercus sp., Baihuashan, Beijing, alt. 1400 m, 19 IX 1995, W.Y. Zhuang & Z. Wang (1233), HMAS 71816; on seeds of Quercus sp., Baihuashan, Beijing, alt. 1400 m, 20 IX 1995, W.Y. Zhuang & Z. Wang (1259), HMAS 71817; on rotten seeds of Quercus sp., Donglingshan, Beijing, 19 VIII 1998, Z. Wang (0264), HMAS 75893. Notes: The collections from Beijing match most of the features of this species described by Dennis (1956, 1968) except that the stipe and asci are shorter. HMAS 75877 on rotten wood from Donglingshan, Beijing has shorter asci and ascospores 37 and somewhat thick-walled cells in the ectal excipulum. We treated it as Hymenoscyphus cf. fructigenus. Hymenoscyphus fucatus (W. Phillips) Baral & Hengstm., Hengstmengel, Persoonia 16: 193, 1996. thE RAR A Specimens examined: on herbaceous stem, Tewo, Gansu, alt. 2600 m, 11 IX 1992, W.Y. Zhuang (1017), HMAS 61883; on rotten twigs, Kangding, Sichuan, alt. 3500 m, 28 VIII 1997, Z. Wang (2178), HMAS 75902. Notes: The Chinese collections has slightly shorter ascospores [21-28.5 x 4-6 um vs. 24-34 x 5-6.8 um and (23-)25-31(-35) um] (White, 1944; Dennis 1956; Hengstmengel, 1996). The fungus was previously recorded as Hymenoscyphus scutulus (Pers.: Fr.) W. Phillips var. fucatus W. Phillips (Zhuang, 1995). Hengstmengel (1996) raised the variety to species level. Hymenoscyphus immutabilis (Fuckel) Dennis, Persoonia 3: 76, 1964. Specimens examined: on rotten leaves of Populus sp., Baihuashan, Beijing, 20 IX 1995, W.Y. Zhuang & Z. Wang (1251, 1273, 1274), HMAS 71806, 71808, 71809; on rotten leaves, Baihuashan, alt. 1400 m, Beijing, 20 IX 1995, W.Y. Zhuang & J.Y. Zhuang (1275), HMAS 71807; on rotten leaves, Baihuashan, Beijing, 19 IX 1995, W.Y. Zhuang & Z. Wang (1230), HMAS 71810; on rotten leaves, Baihuashan, alt. 1400 m, Beijing, 19 IX 1995, W.Y. Zhuang & C.L. Fu (1265), HMAS 71811. Hymenoscyphus lasiopodium (Pat.) Dennis, Persoonia 2: 190, 1962. Specimens examined: on dead root of Carex sp., Tanzhesi, Beijing, 8 X 1988, R.P. Korf & W.Y. Zhuang (465), HMAS 57692; on dead root of Carex sp., Baihuashan, Beijing, alt. 1200 m, 18 IX 1995, W.Y. Zhuang & Z. Wang (1204, 1205, 1196, 1195), HMAS 71803, 71804, 71805, 71819; on dead root of Carex sp., Baihuashan, Beijing, alt. 1400 m, 19 [IX 1995, Z. Wang & W.Y. Zhuang (1221), HMAS 71820; on dead root of Carex sp., Baihuashan, Beijing, alt. 1400-1600 m, 20 IX 1995, W.Y. Zhuang & Z. Wang (1255, 1271), HMAS 71821, 71822; on dead root of Carex sp., Donglingshan, Beijing, 18 VIII 1998, Z. Wang (0242), HMAS 75878; on dead root of Carex sp., Donglingshan, Beijing, alt. 1100 m, 19 VIII 1998, Z. Wang (0261), HMAS 75880. Notes: Compared with the South American material (Dumont, 1982), the Chinese collections occurred on dead roots of Carex sp. from Beijing and have much longer asci [120-180(-200) x 7.5-10(-12) um vs. (90-)110-115(-130) x 6-8(-10) um] and shorter and broader cells in the ectal excipulum (5-33 x 4-17 um vs. 10-40 x 8-15 pm). HMAS 75878 and HMAS 75880 have non-septate ascospores that are possibly young. *Hymenoscyphus menthae wh Phillips) Baral in Baral & een ae BeihieZ: Mykol. 6: 131, 1985. #8 fay Heat Dried apothecia discoid, oe 0.1-0.3 mm in diam, stipe 0.5-0.8 mm long, hymenium light yellow to light brown, receptacle light yellow to light brown, stipe cream to light yellow, red-brown at the base. Ectal excipulum of textura prismatica, 18-40 um thick, cells 3-17 x 2-8 um; covering layer of 1-3 cell layers. Medullary excipulum of textura intricate to textura porrecta, hyphae 1-4 um wide. Hymenium 38 60-105 um thick. Asci cylindric-clavate, 8-spored, J+ in Melzer’s reagent, as two blue lines, 67-88(-112) x 6-8 um. Ascospores elliptic-fusoid, with 1-3 large guttules, non-septate, irregularly biseriate, 14-18 x 3.5-4.3(-5) um. Paraphyses filiform, 1-3 pm thick at apex. Specimens examined: on a stem, Qingchengshan, Sichuan, 18 IX 1981, RY. Zheng & R.P. Korf (2388), HMAS 51841; on stem of Polygonum cuspidatum Sieb. & Zucc., 19 IX 1981, R.Y. Zheng & R.P. Korf (2413), HMAS 51842. Notes: The Chinese collections were previously reported as Hymenoscyphus . scutulus var. solani (P. Karst.) S. Ahmad (Korf & Zhuang, 1985). According to Hengstmengel (1996), H. menthae is the correct name for the fungus. *Hymenoscyphus cf. populneus (Velen.) Svréek, Sborn. Nar. Muz. Praze 40B(3-4): 170, (1984) 1985. (BIR) aia A Apothecia discoid, margin lower than hymenium, subsessile to short-stipitate, 1.5 mm in diam, hymenium yellow when fresh, light yellow to red brown when dry, drying receptacle whitish. Ectal excipulum of textura prismatica, 17-65 wm thick, with short hairs on receptacle surface, cells 13-18 x 3-5.5 um. Medullary excipulum of textura intricata, 12-113 um thick, hyphae 1-6 um wide. Hymenium 62-88 um thick. Subhymenium 0-30 pm thick. Asci cylindric-clavate, 8-spored, J+ in Melzer’s reagent, as two blue lines, 46-55 x 5-5.5 um. Ascospores ellipsoid to broadly fusoid, with 2(-4) polar guttules, non-septate, irregularly uniseriate to biseriate, 5.5-8 x 1.7-2.3 um. Paraphyses filiform, 1-2 um thick at apex. Specimen examined: on rotten leaves of Populus sp., Donglingshan, Beijing, alt. 1100 m, 18 VIII 1998, Z. Wang (0240), HMAS 75881. Notes: Compared with Helotium populneum Velen. (Velenovsky, 1947) from the former Czechoslovakia, our fungus has shorter ascospores and short-stipitate, yellow instead of pure white apothecia. Hymenoscyphus sclerogenus (Berk. & M. A. Curtis) Dennis, Persoonia 2: 190, 1962. Specimen examined: on cane of Rubus sp., E’mei Mountain, 25 VI 1983, W.Y. Zhuang (55), HMAS 45092. Hymenoscyphus scutulus (Pers.: Fr.) W. Phillips, British Discom. p. 137, 1887. Specimens examined: on herbaceous stem, Bailang, Inner Mongolia, alt. 1350 m, 18 VIII 1991, W.Y. Zhuang (679, 681), HMAS 61879, 61880; on herbaceous stems, Tewo, Gansu, alt. 2600 m, 11 IX 1992, W.Y. Zhuang (1018), HMAS 61881; on herbaceous stem, Huangshan, Anhui, alt. 1300-1700 m, 27 IX 1993, Y.R. Lin, Y. Wang, W.Y. Zhuang, S.M. Yu & W.J. Wu (1119), HMAS 61882. Hymenoscyphus sinicus W.Y. Zhuang & Y.H. Zhang, sp. nov. F4EfRAt EI (Figs. 1-3) Ab Hymenoscypho caudato apotheciis magnis, 1-6 mm diam; ascis longis, 125-175 x 8-10(-12.5) ym, ascosporis 0-1-septatis, (16-)18-26 x (4.1-)4.5-5.5(-6) um differt. Apothecia discoid, 2-6 mm in diam, stipitate, stipe 1-7.5 mm long when dry, hymenium yellow to dirty yellow when fresh, yellow to brown when dry, receptacle ay whitish to yellow-orange when dry. Ectal excipulum of textura prismatica, 37-108 um thick, cells 5-33 x 6-15 um, cell walls slightly thickened; covering layer of 0-3 hyphal layers. Medullary excipulum of textura intricata, hyphae hyaline, 2-5 um wide. Hymenium (125-)210-275 um thick. Asci cylindric-clavate, 8-spored, arising from croziers, J+ in Melzer’s reagent, pore walls bluing as two lines, 125-175 x 8-10(-12.5) um. Ascospores fusoid to scutuloid, with 2-4 large guttules or multiguttulate, 0-1(-2)-septate, irregularly uniseriate to biseriate, (16-)18-26 x (4.1-)4.5-5.5(-6) um. Paraphyses filiform, 1-3 um wide. Holotype: on rotten twigs, Baihuashan, Beijing, China, alt. 1400 m, 19 IX 1995, W.Y. Zhuang & Z. Wang (1213), HMAS 71818. Paratypes: on ?root, Zhouqu, Gansu, China, alt. 2400 m, 3 IX 1992, W.Y. Zhuang (929), HMAS 69617; on rotten wood, Donglingshan, Beijing, China, alt. 1100 m, 20 VIII 1998, Z. Wang (0279), HMAS 75876. Notes: The new species is similar to Hymenoscyphus caudatus in ascospore shape and size but differs obviously from the latter in large cells and somewhat gelatinized tissues in the ectal excipulum, longer asci (85-95 x 8-10 um vs. 125-175 x 8-10(-12.5) um), much larger apothecia (1-6 mm diam vs. 0.75-1 mm diam), and occurrence on woody substrates instead of leaf parts or herbaceous stems (Dennis, 1956; Dumont, 1982). Several other species possessing scutuloid ascospores in the genera Hymenoscyphus, Lambertella Héhn., and Lambertellinia Korf & Lizon (Korf & Lizon, 1994; Hengstmengel, 1996; Zhuang, 1999) were also compared. H. sinicus is distinctive in spore morphology, excipular structure, and apothecial dimension. Figs. 1-3. Hymenoscyphus sinicus: 1. Ectal excipular structure. 2. An ascus and three ascospores. 3. Ascospores in a higher magnification. All from holotype. 1-2 x 315, 3 scale = 10 um. 40 Species Belonging to Other Genera Bisporella citrina (Batsch: Fr.) Korf & S.E. Carp., Mycotaxon 1: 58, 1974. Specimens examined: On rotten wood, Jiabigou, Miyaluo, Sichuan, alt. 2850 m, 23 IX 1960, C.M. Wang, Y.X. Han & Q.M. Ma (1317), HMAS 30480; on mossy wood, Hebei, S.C. Teng (489) HMAS 33617; on dead twigs in bamboo forest, Jinzhongshan, Longlin, Guangxi, alt. 1700 m, 20 X 1957, L.W. Xu (125), HMAS 33618; on dead twigs, Lingle, Guangxi, alt. 1000 m, 11 X 1957, L.W. Xu (1153), HMAS 33619; on rotten wood, Laoshan, Lingle, Guangxi, alt. 1300 m, 15 XII 1957, L.W. Xu (368), HMAS 33717. Bisporella cf. citrina Specimens examined: on mossy wood, Miyaluo, Sichuan, 29 VII 1958, Y.L. Hu (98), HMAS 24098; on rotten wood of Larix. sp., Nantianmen, Taibaishan, Shaanxi, alt. 3000 m, 2 VIII 1963, Q.M. Ma & Y.C. Zong (2723), HMAS 33516; on rotten wood of broad-leaved trees, Xiaoxihe, Ledu, Qinghai, alt 2850 m, 10 IX 1959, J.C. Xing & Q.M. Ma (1569), HMAS 33623; on rotten wood, Wulingshan, Hebei, 9 IX 1958, K.G. Ji (176), HMAS 33624; on mossy wood in bamboo forest, Jinzhongshan, Longlin, Guangxi, alt. 1700 m, 20 X 1957, L.W. Xu (107), HMAS 33740; on rotten wood, Jinzhongshan, Longlin, Guangxi, alt. 1720 m, 25 X 1957, L.W. Xu (401), HMAS 33741; on rotten wood, Jinzhongshan, Longlin, Guangxi, alt. 1600 m, 19 X 1957, L.W. Xu (048), HMAS 33744. Notes: The above collections have narrower asci and shorter ascospores than those of Bisporella citrina (Dennis, 1956). A Cystodendron stage was not found. Calycellina populina (Fuckel) Héhn., Akad. Wiss. Wien Sitzungsber., Math. -Naturwiss. K1., Abt.1, 127: 601, 1918. Specimen examined: on rotten leaves, Huang’antuo, Baihuashan, Beijing, 26 IX 1956, Y.C. Wang, L.W. Xu & S.J. Han (223), HMAS 19697. | Calycina herbarum (Pers.: Fr.) Gray, Nat. arr. Brit. pl. 1: 669, 1821. Specimens examined: on herbaceous stem, Yinshengou, Ledu, Qinghai, alt. 2600 m, 9 IX 1959, J.C. Xing & Q.M. Ma (1541), HMAS 31241; on herbaceous stem, Changbaishan, Antu, Jilin, alt. 1280 m, 20 VIII 1960, Y.C. Xiang, J.R. Chen & F.S. Yuan (880), HMAS 30751. Ciboria peckiana (Cooke) Korf, Phytologia 21: 203, 1971. Specimen examined: on rotten wood, Wuchongxiang, Longlin, Guangxi, alt. 1000 m, 31 X 1957, L.W. Xu (638), HMAS 33835. Notes: Baral et al. (1999) assigned this fungus to TJatraea, a genus in the Leotiaceae. We certainly notice the similarities between C. peckiana and the type species of Zatraea but would like to wait for more evidence to support this placement. 4] Dicephalospora rufocornea (Berk. & Broome) Spooner, Biblioth. Mycol. 116: 272, 1987. Specimens examined: on twigs, Dingan, Hainan, IX 1934, S.Q Deng (4306), HMAS 07204; on bark, Xishan, Kunming, Yunnan, 9 VIII 1938, C.C. Cheo (83), HMAS 17063; ibid., 28 VIII 1938, S.Z. Zhao, HMAS 17064; ibid., 9 VII 1938, F.L. Tai (86), HMAS 17067; on twigs, Xishan, Kunming, Yunnan, C.C. Cheo (34), HMAS 17068; ibid., 15 VII 1938, C.C. Cheo (39), HMAS 17069; ibid. 16 VII 1938, C.C. Cheo (50, 46), HMAS 17070, 17074; ibid., 18 VII 1938, C.C. Cheo (55), HMAS 17072; ibid., 9 VIII 1938, C.C. Cheo (81), HMAS 17073; on rotten wood, Xishan, Kunming, Yunnan, 14 VII 1938, C.C. Cheo (26), HMAS 17075; ibid., 27 VII 1941, C.C. Cheo (196), HMAS 17080; ibid., 22 VII 1942, W.F. Chiu (202), HMAS 17081; on small twigs, Wenshan, Yunnan, 17 VIII 1938, Q.H. Wang (97), HMAS 17082; on twigs, Xishan, Kunming, Yunnan, 28 VIII 1938, S.Z. Zhao (123), HMAS 17146; on fallen twig of broad-leaf forest, Huangshan, Anhui, 24 VIII 1957, S.C. Teng, (5052), HMAS 20312; on twigs, Huangshan, Anhui, 30 VIII 1957, S.C. Teng (5158), HMAS 20313; on fallen twigs, Tianmushan, Zhejiang, 9 IX 1957, S.C. Teng (5417), HMAS 20314; on rotten wood, Daji, Nanjing, Fujian, 11 VI 1958, S.C. Teng (5775), HMAS 24099; ibid., Diaoluoshan, Hainan, alt. 350 m, 26 IX 1958, Q.Y. Zheng (184), HMAS 26780; ibid., Nankengdaling, Nanjing, Fujian, 13 VI 1958, S.C. Teng (5840), HMAS 26792; on rotten twigs, Daqingshan, Longjin, Guangxi, 9 IX 1958, G.Z. Jiang (423), HMAS 26793; ibid., Qianlingshan, Guiyang, Guizhou, alt. 1400 m, 13 VIII 1958, Q.Z. Wang (94), HMAS 26794; ibid., Ceheng, Guizhou, alt. 800 m, 14 X 1958, Q.Z. Wang (549), HMAS 27120; on rotten wood, Nankengdaji, Nanjing, Fujian, 14 VI 1958, S.C. Teng (5880), HMAS 27121; on rotten twigs, Guangian, Pucheng, Fujian, alt. 500 m, 28 VIII 1960, Q.Z. Wang & W.L. Yuan & G. He (762), HMAS 28770; ibid., Xipo, Baoting, Hainan, 31 VII 1960, J.H. Yu & R. Liu (1885), HMAS 29387; on twigs, Tianshidong, Qingchengshan, Sichuan, alt.1000 m, 15 VIII 1960, C.M. Wang & Y.X. Han (568), HMAS 30481; ibid., 16 VIII 1960, C.M. Wang & Y.X. Han & Q.M. Ma (586), HMAS 31244; ibid., Dashahe Nature Reserve, Daozhen, Guizhou, 20 VII 1988, Y. Li & Y.C. Zong & J.Z. Ying (500), HMAS 57797; ibid., 19 VII 1988, Y. Li & Y.C. Zong & J.Z. Ying (461), HMAS 58264; ibid., Yunnan, VIII 1992, S.X. Sun (259h), HMAS 61347. Notes: This is a very common species in China. Most of the collections filed under Helotium serotinum (Pers.) Fr. in HMAS turned out to be Dicephalospora rufocornea. Lanzia huangshanica W.Y. Zhuang & Korf forma aurantiaca W.Y. Zhuang, Mycosystema 7: 14, 1995. Specimen examined: on leaf veins, Yanfang, Liuba, Shaanxi, alt. 950 m, 21 IX 1991, W.Y. Zhuang (850), HMAS 61874. Lanzia serotina (Pers.: Fr.) Korf & W.Y. Zhuang, Mycotaxon 22: 506, 1985. Specimen examined: on twigs, Haopingsi, Taibaishan, Shaanxi, alt. 1200 m, 14 V 1963, Q.M. Ma & Y.C. Zong (2230), HMAS 33625. Notes: The stipe base of this collection seems not dark. 42 Acknowledgements The authors would like to express their deep thanks to Prof. R.P. Korf of Cornell University, USA, for his critical review of the manuscript, taxonomic insight, and kindly serving as pre-submission reviewer, and to all collectors of specimens for this study. This project is supported by the National Natural Science Foundation of China and Funds of the Knowledge Innovation Program of the Chinese Academy of Sciences. References Baral, H.-O, R.G. Marquez, I. Krisai-Greilhuber, N. Matoéec & J.T. Palmer (1999) Tatraea dumbirensis, new records of a rare leotialean discomycete in Europe. Osterr. Z. Pilzk. 8: 71-82. Bi, Z.S., G.Y. Zheng, T.H. Li. & Y.Z. Wang (1990) Macrofungus Flora of the Mountainous District of North Guangdong. Guangdong Science and Technology Press. Guangzhou. 1-450. (in Chinese) Carpenter, S.E. (1981) Monograph of Crocicreas (Ascomycetes, Helotiales, Leotiaceae). Mem. New York Bot. Gard. 33: 1-290. Dennis, R.W.G. (1956) A revision of the British Helotiaceae in the Herbarium of the Royal Botanic Gardens, Kew, with notes on related European species. Mycol. Pap. 62: 1-216. Dennis, R.W.G. (1964) Remarks on the genus Hymenoscyphus S.F. Gray, with observation on sundry species referred by Saccardo and others to the genera Helotium, Pezizella or Phialea. Persoonia 3: 29-80 (1963). Dennis, R.W.G. (1968) British Ascomycetes. ed. 2. J. Cramer. Stuttgart. 1-455. Donk, M.A. (1962) The generic names proposed for Agaricaceae. Beih. Nova Hedwigia 5: 1-320. Dumont, K.P. & S.E. Carpenter (1982) Los hongos de Colombia -- VII: Leotiaceae -- IV: Hymenoscyphus caudatus and related species from Colombia and adjacent regions. Caldasia 13 (64): 567-602. Hengstmengel, J. (1996) Notes on Hymenoscyphus — II. On three non-fructicolous species of the ‘fructigenus-group’ with crozier. Persoonia 16: 191-207. Korf, R.P. & P. Lizon (1994) Lambertellinia scutuloides (Sclerotiniaceae), a new genus and species for a discomycete confused with Hymenoscyphus caudatus. Mycotaxon 50: 167-174. Korf, R.P. & W.Y. Zhuang (1985) Some new species and new records of discomycetes in China. Mycotaxon 22: 483-514. Lizon, P. (1992) The genus Hymenoscyphus (Helotiales) in Slovakia, Czechoslovakia. Mycotaxon 65: 1-59. Ou, S.H. (1936) Additional fungi from China. Sinensia 7: 668-685. Tai, F.L. (1979) Sylloge Fungorum Sinicorum. Science Press. Beijing. 1-1527. (in Chinese) Teng, S.C. (1934) Notes on discomycetes from China. Sinensia 5: 431-465. Teng, S.C. (1996) (Korf, R.P. ed.) Fungi of China. Mycotaxon Ltd. Ithaca. 1-586. Velenovsky, J. (1947) Novitates mycologicae novissimae. Opera Bot. Cech. Pragae. 4: 1-158. 43 White, W.L. (1943) Studies in the genus Helotium, III. History and diagnosis of certain European and North American foliicolous species. Farlowia 1: 135-170. White, W.L. (1944) Studies in the genus Helotium, IV. Some miscellaneous species. Farlowia 1: 599-617. Wu, S.H., Y.Z. Wang & W.N. Chow (1996) Catalogue of fungal Specimens and Cultures of NMNS. Natl. Mus. Nat. Sci. Taichung. 1-140. (in Chinese) Yu, Z.H., W.Y. Zhuang, S.L. Chen & C. Decock (2000) Preliminary survey of discomycetes from the Changbai Mountains, China. Mycotaxon 75: 395-408. Zang, M. (1983) Discomycetes. pp 18-29. In Anon. Fungi of Xizang. Science Press. Beijing. 1-226. (in Chinese) Zhuang, W.Y. (1995) Some new species and new records of discomycetes in China. V. Mycotaxon 56: 31-40. Zhuang, W.Y. (1998) A list of discomycetes in China. Mycotaxon 67: 365-390. Zhuang, W.Y. (1999) Discomycetes of tropical China. VI. Additional species from Guangxi. Fung. Divers. 3: 187-196. Zhuang, W.Y. & R.P. Korf (1989) Some new species and new records of discomycetes in China. III. Mycotaxon 35: 297-312. Zhuang W.Y. & Z. Wang (1998) Some new species and new records of discomycetes in China. VIII. Mycotaxon 66: 429-438. cea im Net gh RW oie Bho hereon: were nip ae nena oataes ainea ‘blaine : bib as mea Homi esi a ia Ba ao 7 Any f , y ae As 7 y 72) i a mee 4 ve iy sah basioen ee vein adi ; f i ; Soh nponyn enti: mie eobospnO aR worcnetaninian le ue A A, Sire ig. ersonld alpha aclas th Lon hi Cn ay eue loi elorrenne te he the MeINY DN Jay Si dda MED, minh WN rie BLA ele rca Meek ni Pui \ ie as ed inl at % A} a i Dena a yy! sy Os, anes * poy 7 f ee is, ne ne on aw asi: ce P tay: %, sikh tha) ry ‘shy ‘the ivan iii ; ; Bi ates an ape sib io RE ae ee | sed weit Pili a ma ie heh aha rite as.) boat en . ii Pak ibe Ni Hild eat ibis Opens 4a ie a asl ‘eal ro wi +i pak me "i iM Mh, ‘aan eR pins a i pine, ites fom site pheye Py, FP aed ue ah as Gre ae rae Heh Cael shk aba ie, jt Me hy (Riana ‘I i Mies ce x4 Piss f Ni j | fie 7 {5 “4 ] Ge me , { Bit : ; ; nA Ms aN Ae es Ws fi Lis ee I Be ik ha a es fs ye re Rete ey a wf Oa ace i Oy i J og a hi tp) A ig bau Ay nS iy ut it Oi ai OH Mi ag +l ais ear ie . i ces | « ae “Pree Va # ¥, M4 a Na 1 hit a ah te ie ne gh OR ti jean way ie: oa eel 7 t ; ey ae aint a He’ a aE hie i ‘ iM y a a wei i Ye ae inh 2 us oe i” pay wh pe ey 4 Pere fi atin byte ld ee a . ines on ri Roni es . ‘s A it i A é ide we ies Ae Pitot eer’ " a ie si : Mi ihe “ Maal ‘th vex gs Oi iF gel 4 4 r f _ A ? , ; 4 ms) 4 veel i 7 | ‘ “ sien ve ~ Hey, F)) en ie oe m ie mF iG. Danii on ¥ ia ae aa wee ‘era mai Higa ehhae 40%) Ri ngs shied nde Ai ey Vin) i Pee ie ius ny Nie roa Tyee ke ean igh eee: ca i mete Le ae ‘a a is a p net Pi haan Se re sud en 1a dagiah pel van io Tana nae | Laknnnse anit Hee Ne oh Ways Onis shh Paani rh A ih he Pang » Wy, tone a” bead art ca meri acu se veh ‘ things ih ba erbiae Dake us ah no wot: Abas tie a spaces vend vy ba a ole mah h dayitis Tagine Pn Shaul vlad! a Be ea bedi 2 Khe hs id Shlie me ity tiem sso iene Si ihe) oe Bat MYCOTAXON Volume LXXXI, pp. 45-50 January-March 2002 TWO NEW DERMATEACEOUS GENERA WITH LANCEOLATE PARAPHYSES AIN RAITVIIR Institute of Zoology and Botany Riia Street 181, EE-51014 Tartu, Estonia e-mail: ain@zbi.ee ABSTRACT: The taxonomic position of two species currently assigned to the Hyaloscyphaceae, Dasyscyphus involucratus and Trichopezizella myricae is reassessed, two new Dermateaceous genera Involucroscypha and Obscurodiscus are erected to accommodate them and the corresponding new combinations /nvolucroscypha involucrata and Obscurodiscus myricae are proposed. Key words: taxonomy, Hyaloscyphaceae, Dermateaceae, new genera. INTRODUCTION There are two very rare and curious, dark, hairy Helotiales with lanceolate paraphyses, Dasyscyphus involucratus B. Erikss. and Trichopezizella myricae (P. Karst.) K. & L. Holm, currently assigned to the Hyaloscyphaceae which, however, do not fit the concept of genera they are placed in and the family as whole, too. Recently the author had an opportunity to study representative material of both species. In the result of this study two new genera of the Dermateaceae are proposed to accommodate those species. 46 MATERIAL AND METHODS. The freshly collected living material was mounted in tap water, the dry herbarium material was resoaked in 3% aqueous solution of KOH. The Meltzer’s reagent (MLZ), Congo Red (CR) and Cotton Blue (CB) were used for histochemical reactions. The mounts were examined by a Nikon Labophot- 2 microscope equipped with a drawing tube. The size of microscopical structures was measured in 3% KOH using a 100x oil immersion objective lens if not stated otherwise. TAXONOMY Involucroscypha Raitv., genus novum. Apothecia superficilia, breviter stipitata, minuta, primo obovoidea, cupulata, opaco nigra, extus furfuracea. Excipulum ectale ex cellulis globosis tenuiparietalis fuscis compositur, ad margine cum processibus fuscis tenuiparietalibus constrictoseptatis. Medulla ex cellulis globosis crassiter hyalinotunicatis amyloideis. Pili clavati, tenuiter vel subcrassiter brunneotunicati, verrucis hyalinis crustiforme confluentibus obtectis. Asci clavati, octospori, poro iodo caerulescentia praediti. Sporae hyalinae, aseptatae. Paraphyses lanceolatae, ascos longe superantes. Typus generis: Dasyscyphus involucratus B. Erikss., Acta Univ. Ups. Symb. Bot. Ups. 19 (4): 16, 1970. Apothecia superficial, shortly stipitate, often involucrate, a new apothecium proliferating from the centre of the old apothecium shell which persists as an involucrum, externally hairy and medium brown when fresh, dark brown when dry, with whitish to greyish hymenium. Ectal excipulum composed of textura globulosa, cells with thin, pale, yellowish walls. Externally the excipulum is densely covered with fascicles of adpressed brown hairs. Hairs clavate, arising from a wide basal cells at the flanks, narrowly rooting at the margin, with brown to dark brown slightly thickened walls, externally covered with a hyaline, strongly refracting, warty crust. Medulla composed of textura globulosa, cells hyaline, thick-walled, dextrinoid in MLZ. Asci cylindrical-clavate, with a short distinct stipe, 8-spored, apical pore blue in MLZ. Spores biseriate or obliquely uniseriate, hyaline, aseptate. Paraphyses lanceolatae, exceeding the asci. Etymology: referring to the peculiar growth habit of the apothecia. A single known species. 47 Involucroscypha involucrata (B. Erikss.) Raitv. comb. nov. Basionymum: Dasyscyphus involucratus B. Erikss., Acta Univ. Ups. Symb. Bot. Ups. 19 (4): 16, 1970. Figs. 1-4. Apothecia superficial, shortly stipitate, 0.3-0.6 mm diam., often involucrate, a new apothecium proliferating from the centre of the old apothecium shell, externally hairy and medium brown when fresh, dark brown when dry, with whitish to greyish hymenium. Ectal excipulum composed of textura globulosa, cells 5-8 um diam., with thin, pale, yellowish walls. Externally the excipulum is densely covered with fascicles of adpressed brown hairs. Medulla composed of textura globulosa, cells hyaline, thick-walled, dextrinoid in MLZ, 5-8 pm diam. Hairs clavate, arising from a wide basal cells at the flanks, narrowly rooting at the margin, 35-60 x 4-6 um, apically swollen up to 8 ym, with brown to dark brown slightly thickened walls, externally covered with a hyaline, strongly refracting, warty crust. Asci arising from croziers, cylindrical-clavate, with a short distinct stipe, 8-spored, 40-55 x 5-7 um, apical pore blue in MLZ. Spores biseriate or obliquely uniseriate, ellipsoid-fusoid, often slightly clavate and inequilateral to bemt,hyaline, aseptate, with 2 big polar lipid globules, 7-12 x 2-3 wm. Paraphyses lanceolatae, often 1-septate, without inclusions, 55-70 x 3-4 wm, exceeding the asci by up to 15 um. On dead stems of Ericaceae. Specimens examined: Austria, Kartnen: Koralpe,Pomseben,oberhalb des Wassenfalls, bei 1650 m, on dead stems of Vaccinium vitis-idaea, 5. VIII 1975, P. Remler (GZU, s.n.) Finland, Edela-Hame, Lammi Biological Station, on dead stems of Calluna vulgaris, 12. IX 2001, K. Leenurm (TAA 165831 This species with a curious habit and structure was described as a Dasyscyphus by Eriksson (1970) and treated under the same name by Remler (1979), too. But, in spite of its lanceolate paraphyses and rough “hairs”, it is not a Lachnum (=Dasyscyphus) and does not belong to the Hyaloscyphaceae at all. Its ectal excipulum, hidden under dense cover of adpressed hairs, is composed of textura globulosa and the hairs are different from typical hairs of the Hyaloscyphaceae. Its asci are different from asci found in the Hyaloscyphaceae but resemble closely the asci of a Dermateaceous genus Coronellaria which has also lanceolate paraphyses. The dense hair cover and involucrate growth of apothecia distinguish it clearly from the latter genus. But the involucrate growth of new apothecia from the shells of old apothecia can be sometimes observed in some other members of the Dermateaceae. The author has seen it in Tapesia livido-fusca (Fr.) Rehm and in some species of Mollisia. A characteristic feature of J. involucrata is dextrinoid medulla, Figs. 1-4. Involucroscypha involucrata. 1. A paraphysis and two asci; 2. Marginal hairs. 3. Spores; 4. Detail of ectal excipulum. Figs. 5-7. Obscurodiscus myricae. 5. Asciand paraphyses; 6. Spores; 7. Ectal excipulum and marginal hairs. 49 overlooked by the earlier authors. The thick-walled cells of medulla turn beautiful light reddish brown, sometimes with rose-coloured tint in MLZ. Obscurodiscus Raitv., genus novum. Apothecia superficilia, sessilia, muinuta, primo obovoidea, subsphaeroidea vel urceolata, dein cupulata, opaco nigra, extus furfuracea. Excipulum ectale ex cellulis globosis tenuiparietalis fuscis compositur, ad margine cum processibus fuscis tenuiparietalibus constrictoseptatis. Asci clavati, octospori, poro iodo caerulescentia praediti. Sporae hyalinae, aseptatae. Paraphyses lanceolatae, ascos longe superantes. Typus generis: Godronia myricae P. Karst., Hedwigia 23: 5, 1884. Apothecia superficial, sessile, minute., at first obovoid, subspherical or urn-shaped, later deeply cupulate or cupulate, opaque black when fresh and dry, externally furfuraceous. Ectal excipulum composed of textura globulosa, cells with firm, dark smoky brown walls, turning at the margin into hair-like hyphal processes with constricted septa and dark smoky brown, thin to slightly thickened walls. Asci cylindrical-clavate, 8-spored, apical pore deeply blue in MLZ. Spores hyaline, aseptate. Paraphyses lanceolate, hyaline, without inclusions, exceeding the asci. Etymology: referring to the dark colour of apothecia. A single known species. Obscurodiscus myricae (P. Karst.) Raitv. comb. nov. Basionymum: Godronia myricae P. Karst., Hedwigia 23: 5, 1884. Syn.: Trichopezizella myricae (P. Karst.) K. & L. Holm, Nord. J. Bot. 11: 678, oot Figs. 5-7. Apothecia superficial, sessile, 0.2-0.4 mm diam., at first obovoid, subspherical or urn-shaped, later deeply cupulate or cupulate, opaque black when fresh and dry, externally furfuraceous. Ectal excipulum composed of textura globulosa, cells with firm, dark smoky brown walls, 5-8 um diam., turning at the margin into hair-like chains of cells, composed of 2-4 cells with constricted septa and dark smoky brown thin to slightly thickened walls, 15-50 x 4-7 um. Asci arising from simple septa, cylindrical-clavate, 8-spored, 25-30 x 4-5 uum, apical pore deeply blue in MLZ. Spores hyaline, clavate-fusoid, aseptate, without inclusions, 6-8 x 1.5 um. Paraphyses lanceolate, hyaline, without inclusions, 1-3-septate at the base, 40-50 x 3-3.5 um, exceeding the asci by up to 20 um. On dead branches of Myrica gale. 50 Specimens examined: Finland, Waasa, Qvarken (Wallgrund), on dead branches of Myrica gale, 20. VII, 1859, P. Karsten 3891a (syntype in H, annot. S. Huhtinen). Sweden, Dalarna, Sundborn parish, between Lake Langtjarn and the road Karlsbyheden-Korsan, on dead branches of Myrica gale, 18. VII 1978, K. & L. Holm, Fungi Exsiccati Suecici no. 3475 (UPS). This extremely rare species, originally described as a Godronia (Karsten, 1884), is surely not Hyaloscyphaceous and should be placed into the Dermateaceae as it has typical Mollisiaceous hymenium and its marginal hairs are similar to those of some Pirrotaea species. It differs, however, from Pirrotaea in lack of grana and lanceolate paraphyses. The Karsten’s original material in H is mostly juvenile and rather scanty but Fungi Exsiccati Suecici no. 3475 presents rich fully mature material in excellent condition. It seems to be a midsummer species which should be looked for in Myrica gale stands. ACKNOWLEDGEMENTS The author is indebted to the curator of H for the opportunity to visit the herbarium and study Karsten’s type material, and the curators of UPS and GZWU for the loan of valuable specimens. Special thanks are due to Kadri Leenurm (Tartu) for an excellent collection of Jnvolucroscypha involucrata. I thank Dr. E. Kutorga (Vilnius) for valuable critical comments. The study was supported by the Estonian Science Foundation grant 4078 to the author. REFERENCES Eriksson, B. 1970. On Ascomyceetes on Diapensiales and Ericales in Fennoscandia. 1. Discomycetes. Acta Univ. Ups. Symb. Bot. Ups. 19 (4): 1-42. | Karsten, P.A. 1884. Fragmenta mycologica. 7. Hedwigia 23: 5-7. Remler, P. 1979. Ascomyceten auf Ericaceen in den Ostalpen. Bibl. Mycol. 68: 1-321. MYCOTAXON Volume LXXXI, pp. 51-67 January-March 2002 THE PLEUROTUS ERYNGII SPECIES-COMPLEX IN ISRAEL: DISTRIBUTION AND MORPHOLOGICAL DESCRIPTION OF A NEW TAXON LEWINSOHN D.!, WASSER S.P.!, RESHETNIKOV S.V.!. HADAR Y.2 AND NEVO E|! 1. International Center for Cryptogamic Plants and Fungi, Institute of Evolution, University of Haifa, Mount Carmel, Haifa 31905, Israel 2. Department of Plant Pathology and Microbiology, Faculty of Agricultural, Food and Environmental Quality Sciences, The Hebrew University of Jerusalem, Rehovot 76100, Israel Dalia Lewinsohn - E-mail: dlevin01@study.haifa.ac.il ABSTRACT Thirteen wild populations of the Pleurotus eryngii species-complex (Pleurotaceae, higher Basidiomycetes) were collected in various geographical regions during an extensive survey on the occurrence of this complex in Israel. We sampled populations from the Golan Heights in northern Israel extending to the south steppic area of Lahav, in the northern Negev. We defined the exact geographical distribution of this complex in Israel. Previous studies described P. eryngii var. ferulae in close association only with Ferula communis. In our present study we found one population of P. eryngii growing in close association with Ferula tingitana in the Sataf area (Judean Mountains, Israel). The taxonomic study of this peculiar collection belonging to P. eryngii species—complex and the collection from Sataf is described as a new variety P. eryngii var. tingitanus. A comparative morphological analysis and mating experiments were conducted and microscopic features are described. The mycelium growth rate of isolates that were associated with Ferula communis were higher after 7 and 12 days of incubation than of isolates that were associated with Ferula tingitana. The new variety, P. eryngii var. tingitanus is closely related to P. eryngii var. ferulae, but it differs 52 from the latter in spore size, mycelial characteristics, habitus of basidomata, or associated host plant and ecology. KEYWORDS: Pleurotus eryngii var. tingitanus, Ferula communis, F. tingitana, taxonomy, morphology. INTRODUCTION The genus Pleurotus (Jacq.: Fr.) P. Kumm. (Pleurotaceae, higher Basidiomycetes) covers a group of lignotrophic species growing as saprotrophs on deciduous trees and rarely on coniferous trees, or as weak parasites on the lower part of the stems and roots of some Apiaceae species (Hilber 1982, 1997; Singer, 1986; Watling and Gregory, 1989; Zervakis and Balis, 1996). All Pleurotus species are edible or medicinal. Six Pleurotus species: P. ostreatus, P. pulmonarius, P. eryngii, P.cornucopiae, P.cornucopiae var. citrinopileatus, P. dryinus and P. calyptratus, are used in traditional medicine for at least 35 disorders or diseases (Ying et el., 1987; Stamets, 1995; Royse, 1997; Chang, 1999 a,b; Gunde-Cimerman, 1999; Guzman, 2000). Over the last twenty-five years, great efforts have been made to cultivate some taxa of Pleurotus species on a commercial scale. Pleurotus species account for 15% of worldwide mushroom production (Chang, 1999 a,b). Over 1000 species epithets have been proposed in Pleurotus, including names transferred into and out of the genus over the past two centuries (Petersen and Hughes, 1993, 1997; Petersen 1995). Newly proposed names continue to appear. Singer (1986) reported about 40 Pleurotus species for the world mycobiota. Recently, Petersen and Hughes (1997), considered only 15 Pleurotus species, including some North American species. Hilber (1982, 1997), and Moser (1983) reported 11-13 species for Europe, including seven species for the Nordic countries (Kaarik, 1992), five species for Great Britain (Watling and Gregory, 1989), ten species for the former Soviet Union (Michailovsky, 1974). Zervakis et al. (1998) recognized four species in Greece, and eight intersterity groups in Europe (Zervakis and Balis, 1996). Considering that there is no modern complete revision on the genus Pleurotus, there is much taxonomic confusion on the state of several species. It is certainly necessary to revise several described species and also to propose new species by combining classic and modern methodologies. We agree with Bresinsky et al. (1977, 1987) who pointed out that the species concept should be based primarily on macroscopical and 53 microscopical morphological signs, chemical characters, and on ecological observation. Pleurotus eryngii (DC.:Fr.) Quél. species-complex parasitizes the stem and the roots of some Apiaceae and is especially distributed in the subtropical-temperate climate of the Mediterranean region (Italy, Spain, Greece, Cyprus, France, former Yugoslavia, Israel, Lebanon) and central Asia (Uzbekistan, Tadzikistan, Turkmenistan, and Iran). It is also known from central Europe: Hungary (Rimoczi, 1992), Austria (Krisai-Greilhuber and Vogelmayer, 1995), Germany (Hilber, 1982, 1997; Krieglsteiner, 1991), Netherlands (Boekhout, 1990), Czech and Slovak Republics (Dermek, 1973; Klan, 1990), Ukraine (Wasser, 1971), and North Africa (Maléncon and Bertault, 1975). Pleurotus eryngii species-complex are known from the following hosts: Eryngium campestre L., E. maritinum L., Ferula communis L., Cachrys ferulacea (L.) Calestani, Thapsia garganica L., T. villosa var. fuscus, Elaeoselinum asclepium Bert., Diplotaenia spp., Ferulago spp., Laserpitium siler L., L. latifolium L., Pimpinella spp., Opopanax chironium Roch., Prangos ferulacea DC., Peucedanum cervaria (L.) Lapeyr, etc. The P. eryngii species—complex has been described by many authors (Maléngon and Bertault, 1975; Binyamini, 1975; Hilber, 1982, 1997; Moseraybos3 singer yao sou Candusomand sy Basso, 1.9.9,52 Krisai-Greilhuber and Vogelmayer, 1995). The content of the P. eryngii species-complex is interpreted in different ways. This conclusion is confirmed by the observation that the P. eryngii species-complex continues to be designated by one species (P. eryngii) which contains from five varieties to five independent species [(P. eryngii, P. ferulae (Lanzi) Sacc., P. nebrodensis (Inzenga) Sacc., P. fuscus (Battarra) Bres., P. caespitoso-terrester (P. Henn.) Pil.)]. According to Maléncgon and Bertault (1975) there are four different varieties of P.eryngii in Morocco and Algeria (var. eryngii, var. fuscus, var. caespitoso-terrester, and var. ferulae). Hilber (1982, 1997) distinguished in P. eryngii only var. ferulae. Moser (1983) considered all types as one species P. eryngii, while Candusso and Basso (1995), stated that two species are valid, P. eryngii and P. nebrodensis for the mycobiota of Italy. Venturella (1998) illustrated the typification of P. nebrodensis and Venturella et al., (2000) described new var. of P. eryngii, P. eryngii var. e/aeoselini Venturella et al., for the mycobiota of Sicily. Zervakis and Venturella (1998) presented their results of the systematics of the Pleurotus taxa growing on Umbellifers. 54 Prior authors disagreement on the taxonomical status of P. eryngii species-complex has led us to initiate a study of this taxon, which is widely distributed in Israel. Determination of a variety must consider the morphological structures, anatomical features, and other very important aspects such as the ecology and biology of the associated substrates. | In Israel, most of the P. eryngii species—complex grow in close. association with Ferula, where it fruits from November to March. F. communis is the most widespread species of the genus Feru/a in Israel. It is circum-mediterranean, and considered a poisonous plant. The distribution of Ferula communis in Israel has been described by Danin (1982). P. eryngii species-complex in Israel has not hitherto been critically studied. Particular data on P. eryngii appear in some publications devoted to the higher Basidiomycetes of Israel (Binyamini, 1975; Wasser, 1995). In this study we examined the ecogeographical variation and the genetic diversity in populations of the P. eryngii species-complex in Israel (Lewinsohn et al., 2000). Herein, we studied the morphology and defined the geographical distribution of this complex in Israel, and we first reported that in the Judean Mountains P. eryngii species-complex is associated with Ferula tingitana. The taxonomic study and the collection from Sataf is described as a new variety. MATERIALS AND METHODS Biological material During the winters of 1997-99, a total of 226 basidiomata of Pleurotus eryngii species-complex were collected (and 226 dikaryotic strains were isolated) from various geographical regions of Israel (Fig. 1, Table 1). Thirteen populations were delimited among them on the basis of RAPD-PCR analysis of DNA polymorphism (Lewinsohn et al., 2001). These collections have been deposited in the Herbarium of the International Center for Cryptogamic Plants and Fungi, Institute of Evolution, University of Haifa (HAI), Israel, and documented in the “Catalogue of Cultures” (Wasser et al., 2002). 3p) 4 < Aes & Vas = Aa "413 gceer Sheva Negev Fig.1: The distribution of Ferula communis and F. tingitana in Israel. [Marked with small dots on map). Numbers indicate localities where populations of P. eryngii species-complex were sampled (Table 1)]. Note: Population no. 7 grows in close association with Ferula tingitana. All other populations grow in close association with fF. communis. 56 Table 1. Origin of the 13 populations of Pleurotus eryngii species- complex. Population no. Host/Substrate Location Total number on map of isolates | Ferula communis Dishon 6 2 Ferula communis Golan 3 Ferula communis Mt.Tabor 4 Ferula communis | Menahchamyiah 5 Ferula communis Tel-Bet She’an 6 Ferula communis Mt. Gilboa 4 | Ferula tingitana Sataf 8 Ferula communis Har-Hatayasim 9 Ferula communis Zorah 10 Ferula communis Harel Ferula communis Gedera Amazia-Lachish Lahav-Devira Ferula communis Ferula communis Media MEA (Malt extract agar) was prepared according to Gams et al. (1978). The pH of MEA was 6.5+0.4. The medium was autoclaved for 30 min. at 121°C, then cooled and poured into sterile test tubes or Petri dishes. Strain isolation and maintenance of culture stocks Small pieces (about 1 cm in diameter) of the collected basidiomata were removed using aseptic techniques and placed on Petri dishes containing MEA. The cultures were kept at room temperature and transferred several times until pure cultures were obtained. MEA slant tubes were inoculated from Petri dish colonies, incubated at 4° C, and transferred every 6 months for routine maintenance. Actively growing cultures were incubated at 27° C (Gams et al., 1978). A total of 226 cultures of P. eryngii complex were isolated and kept at 4° C for further tests. a Measurements of growth rate of fungal isolates Four dikaryotic isolates obtained from Zorah (associated with F. communis) and four dikaryotic isolates obtained from Sataf (associated with F. tingitana) were inoculated from Petri dishes containing MEA in four replicates (Table 1, Fig.1). The cultures were incubated at 27°C for 10 days. Then, 7 mm disks were cut off with a sterile cork borer from the border of the mycelial colony. Each disk was placed into the center of a sterile Petri dish with Malt extract agar by using a sterile scalpel and incubated at 27°C, until the first strain colonized the entire Petri dish. The growth rate of the cultures were compared after 7 and 12 days of incubation. Mating experiments Freshly collected naturally growing basidiomata of P. eryngii were placed over a Sterile Petri dish in a moist chamber at conditions of slowly decreasing humidity. Basidiospore prints obtained 12-24 hours later were kept in a refrigerator at 4°C. Basidiospore suspensions were prepared in sterile water (1-2 microbiological loops per 5 ml of water), and up to four serial dilutions 1:10 were developed. Basidiospore Suspensions from each dilution of 0.5 ml were spread uniformly on MEA surface on a Petri dish. Inoculated dishes were incubated at 24° C in the dark. One week after incubation, germlings were investigated under the stereomicroscope, and individual microcolonies were transferred by sterile needle into Petri dishes with fresh agar. Colonies obtained were microscopically investigated for clamp connection absence, and 12-14 of them, derived from a single basidomata, were used for determination of mating types in monosporous isolates. Pairings were set up by placing mycelia plugs freshly cut from Petri dish colonies with a cork borer (5 mm diameter) about 15-20 mm apart on MEA Petri plates. After one week when the colonies had grown over the zone of contact, the mycelium was examined for the occurrence of clamp connections. Microscopic features In order to obtain spore-prints, pilei were placed (gills facing down- wards) over white Whatman filter paper #1. The micromorphology, length and width of sixty basidiospores of specimens from each population were measured with the aid of the Olympus Bx100 microscope (magnification 100x). 58 RESULTS AND DISCUSSION The geographical distribution of Pleurotus eryngii species-complex in relation to Ferula spp. and the association between Pleurotus eryngii and Apiaceae Israel is located in an area where the monthly maximum and the total annual values of solar radiation are among the highest in the world, resulting in high annual temperatures and extensive aridity. It consists of three main climatic regions, Mediterranean, steppe, and desert climates (Atlas of Israel, 1970). Our previous study and publications (Lewinsohn et al., 2001) showed a high positive polymorphism correlation with the trend of drought in P. eryngii species-complex. We also found that Israeli populations are well adapted to the hot and dry climate and can tolerate extreme temperature conditions. The geographical distribution and the ecological habitats of Pleurotus eryngii species—complex in Israel parallel the three main climatic regions: Mediterranean, steppe, and desert climates (Lewinsohn et al., 2000) and the distribution of F. communis in Israel (Fig.1). This includes the area of the lower Galilee, the Hula Valley and the slopes of the Golan Heights and Mt. Gilboa, and the southern coastal plain on the sandy stone hills (Gedera area) to the northern Negev (Lahav steppic area). In all these locations, P. eryngii was found in close association with F. communis, in areas heavily grazed by cattle. Only in one place near the water springs of Sataf (located in the Judean Mountains, population 7, Fig. |), we found many individuals of P. eryngii growing in close association with F. tingitana. This is the first report in Israel that describes another Apiaceae host of the P. eryngii. In Israel P. eryngii has not been found in the northern coastal plain, western and upper Galilee, hence its distribution is assumed to be primarily in the Irano-Turanian and marginal Mediterranean phytogeographical regime (Fig.l; Atlas of Israel, 1970). It is interesting how much this complex has a unique association with the higher plants of the Apiaceae species, especially in Israel where many other species in this family grows in the wild and the fungi have selected one species to be associated with, Ferula communis. Such phenomena can raise evolutionary questions, and it could be a result of co-evolution between the host-plant and fungus. The presence of cattle positively selects for this plant, since it is poisonous and normally avoided by cattle. oe) The reason why we found only one place where P. eryngii associates with Ferula tingitana could be, that F. tingitana grows mainly in the Mediterranean climatic regions, on chalk slopes of mountains (Danin, 1982). Possibly the Mediterranean distribution of F. tingitana and the vertical rocky cliffs on which it grows do not provide appropriate region, soil, and habitat, for P. eryngii. Growth rate Comparing the growth rate of cultures obtained from Zorah with those of Sataf shows that: the Zorah isolates grew about 7 mm/day but after 7 days of incubation they reached up to a maximum 50x45 mm in diameter. The Sataf isolates grew about 4.5 mm/day and reached a maximum of 35x32 mm in diameter after 7 days with in the same conditions. After 12 days of incubation the growth rate of the Zorah isolates reached a maximum 80x78 mm in diameter while the Sataf isolates reached only 54x53 mm in diameter. So clearly the Sataf associations grow slower than those of Zorah. Mating experiments Tetrapolar heterothallism as system of sexual reproduction Mating experiments within Zorah and Sataf stocks revealed all 4 mating types of haploid cultures obtained from the basidiospore print of each stock. The mating reactions of Sataf homokaryons can be recognized macroscopically, as positive or negative, while sometimes “barrage” formation was observed. “Barrage” formation in Pleurotus ostreatus species is generally associated with the common presence of B factor (Eger, 1978). We classified mating groups according to this type of reaction. A positive mating was not followed by aerial dikaryotic mycelium formation thus, a careful microscopic examination was needed for clamp detection on hyphae at the contact zone. Monokaryons of Zorah origin developed no “barrage”-type reaction on agar plates. The type of mating reaction can be detected only microscopically, since dikaryotic mycelium was not formed in abundance when compatible monokaryotic hyphae came into contact. In order to investigate the level of biological differentiation of two stocks, four tester monokaryons from each stock were crossed in all possible combinations (Fig. 2). Nevertheless, microscopic examination aided determination of clamp formation in all matings types (Table 2). 60 Fig. 2: Mycelial interaction in agar plates (85 mm diameter) at 27° C on MEA of Zorah (Z on the left side of the plate) and Sataf (S on the right side of the plate) haploid cultures. (Note the slow growing cultures of Sataf compared to those of Zorah). Ar ZyiSs1Z, 1 Ss °ZyaSeh Zi LSB By ZayiSduZ, UeSanZ ates etZaesis 61 Table 2: Results of matings between Sataf (S) and Zorah (Z) stocks tester homokaryons. Mating types and ES vee A»B> strain number Z11 Compatible mating producing clamp connections are represented by a “+”. The results of crossing experiments indicated that no common incompatibility alleles existed between the Sataf and Zorah stocks. This means that at least the two stocks investigated do not belong to a common population, but they are still members of the same biological Species. Basidiospores measurements Tukey-Kramer statistics test indicated that the Sataf basidiospores are significantly (p<0.05) larger [(10) 12-15(15.8) x (3.8) 5-7.2 (7.9) um] (Fig. 3), than the rest of the 13 populations that were examined. Additionally, basidiospores from the Golan were smaller [(7.6) 8.8-10.6(11) x (4.7) 5-6 (6.9) uum] and from Har Hataysim (located only about 10 km apart from Sataf) they were significantly smaller [(6.3) 7.5-9.6(10.6) x (3.9) 4.5-5.8 (6.7) um] than the other basidiospores from all other populations. Morphological description A study of morphological characteristics (basidiomata, microstructures, mycelial cultures) and ecology showed that the P. eryngii species-complex is represented in Israel by two well-distinguished taxa: P. eryngii var. ferulae and P. eryngii var. tingitanus var. nov. Below, the descriptions of a new variety for science is presented. Pleurotus eryngii (DC.:Fr.) Quél. var. tingitanus Lewinsohn var. nov. (Figs. 3, 4). Fig 3: Pleurotus eryngii var. tingitanus (HAI, Typus). a. elements of pileal cuticle c. basidia b. spores d. cheilocystidia Fig. 4: Basidiomata of P. eryngii var. tingitanus (growing in close association with Ferula tingitana). A tipo (P. eryngii var. eryngii) differ: Pileus 6.5-26 cm latus. Basidia tetrasterigmata 23-33x8-11 um, clavate. Cheilocystidia 24-43x9-16 um. Sporae 10-15.8x5.0-7.9 um, cylidraceae, ellipsoideae, hyalinae. Typus: Israel, Sataf, ad radices Ferula tingitana L., 02.02. 1999, leg. et det. D. Lewinsohn, in herbario INST. Evolution (HAI) conservatur, N-3399. Etymology: P. eryngii var. tingitanus, named after the host that it was found to be associated with, Ferula tingitana. Pileus: 6.5-26 cm in diameter, thick-fleshy, first hemispherical, later flat and fully expanded to concave, slightly centrally depressed, with inrolled and sometimes remaining down-turned margin, or lobed at the margin or splitting; surface in young specimens smooth, radially fibrillose, or with appressed squammules, especially to the margin, sometimes at the margin becoming areolate-cracked, uniformly colored, snuff brown buff, ochraceous brown, cream. Pileal cuticle consisting of ochraceous brown, cream, thin-walled, cylindrical, septate hyphae 3-16 um in diameter with clamp connections. Lamellae: decurrent, often slightly anastomosing near stipe, rather thin, up to 10mm wide, cream when young, cream-clay, later pale pinkish 64 cream with somewhat brownish edge. Lamellar trama: monomitic, generative hyphae densely interwoven, 3-13 um in diameter, thin-walled, with clamp connections. Basidia: 4-sterigmate, 23-33x8-11 wm, clavate. Cheilocystidia: 24-43x9-16 bm, irregularly clavate, irregularly distributed (some parts of the gill edges even without cheilocystidia). Spore print white (fresh), later slightly ochraceous. Basidiospores: (10) 12-15 (15.8)x(3.8) 5-7.2 (7.9) um, hyaline, oblong to cylindrical. Stipe: 1.3-12x1.5-3.8 cm subcentral to eccentric, rarely central, cylindrical, tapered from apex to base, whitish-cream, ochraceous, smooth or subpubescent with squamules. Flesh solid, elastic, whitish, becoming yellow; mild taste: anise-like odor. Mycelial cultures: pink to yellow-orange pigments especially on monokaryotic mycelial cultures. Mycelial colonies mostly presenting a flat and loose aerial mycelium. Growth rate of mycelial cultures about 4.5 mmi/day. Type: Israel, Sataf springs, in the Judean Mountains, associated with Ferula tingitana 02.02.1999, coll. D. Lewinsohn. Note: This new variety is morphologically related to P. eryngii var. ferulae, but differs in spore size, mycelial characteristics, and growth rate of the mycelia, habitus of basidiomata, associated host plant, and ecology. The data about P. eryngii var. ferulae elucidated on the basis of previously reported literature (Hilber, 1982, 1997; Boekhout, 1990; Krisai-Greilhuber and Vogelmayer, 1995). CONCLUSION In this study we define the geographical distribution of the P. eryngii Species-complex in Israel. We examined the morphological and cultural characteristics and mating system of a P. eryngii population, which enabled us to describe a new variety of P. eryngii complex in Israel. This is the first report on an association between P. eryngii and Ferula tingitana. The morphological differences enabled us to distinguish between the two varieties of P. eryngii for the mycobiota of Israel; P. eryngii var. ferulae and P. eryngii var. tingitanus. ACKNOWLEDGEMENTS The authors thank Prof. G. Zervakis for his useful comments and review of the manuscript; Prof. A. Danin for his aid in the identification of Ferula tingitana and for the information on the distribution of F. communis in Israel; Prof. A. Buchalo for her aid in morphological studies; Dr. E. Lewinsohn for editorial comments; Dr. Y. Tadmor, for his invaluable assistance with the statistical analysis; and Mr. M. Margulis for his assistance with the photographs. We also thank Mr. David Shahak of Kibbutz Tirat-Zvi for his help during sampling. This study was also 65 supported by the Israel Discount Bank chair of Evolutionary Biology and the Ancell-Teicher Research Foundation for Genetics and Molecular Evolution. BIBLIOGRAPHY Atlas of Israel (1970). Surveys of Israel Ministry of Labor. Ed. Amiran D.H.K., J. Elster, M. Gilead, N. Rosenan, N. Kadmon and U. Paran, Jerusalem and Elsevier, Amsterdam. Bresinsky, A., O. Hilber and H.P. Molitoris (1977). 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Zervakis and S. La Rocca (2000). Pleurotus eryngii var. elaeoselini var. nov. from Sicily. Mycotaxon 76: 419-427. Watling, R. and N.M. Gregory (1989). 6/Crepidotaceae, Pleurotaceae and other pleurotoid agarics. In: British Fungus Flora, Agarics and Boleti. Royal Botanic Gardens. Edinburgh, 157 pp. Wasser, S.P. (1971). New and little known species of the Agaric fungi for the USSR from the steppe zone of the Ukraine. Ukrainian Journal of Botany, 28: 299-304. Wasser, S.P. (1995). Edible and Poisonous Mushrooms of Israel. Modan Publ. House, Tel Aviv, 185 pp. (In Russian). Wasser, S.P., I. Dukman and D. Lewinsohn (2002). Catalogue of Cultures (Higher Basidiomycota). Ed. E. Nevo., Second edition (in press). Ying, J., X. Mao, Q. Ma, Y. Zong and H. Wen (1987). Icones of Medicinal Fungi from China. Science Press, Beijing, 575 pp. Zervakis, G. and C. Balis (1996). A pluralistic approach in the study of Pleurotus species with emphasis on compatibility and physiology of the European morphotaxa. Mycological Research, 100: 717-731. Zervakis, G. and G. Venturella (1998). Towards the elucidation of systematics of the Pleurotus taxa growing in Umbellifers. In Abstracts of the Sixth International Mycological Congress. Jerusalem, Israel, p. 7. Zervakis, G., Dimou D. and Balis C. (1998). A check-list of the Greek macrofungi, including hosts and biogeographic distribution: Basidiomycotina. Mycotaxon, 66: 273-336. ry 4) ; hg : he ¥ : ae aes i val vey ih 7 wt Mt , ie Paina Beatin -d or i te ty hidy ake eo ote yin kines " Wis Naina i dC al | re a Say vi ns ent Li Lewy ba sith! V 4s eee ir ian ~, i a os a bk FY. i | enti Menthyrhe sh 5, saa Dv deh he bine ‘Pim | te Se a aaa ab Weondatinn vihats er ARETE a ea . Sidkieis'c oh iy We ee cupid ihe Bene rt ae ins ht i 1am vata yyt teh: a t ati Way yp cho aa i'd : ~ te acs: ae th hee Prete sah ‘ jars: {Stns ateRRATA ‘* ey ee Mar: hile ick ceeee Pana agit? gr the Sobrnyt %. Py Ae! Pi Was ik af cand ae a : ; j rh ae dee, WR AYA sialic: SORE Ae ae iifes iy, ein toy? ae ; at (evnn weet Tt vy wits rears hp eared CGY Piha yh, ae i SRR AON a ee Pe SE tii, e Lee ae a \eNesdie Ahab ais 4s, aA sO simcnih-a iat 4A " wot hing sw sresaienbd heute . te) ee tet te BS” Manly a ieny ree perl a aie mM incerdaayn te. wing, ek ey acre mer ee | eeary ee RN ie Dia 4 iMiartees ph ee | ot mas (A < ol rie Mim 1" qa et ccc Wie spi ue inky vain’. Varia |: te Cae O ERM ear i iat aie Bao EL eyed, ri Hy! pres ‘ & | ys Shy) RPM aa ash hid oti + LR Sal a pint 1 ee ot ar ae eae OWI eet, My obn eee ae atl beeen | ini ie] ieerirader,” ha: ty gS rate im byt tae 4 eee’), ba tonsa id z wal de B oa vay, actin ou ear pe ( ' pie a iia Ancarwestatll at ert & E i ae d .* 5 Wiahe sage rw ten oft i oe tyes te es. ws hy Sih aa ane pride Abb caddis, Ei a abcagl HE Owe Ia Al. ty ath az Pet ae | +h ty Tie iis Be aan ae Fey aids ll ay enero pr vA tani res aed pe iat aR al Alek gate cine dee I a ™ a fare arini wan set im . Wit LE Legh Boling Be fe eo sky i pean : ‘eeebanrs ay igh i blue ‘a divest $ wt ipa Pai . Ci ha: a re agp en a f nl rane iti Yi ens ie x he) eee aerate Peers # i than i bis at }e TR Peg Hh re IN Saline ena: tt ee a ‘ebibast ia hea vehi moneys Rest 2) fiche: i Sesto unaia Seey } m yt Nate ciple skit is Nile SA Aer eee ea ‘gts me ftil: Lowney aN Bhp cos sate lod ay. iit + PNM ate Aerts: hen Hy Ry ae bes Hest, Th et sey hsp hC LAS ORE NN ET MRT, wis Tpit yf ibis kant His sil Gane is: me ia jak Wi i hte te “2 ara el Ri gintrp wn aya iy eS Sparen. Sygate Bata Me SN Bega o y b a ian wi Moa pv adrian Bat IS | Hijet ah axe 1 Ne ‘ caret ru am ; vie coe — ee ; cia eh ms} Al «a ths A > ath 7 y a acne f is MYCOTAXON Volume LXXXI, pp. 69-74 January-March 2002 PLEUROTUS ERYNGII VAR. THAPSIAE VAR. NOV. FROM SICILY G. VENTURELLA!, G. ZERVAKIS?2 & A. SAITTA! 1Dipartimento di Scienze Botaniche, University of Palermo, Via Archirafi 38, I-90123 Palermo, Italy. gvent@unipa.it 2National Agricultural Research Foundation, Institute of Kalamata, Lakonikis 85, 24100 Kalamata, Greece. zervakis@kal.forthnet.gr ABSTRACT Field investigations in the Sicilian territory allowed the study of Pleurotus eryngiu basidiomata growing in association with Thapsia garganica. The evaluation of morphological, ecological, distribution, and isozyme and PCR- RAPD data together with a revision of Bresadola’s herbarium specimens kept in the Herbarium Botanische Staatssammlung (M) led to the description of a new variety, Pleurotus eryngt var. thapsiaevar. nov. KEY WORDS: fungal taxonomy, ecology, Mediterranean area, Pleurotus eryngit, Umbellifers, edible mushrooms, Apiaceae. INTRODUCTION Recent mycofloristical, geographical and ecological studies together with RAPD-PCR and isozyme analysis were carried out to elucidate the perplexed relationships among the different taxa growing in association with Umbellifers and forming part of the so-called “Pleurotus eryngii species-complex” (Zervakis & Balis, 1996; Zervakis & Venturella, 1998; Venturella, 2000; Venturella & al., 2000; Zervakis et al., in press). According to these investigations the infrastructure of the P. eryngii species-complex in the Mediterranean area is composed of the following host- specific taxa: Pleurotus eryngt (DC. : Fr.) Quél. var. eryngiu growing on root residues of Eryngium campestre L., P. eryngii var. ferulae Lanzi on Ferula communis L. subsp. communis, P. nebrodensis (Inzenga) Quél. on Cachrys ferulacea (L.) Calestani, and P. eryngti var. elaeoselini Venturella et al. on Elaeoselinum asclepium(L.) Bertol. subsp. asclepium. In addition, an interesting and misappreciated P. eryngii population growing on Thapsia garganicaL. in Sicily, which was considered to belong to P. eryngl var. eryngit was studied in detail. A re-evaluation of morphological, ecological, distribution and other taxonomic data permitted its proposal as a new variety, Pleurotus eryngti var. thapsiae 70 MATERIALS AND METHODS Periodical field observations and collection of Pleurotus basidiomata growing on Thapsia garganica L. have been carried out in Sicily from 1995 until the present day. Morphological, distributional and ecological characters were also evaluated in conjunction with the examination of Pleurotus specimens kept in the Herbarium Botanische Staatsammlung (M). Microscopical observations were made in an aqueous solution with the aid of a micrometer (x1000). Spores were taken directly from the spore prints which were obtained on glass slides. For examination of herbarium material a piece of lamellae was removed from each sample and rehydrated in L4 or KOH 5%. Mycelium cultures were maintained on PDA (Bacto potato dextrose agar, Difco), and are stored in the Culture Collections at the authors’ Institutions. The type specimen is kept in the Herbarium Mediterraneum (PAL). DESCRIPTION Pleurotus eryngiivar. thapsiae Venturella, Zervakis et Saitta, var. nov. (Fig. 1). Etym.: from the name of the genus of the associated plant (i. e. Thapsia). SyNn.: Pleurotus fuscus (Batt.) Bres. var. ferulae Lanzi s.1. Bres.; Pleurotus fuscus (Batt.) Bres. var. nebrodensis Bres. s.1. Bres. Basidiomata valida atque robusta, una, haud saepe luxuriosa. Pileus 2-10 cm, convexus dein explanatus, postremo depressus, perraro_ reflexo- infundibuliformis. Pilei pellis haud facile separabilis, velutina atque opaca, cerea cum pigmentis, et laxatis squamis, fusca, margine tenui acutaque, maxime involuta, subtilibus venis ornata, in senectute irregulariter plana et interdum lobata. Lamellae, cineraceo colore, spissae, arcuatae, cum lamellulis, decurrentes, anastomosantes in supera parte stipitis, plus minusve evidentes. Stipes 4-7,5 x 1,2-2,8 (3,6) cm, solidus, plenus, irregulariter cylindricus, ad basim attenuatus vel subradicatus, medianus vel excentricus, albus paene pullus colore, inde levis et glaber. Caro compacta, fibrosa, elastica, alba; odor saporque debiles, vulgares. Sporae 10-14 x 5-7 um (Q = 1,9-2,1), hyalinae (in KOH etiamque in Melzer), leves, cylindraceae vel irregulariter ellipsoideae, pluriguttulatae, apiculo manifesto praeditae. Basidia 32-54 x 6-9 um, tetraspora, clavata, cum sterigmatibus 4-6 ym longis. Cheilocystidia 8-12 x 40- 65 um, basidiis similia, superne attenuata, interdum acute mucronata. Pileicutis ex hyphis cylindricis radialibus, 5-12 pm latis, constituta. Fibulae praesentes. Locus: graegaria in herbidis locis, cum Thapsiae garganicae consociata. Typus — Madonie Mts, locality Piano Zucchi (Palermo, Sicily), 37° 54’ 01” N 13° 59’ 40” E, arid pastures with Thapsia garganica on calcareous soil, 1000 m, 11 May 1995, leg. G. Venturella (holotypus in PAL). Basidiomata sturdy and fleshy, single, seldom caespitose. Pileus 2-10 cm, convex then flat, finally depressed, seldom reflexed or funnel-shaped. Cuticle not easily detachable, velvety, pruinose with pigments, opaque, with scattered squammules, dark brown, then smooth, bright and greasy-like appearance in 71 ripe basidiomata. Margin thin, acute, deeply involute and adorned by thin ribs, then flat, irregular, and sometimes also lobate. Lamellae greyish white, thick, arcuate, mixed by lamellulae, deeply decurrent with anastomoses, more or less evident, at the point of attachment with stipe. Lamellae margin entire, subconcolorous to the pileus. Stipe 3-5 x 1-3 cm, sturdy, filled and firm, irregularly cylindrical, attenuate at the base, sometimes radicating, central and sometimes eccentric, white-brownish, then smooth and glabrous. Flesh firm and compact, fibrous, elastic, white, with a fungus-like smell and taste. Spores 10-14 x 5-7 pm (Q = 1,9-2,1), cylindrical to irregularly ellipsoid, smooth, hyaline, guttulate with pronounced apiculus. Basidia, club-shaped, 4- sterigmata, 32-54 x 6-9 um, sterigmata 4-6 um. Cheilocystidia, 8-12 x 40-65 um, rare, club-shaped, with irregular apex, acute to subglobose, sometimes showing a thin acute beak. Epicutis with cylindrical hyphae, 5-12 um wide, closely appressed. Texture cutis-like. Clamp connections abundant in all parts of the basidioma. Basidiospores, basidia and trama (with occasional presence of oleiferous hyphae) hyaline in aqueous KOH and Melzer's reagent. Culture characteristics on PDA: pigments absent, reverse colour unchanged, dikaryotic colonies mostly presenting a loose submerged and suppressed aerial mycelium, more or less zonate and radial, growth margin even and regular, colour white to cream; thin-walled hyphae, hyaline in aqueous KOH and Melzer's reagent, with abundant clamp-connections; occasional production of microdroplets, singly on short secretory sterigmata on aerial hyphae (nematode trapping devices); optimal fungal growth 25°C (2.2 mm day). Fig. 1. Basidioma of P. eryngii var. thapsiae. <P Specimens examined MUNCHEN: Fiemme alla Bellamonte, ad radice, Laserpitii latifoliae in pratis, IX. 1925, Pleurotus fuscus (Batt.) Bres. var. ferulae Lanzi (M). PALERMO (SICILY): Piano Zucchi, on root residues of Thapsia garganica, calcareous soil, 1100 m, 27 Apr 1995, 11 May 1995, 06 May 1996, 29 Apr 1997, 13 May 1997, 24 May 1998, 16 May 1999, 25 Apr 2000, 08 May 2000, leg. G. Venturella (PAL); Piano della Madonna, on root residues of Thapsia garganica, calcareous soil, 1500 m, 12 May 2001, leg. A. Saitta (PAL). DISCUSSION In the Mediterranean area the Pleurotus eryngii species-complex includes many host-associated taxa. Particularly in Sicily, the Pleurotus populations associated with Umbellifers show a high degree of diversity (Zervakis & Venturella, 2002). Bresadola (1932) considered Pleurotus nebrodensis (sub Agaricus eryngt var. nebrodensis) as a synonym of P. fuscus (Batt.) Bres. var. ferulae Lanzi. Within P. fuscus var. ferulae s. 1., two ecotypes were distinguished by Bresadola: P. fuscus var. ferulae growing on Laserpitium latifolium (the so- named “fungo d’Ampezzo” or “fungo della Bellamonte”) in central Europe (including northern Italy) and P. fuscus var. nebrodensis on Ferula communis ssp. communis, Elaeoselinum asclepium ssp. asclepium, Opopanax chironium, etc., in Sicily on the Madonie Mts. Mattirolo (1938) realized that “probably the P. fuscus varieties reported by Bresadola could be considered as local varieties of P. eryngit which are associated with different Umbellifers and habitats’. Mattirolo’s interpretation is indeed in accordance with the individual population structure within the Pleurotus eryngii species-complex in Sicily, where basidiomata growing on various host-plants (other than Eryngium and Ferula spp.) possess intermediate characters between P. eryngii var. eryngii and P. eryngu var. ferulae (Zervakis & Venturella, 1998). Besides, recent investigations demonstrated that Pleurotus nebrodensis is strictly associated with Cachrys ferulacea, whereas Elaeoselinum ascelpium ssp. asclepium is the host-plant of P. eryngi var. elaeoselini (Venturella, 2000; Venturella et al., 2000). The same studies could not reconfirm the occurrence of P. nebrodensis on Opopanax chironium, while they provided the evidence that Ferula communis is the only plant species associated with P. eryngti var. ferulae. In addition, RAPD-PCR and isozyme analysis (Zervakis & Venturella, 1998; Zervakis et al., in press) confirmed a clear separation of P. nebrodensis from the “P. eryngii species-complex”. Furthermore, several morphological characteristics and particularly the dark colour of the pileus of P. fuscus var. ferulae (Bresadola, 1932) excluded any resemblance to the current notion of P. nebrodensis and P. eryngii var. elaeoselint which are both characterized by white basidiomata (Venturella, 2000; Venturella et al., 2000). Hence, the description of P. juscus var. ferulae should correspond to roe taxon. Indeed, the macroscopic and microscopic features of P. fuscus var. ferulae (sensu Bresadola) are very similar to those of Pleurotus basidiomata growing on Thapsia garganica Moreover, the observations on the Pleurotus fuscus var. ferulae herbarium specimens, collected in 1925 by Bresadola on Laserpitium 73 latifolium, and kept in the Herbarium Botanische Staatsammlung (M), sub P. eryngu var. nebrodensis (a Hilber’s revised attached label), show that the spore size is smaller than in P. nebrodensis and corresponds to the spore range of taxa included in the P. eryngii species-complex. It also evidences that Pleurotus strains growing on Laserpitium spp. in central and northern Italy present identical morphological characteristics to those of Pleurotus populations associated with T. garganica in Sicily. Hence, these two allopatric populations growing on different host-plants belong to the same taxon, P. eryngii var. thapsiae. Pleurotus eryngtu var. thapsiae is a spring mushroom, locally named “firrazzolu” with reference to the vernacular name used for Thapsia garganica in the Madonie Mts. (north Sicily). P. eryngii var. thapsiae was collected on calcareous soils, in arid pastures with Thapsia garganica and Eryngium campestre, and at elevations ranging from 1000 to 1500 m. The appearance of the basidiomata is restricted, from the end of March until May. The basidiospores in P. eryngi var. thapsiae are irregularly ellipsoid to cylindrical (10-14 x 5-7 pm, Q = 1.9-2.1) (Fig. 2). This variety possesses also basidiole-like cheilocystidia with an attenuate apex at the top, and sometimes with an acute beak, showing a close resemblance to P. eryngii var. elaeoselini microscopic features. The macroscopic and microscopic features of this new taxon correspond to those described for P. fuscus var. ferulae s. Bresadola (1932), as confirmed by the observations made on the specimens kept in M. Fig. 2. Pleurotus eryngii var. thapsiae- a. spores, b. basidia, c. cheilocystidia, d. epicutis. 74 P. eryngii var. thapsiae differs from P. eryngii var. eryngi and from P. eryngu var. ferulae in producing significantly smaller and darker basidiomata. In addition, the P. eryngii var. eryngii basidiomata are produced singly or in clusters from early autumn to winter, while basidiomata of P. eryngu var. thapsiae appear mostly singly and in spring. Moreover, P. eryngii var. eryngiiis quite common and occurs from sea level to 1500 m while P. eryngii var. thapsiae is infrequent and restricted to the mountainous zone. P. eryngiti var. thapsiae is also ecologically separated from P. eryngi var. ferulae; the latter grows on Ferula communis ssp. communis root residues in garrigues, waste lands and arid pastures, and it is widespread throughout Sicily, mainly on calcareous soils from sea level to 1000 m. Analysis based on isozyme and PCR- RAPD data (Zervakis & Venturella 1998; Zervakis et al., in press) further confirmed the distinct taxonomic position at the varietal level of Pleurotus populations growing in association with Thapsia garganica Therefore, the interpretation of all available data support the hereby proposed description of a new Pleurotus taxon from Sicily, P. eryngiivar. thapsiae ACKNOWLEDGEMENTS Financial support by Assessorato Agricoltura e Foreste della Regione siciiana, P.O.P. 1994/99 -— Misura 10.4 - Ricerca applicata, indagini e sperimentazione di interesse regionale. “Progetto di sviluppo per la fungicoltura in Sicilia (Progetto FUNGIS)” is gratefully acknowledged. LITERATURE CITED Bresadola, G. 1932. Funghi mangerecci e velenosi. 1. Societa Botanica Italiana, Sezione Lombarda — Museo di Storia Naturale della Venezia Tridentina, pp. 231-232, Tav. 75. Mattirolo, O. 1938. I] Pleurotus fuscus (Batt.) Bresadola var. ferulae Lanzi nelle Alpi occidentali. Nuovo Giornale Botanico Italiano65(3): 357-363. Venturella, G. 2000. Typification of Pleurotus nebrodensis. Mycotaxon 75, 229- PAS by Venturella, G., Zervakis, G. and La Rocca, S. 2000. Pleurotus eryngiti var. elaeoselinivar. nov. from Sicily. Mycotaxon 76: 419-427. Zervakis, G. and Balis, C., 1996. A pluralistic approach on the study of Pleurotus species, with emphasis on compatibility and physiology of the European morphotaxa. Mycological Research 100: 717-731. Zervakis G. and Venturella G. 1998. Towards the elucidation of the systematics of the Pleurotus taxa growing on Umbellifers. In Proceedings of the Sixth International Mycological Congress(abstract). Jerusalem, 23- 28 August 1998. Zervakis G. and Venturella G. 2002. Mushroom breeding and cultivation favors ex situ conservation of Mediterranean Pleurotus taxa. In Managing Plant Genetic Resources pp. 351-358. Eds. J.M.M. Engels, A.H.D. Brown and M.T. Jackson. CAB International. Zervakis, G., Venturella, G. and Papadopoulou, K. in press. Genetic polymorphism and taxonomic relationships of the Pleurotus eryngii species-complex as resolved through the analysis of random amplified DNA patterns, isozyme profiles and ecomorphological characters. Microbtology-UK. MY COTAXON “Volume LXXXI, pp. 75-82 January-March 2002 MATERIALS FOR A HYPOGEOUS MYCOFLORA OF THE GREAT BASIN AND ADJACENT CORDILLERAS OF THE WESTERN UNITED STATES. VII: A NEW TRUFFLE-LIKE PEZIZA (ASCOMYCOTA, PEZIZALES) Robert Fogel University of Michigan Herbarium, North University Building, Ann Arbor, Michigan 48109-1057 Jack States Department of Botany, University of Wyoming, Laramie, Wyoming 82071! ABSTRACT A new truffle-like species of Peziza, P. quercicola, associated with Quercus gambelii and Q. turbinella from southwestern Utah, United States, is described and illustrated. Key Words: hypogeous fungi, sequestrate, biogeography. This is the seventh in a series containing materials for a hypogeous mycoflora of the Great Basin and adjacent Cordilleras (Fogel 1994; Fogel and Pacioni, 1989). The region has received little attention from mycologists despite its potential for studies of fungal biogeography, evolution, and coevolution. The Great Basin, as defined by Cronquist et al. (1972), occupies an area of 714,854 square km. The vegetation in the region contains many suitable ectomycorrhizal hosts for hypogeous fungi patchily distributed. The species composition of a specific stand reflects the interaction of environmental optima for species, soil factors, pre-glacial flora, extent of glacial advance, pluvial lake levels, and subsequent re-invasion from glacial refugia (Wells, 1983). Another significant factor affecting the presence of hypogeous fungi is the efficiency of mycophagous rodents, and perhaps insects, in spore dispersal. ‘Address for correspondence: 2 Canyon Shadows Rd., Lander, WY 82520. 76 Our survey of the Great Basin and adjacent Cordillera has involved collecting in a large number of habitats. One of the most interesting and difficult habitats to- collect is the Interior Chaparral (133.3) vegetation-type (Fig. 1) (Brown, 1994). This vegetation-type is widely scattered along the forest islands comprising the axial route for conifer re-invasion of the Great Basin in southwestern Utah and extends south into Arizona (Wells, 1983). The stands are usually open becoming quite dense along permanent streams. The larger shrubs reach 2 to 2.5 m in height. We were interested in these stands because they contain Shrub Live Oak (Quercus turbinella Greene) a putative host for ectomycorrhizal truffles and false-truffles. The windows for sporocarp production are very limited because the monthly precipitation during the growing season averages between 5 and 16 mm (http://www. utah.com/weather/stgeorge.htm). Another interesting habitat in the same area is the Great Basin Montane Scrubland (132.1) vegetation-type (Fig. 2) (Brown, 1994). These stands contain the deciduous Gambel’s oak (Q. gambelii Nutt.) and evergreen mountain mahogany (Cercocarpus ledifolius Nutt. in T. & G.) a member of the Rosaceae. Both of these shrubs are putative hosts for ectomycorrhizal hypogeous fungi. Often the stands consist of thickets 1 to 6 m in height. The best collecting is around springs and seeps where the Gambel’s oaks are larger, up to 15 cm in diameter, and the stands are more open. The windows for collecting are longer, but sufficient precipitation is often limiting. We visited these stands on numerous occasions but rarely collected any hypogeous fungi. The truffle-like Peziza described below is one of the hypogeous fungi that we did collect. MATERIALS AND METHODS Methods of collection and study of the morphological and anatomical features are essentially those of Smith (Smith and Zeller, 1966). Spores were measured in freehand sections mounted in Melzer’s reagent. Lengths and widths of 20 ascospores randomly selected from a mature ascoma in each collection were measured for statistical comparisons. Color names are given as the ISCC-NBS names with the numerical equivalent in parentheses (Kelly and Judd, 1965). Herbarium names are abbreviated according to Holmgren, Holmgren, and Barnett (1990). Fig. 1. Interior Chaparral (133.3) vegetation-type, Tobin Wash, Utah. Fig. 2. Great Basin Scrubland (132.1) vegetation-type, Oak Grove Campground, Utah. Fig. 3. Sectioned ascoma, note large locules formed by infolding, Fogel F4971 (MICH). 78 The distribution map was created by using ArcView 3.1 Geographic Information System personal computer software (ESRI Inc., Redlands, California) to overlay coordinates of localities on a political boundary base map. The shaded contour map theme for Utah is an uncompressed TIFF image in 24-bit color from Chalk Butte Inc., Boulder, Wyoming. Specific locality data available from MICH. TAXONOMIC TREATMENT PEZIZA QUERCICOLA Fogel & States, sp. nov. Figs. 1-6. Ascocarpia usque ad 30 x 13 x 15 mm, profunde sulcata. Peridium glabrum, album, lutescens. Loculi magnum, cavus. Asci cylindrici, octospori, .amyloidei (fortissime ad apices), ad apices indentati. Sporae ellipsoideae, 13-17(-19) x 10-13 yum, ornamentis verrucarum <0.2 um, hyalinae, uniguttulatum. Paraphyses 6-8 um in diam., apicibus clavatis. Holotypus: USA, Utah: Kane Co., 22 May 1995, Fogel F4968 (MICH). Etymology: Dweller in Quecus (oak) forests. Ptychothecia of irregular symmetry, oblong to turbinate, up to 30 x 13 x 15 mm; multichambered, chambers empty, formed from infolding of ascoma wall, opening to chambers not apparent, lined by a farinose, white to translucent hymenium. Ascoma wall 1 mm thick, fleshy, brittle; abhymenium white with brilliant yellow patches (83) or bruising brilliant yellow, glabrous, farinaceous at X10. Medullary excipulum white, turning brilliant yellow when sectioned. Hymenium white. Attached by a prominent basal tuft of hyphae holding a small “button” of soil. Odor fungoid to strong, sweet, phenolic or dimethyl! sulfide-like. Ascospores ellipsoid, verruculose, 13-17(-19) x 10-13 um, mean length 15.5 + 0.8 wm; width 11.2 + 0.7 um, length to width ratio 1.39 + 0.1 um, n=140 spores from 7 collections, wall 0.5 um thick including ornamentation; ornamentation of warts < 0.2 um tall, weakly cyanophilic; containing a single large guttule. Asci arranged in palisade, cylindrical, 156-198 x 13-15 um, operculate with 8 uniseriate spores, opercula 7-8 um broad, indented tip and apical pore blue in Melzer’s reagent, immature asci dextrinoid (dark orange), 17-25 wm broad; walls thin; croziers lacking. Paraphyses filamentous, capitate, tip 9-10 um broad, stalk 6-8 um broad, extending slightly past asci (10 um), not abundant. Ascomata walls 550-650 um thick, no clear demarcation between ectal excipulum and medullary excipulum; surface sparsely covered with short, blunt, hyaline hairs, terminal cell 15-30 x 8 wm; context pseudo- parenchymous, cells of hyaline, thin-walled, 5-6 um broad hyphae swelling to 20-30 x 21-23 um. 79 BRR Ri a a ce i i Hy _ ; : i Hi = i / 7 i i / i He oS Co _ Hh | a | _ | | He Hi i HH 7 i 7 | i bid i i i He Hi | _ / . ee . Ht i | _ _ _ na ~ / fod bale kde PMNL AIP onl ON eye RN ok Na ec ae ERY ate a Fig. 4. Spores mounted in Melzer’s reagent, 1600X, Fogel F4968. Fig. 5. Ascus tip bluing in Melzer’s reagent, 1600X, Fogel F5294. Fig. 6. Distribution of Peziza quercicola in southwestern Utah, USA. Outline (white) of Great Basin in upper left superimposed on political boundaries (black) and shaded topography. Large low relief areas inside the white outline are dry, pluvial lake beds. Mapped localities are in Washington (W) and Kane (K) Counties. 80 Material examined: USA, UTAH, Kane Co., Fogel F4968 (Holotype). Washington Co., Fogel F4851, F4852, F4881, F4907, F4908, F4971, F5225, F5294 (all MICH). Putative mycorrhizal hosts and phenology: Hypogeous most commonly under Shrub Live Oak (Q. turbinella) in the Interior Chaparral (133.3) vegetation-type, found less frequently under Gambel’s oak (Quercus gambelii) in the Great Basin Montane Scrubland (132.1) vegetation-type (Brown, 1994). Fruiting from early May to late June at 1300-1800 m elevation in southwestern Utah (Fig. 6). Discussion: Two of the collections examined, F4852 and F5225, were immature but matched the description of Peziza quercicola in every other detail. F4968 was the most mature collection and had larger spores on average than the other collections The ellipsoid, uniguttulate spores suggest Peziza quercicola is more closely related to P. shearii (Gilkey) Korf, P. whitei (Gilkey) Trappe, P. stuntzii Trappe and P. ellipsospora (Gilkey) Trappe than either P. sparassiformis (Henn. in Warb.) Sacc. & Syd. in Sacc. or P. proteana (Boud.) Seaver. P. shearii from Virginia differs from P. quercicola in having subglobose spores (I/w = 1.1-1.2 vs. 1.39) with large tubercules. P. whitei has shorter, narrower spores (11.3-14.0 x 8.1-11.3 um) and is only known from Australia and New Zealand. P. stuntzii from central Washington State differs macroscopically in having a solid gleba and pink flesh. The spores of P. stuntzii are ornamented with cyanophilic warts and short ridges. P. ellipsospora from California and Oregon has “purplish brown’ ascoma, smaller spores (“14 x 10 wm” vs. 13-17(-19) x 10-13 pum), taller ornamentation (< 1.0 tm vs. < 0.2 um) and narrower paraphyses (2-5 um vs. 6-8 pm, swelling to 9-10 um broad at tip). Arora (1986) describes a “Peziza sp. (unidentified)” from under oaks in coastal California that might be our P. quercicola. His photograph on p. 822 very Closely resembles P. quercicola. Analysis of fossil packrat middens provides evidence for a much wider distribution of P. quercicola than our collecting indicates (Fig. 6). P. quercicola may have accompanied the northward spread of oaks during the late Wisconsin and early Holocene (Spaulding 1990) into the Great Basin. Contact between oaks in the Great Basin and California could have occurred at this time. Pifion pine (Pinus monophylla Torr. & Frém.) another member of the Mojavean Woodland grew on the west side of the Sierra Nevada in the late Wisconsin (Spaulding, 1990). | The addition of Peziza quercicola to a growing list of sparrassoid and ptychothecial taxa in Peziza raises questions about relationships within the Pezizaceae. Peziza is a large, common genus producing mostly epigeous 81 apothecia that forcibly release spores to the atmosphere. There are taxa however that produce ptychothecia (sparassoid ascomata) characterized by infolding of the hymenium to produce a urnulate to turbinate or more-or-less spherical ascoma (Korf, 1973). The ptychothecia may be rather loosely folded, creating empty chambers, or more compact, with veins of fertile tissue opening externally. Trappe (1979) states that there are six hypogeous to subhypogeous Peziza taxa reported from North America, Europe, Australia, and New Zealand. Castellano et al. (1989) report that there are at least three hypogeous species in North Temperate forests. Two of these in western North America. All are ptychothecial. The stereothecial Cazia flexiascus Trappe was recently transferred to the Pezizaceae after 18S and 28S rDNA _ sequences were analyzed (ODonnell et al., 1997). The Pezizaceae thus contains apothecial, ptychothecial and stereothecial taxa. Asci forcibly discharge spores or are indehiscent. It is tempting to hypothesize an evolutionary series from apothecial Peziza taxa through ptychothecial taxa with a small number of large open chambers (e.g., P. quercicola), ptychothecial taxa with complex folding and small chambers (e.g., P. stuntzii) culminating in stereothecial taxa (e.g., Cazia). A similar evolutionary "series" has been reported in_ the Pyronemataceae from apothecial taxa (e.g., Humaria, Hydnocystis) to ptychothecial taxa (e.g., Geopora, Labyrinthomyces, Paurocotylis, Petchiomyces). It is also tempting to hypothesize that loss of active spore discharge is correlated with ascomata type. Trends in spore discharge are unclear in the Pezizaceae. The epigeous ptychothecial taxa have forcible spore discharge. The mechanism in P. stuntzii is unclear although it has indented tips structured for opercula. P. quercicola is operculate, but spore release, "puffing", has not been observed. The mechanism in Cazia is unknown, although the rounded, entire tip in the micrograph published by Trappe (1989) suggests it is indehiscent. No correlation exists between ascomatal type and spore discharge mechanism in the Pyronemataceae. Geopora forcibly discharges spores. Hydnocystis, Labyrinthomyces, Paurocotylis and Petchiomyces have indehiscent asci. Clearly, the presence of indehiscent asci and epigeous ascomata are not sufficient on their own to separate taxa into families. Although these extant taxa may represent steps in an evolutionary series from apothecia containing operculate asci to stereothecia containing indehis- cent asci, they are the result of evolution not "missing links." Elucidation of evolutionary relationships within the Pezizaceae and Pyronemataceae awaits phylogenetic analysis utilizing molecular data, at least in part. 82 ACKNOWLEDGMENTS This study was supported in part by grants from the National Science Foundation (BSR 90-24572, DEB 94-00871). The generous assistance of Clark T. Rogerson is also gratefully acknowledged. The authors also wish to thank Drs. Nancy Smith Weber and Bradley Kropp for their comments on the manuscript. LITERATURE CITED Arora, D. (1986). Mushrooms demystified. Ten Speed Press, Berkeley, California. Brown, D. E. (1994). Biotic Communities: Southwestern United States and Northwestern Mexico. Univ. Utah Press, Salt Lake City, Utah. Castellano, M. A., J. M. Trappe, Z. Maser, and C. Maser. (1989). Key to spores of the genera of hypogeous fungi of North Temperate forests with special reference to animal mycophagy. Mad River Press, Eureka, California. Cronquist, A., A. H. Holmgren, N. H. Holmgren, and J. L. Reveal. (1972). Intermountain flora; vascular plants of the Intermountain West, U.S.A. Vol. 1. Hafner Press, New York. Fogel, R. (1994). Materials for a hypogeous mycoflora of the Great Basin and adjacent cordille- ras of the Western United States. II: Two sub-emergent species: Cortinarius saxamon- tanous sp. nov. and C. magnivellatus, plus comments on their evolution. Mycologia 86:795-801. Fogel, R. and G. Pacioni. (1989). Materials for a hypogeous mycoflora of the Great Basin and adjacent Cordilleras of the western United States. Mem. New York Bot. Gard. 49: 119- 128. Gilkey, H. M. (1954). Tuberales. North American Flora 2, 1: 1-36. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. (1990). Index Herbariorum Part I: The Herbaria of the world. 8 ed. Regnum Veg. 120: 1-693. Kelly, K. L. and D. B. Judd. (1965). The ISCC-NBS method of designating color and dictionary of color names. Natl. Bur. Standards, Circ. No. 553, U. S. Government Printing Office, Washington, D.C. Korf, R. P. (1973). Sparassoid ascocarps in Pezizales and Tuberales. Rep. Tottori Mycol. Inst. 10: 389-403. O”Donnell, K. O., E. Cigelnik, N. S. Weber and J. M. Trappe. (1997). Phylogenetic relation- ships among ascomycetous truffles and the true and false morels inferred from 18S and 28S ribosomal DNA sequence analysis. Mycologia 89: 48-65. Smith, A. H., and S. M. Zeller. (1966). A preliminary account of the North American species of Rhizopogon. Mem. New York Bot. Gard. 14: 1-177. Spaulding, W. G. (1990). Vegetational and climatic development of the Mojave Desert: The last glacial maximum to the present. In: Packrat middens: The last 40,000 years of bi- otic change. Edited by Betancourt, J. L., T. R. Van Devender and P. S. Martin. Univ. Arizona Press, Tucson. pp. 166-199. Trappe, J. M. (1979). The Orders, Families and Genera of hypogeous Ascomycotina (truffles and their relatives). Mycotaxon 9: 297-340. Trappe, J. M. (1989). Cazia flexiascus gen. et sp. nov., a hypogeous fungus in the Helvellaceae. Mem. New York Bot. Gard. 49: 336-338. Wells, P. V. (1983). Paleobiogeography of montane islands in the Great Basin since the last glaciopluvial. Ecol. Monogr. 53: 341-382. MYCOTAXON Volume LXXXI, pp. 83-89 January-March 2002 MATERIALS FOR A HYPOGEOUS MYCOFLORA OF THE GREAT BASIN AND ADJACENT CORDILLERAS OF THE WESTERN UNITED STATES. VIII: PACHYPHLOEUS LATERITIUS SP. NOV. AND CAZIA QUERICOLA SP. NOV. (ASCOMYCOTA, PEZIZALES) Robert Fogel University of Michigan Herbarium, North University Building, Ann Arbor, MI 48109-1057 Jack States Botany Department, University of Wyoming, Laramie, WY 82071! ABSTRACT Two new taxa, Pachyphloeus lateritius and Cazia quericola, associated with Quercus turbinella and Pinus edulis from southwestern Utah, United States, are described and illustrated. Key Words: truffle, Utah, biogeography. This is the eighth in a series containing materials for a hypogeous mycoflora of the Great Basin and adjacent Cordilleras (Fogel 1994, Fogel and Pacioni, 1989). The region has received little attention from mycologists despite its potential for studies of fungal biogeography, evolution, and coevolution. The Great Basin, as defined by Cronquist et al. (1972), occupies an area of 714,854 square km. The species composition of a specific stand reflects the interaction of environmental optima for species, soil factors, pre- glacial flora, extent of glacial advance, pluvial lake levels, and subsequent re-invasion from glacial refugia (Wells, 1983). The vegetation in the region contains many suitable ectomycorrhizal hosts for hypogeous fungi. One of the most interesting and difficult habitats to collect is the Interior Chaparral (133.3) vegetation-type (Brown 1994). This vegetation-type is widely scattered along the forest islands comprising the axial route for conifer re-invasion of the Great Basin in southwestern Utah and extends south into Arizona (Wells, 1983). The stands are usually open becoming quite dense along permanent streams. The larger shrubs reach 2 to 2.5 m in height. Shrub live oak (Quercus turbinella Greene), a putative host for ectomycorrhizal truffles and false-truffles, is one of the dominant shrubs. Collecting in these stands has produced a number of interesting truffles including the new Pachyphloeus and Cazia taxa described below and Peziza quercicola described earlier (Fogel and States, 2002). ' Address for correspondence: 2 Canyon Shadows Rd., Lander, WY 82520. 84 The apparent high degree of endemism suggested by the three new taxa being from the same stand may only reflect the lack of collecting in this vegetation type. — MATERIALS AND METHODS Methods of collection and study of the morphological and anatomical features are essentially those of Smith (Smith and Zeller, 1966). Spores were measured in freehand sections mounted in Melzer’s reagent. Lengths and widths of 20 ascospores randomly selected from a mature ascoma in each collection were measured for statistical comparisons. Color names are given as the ISCC-NBS names with the numerical equivalent in parentheses (Kelly and Judd, 1965). Herbarium names are abbreviated according to Holmgren, Holmgren, and Barnett (1990). TAXONOMY PACHYPHLOEUS LATERITIUS Fogel & States sp. nov. Figs. 1-4 Pytchothecia usque ad 33 x 28 mm, subglobosum ad cerebriformum, sulcata. Peridium glabrum, lateritium. Gleba album. Asci saccati, quadrispori ad octospori, non-amyloidei. Sporae globosoideae, 18-27 um, ornamentis echinulatum, <0.5 um, hyalinae, uni-guttulatum. Paraphyses 4 um in diam., apicibus rotundatum. Holotypus: USA, Utah: Kane Co., 13 May 1995, Fogel F4887 (MICH). Etymology: lateritius (L.) dark brick color, in reference to the color of the peridium. Pytchothecia subglobose to craniform, deeply furrowed with invaginations on apices, up to 33 mm wide x 28 mm high, light yellow brown (ISCC-NBS 76), strong brown (ISCC-NBS 55) to deep brown (ISCC- NBS 56) with strong red tones, to dark grayish brown (ISCC-NBS 62), granulose with small dark brown warts, sparsely tomentose. Gleba solid, white (ISCC-NBS 263), veins suggested in dried pytchothecia by slight depressions or wrinkles. Hymenium irregular, interrupted by sterile veins. Attached by a single 0.5 mm diam. rhizomorph. Odor not noted. Spores globose, 18-27 um diam., averaging 19.95 + 1.00 um, hyaline, immature spores dextrinoid, ornamented with mucilage-embedded spines <0.5 um long, wall 1 um thick including mucilage. Asci broadly clavate, 85- 120 x 35-45 um wide, non-bluing (I), immature asci dextrinoid, containing 4 uniseriate to 8 biseriate spores, base stalked, buried in sinuous 4 um diam. septate paraphyses. Tips of paraphyses rounded, not swollen, projecting beyond the asc. Ectal excipulum 80-100 um thick, composed of subglobose to radially elongate, 17-35 um long x 12-30 um wide, thick-walled (1 um) cells, walls dull orange to brownish black (KOH and I). Parenchymous below, cells hyaline, grading into a 100 um thick layer of periclinal, hyaline, 4-5 um diam. hyphae, some cells inflated to 8 um. Content of some cells in the 85 if Figs. 1-4. Pachyphloeus lateritius. Fogel F4981 (MICH). Fig. 1. Ectal excipulum and gleba mounted in Melzer’s reagent, 118X. Fig. 2. Thick walled cells on surface of ectal excipulum mounted in KOH, 640X. Fig. 3. Asci mounted in KOH, phase contrast, 235X. Fig. 4. Spores mounted in Melzer’s reagent, 1600X. 86 center of sterile veins dull orange (I). Warts 100 um wide x 80 um high composed of thick-walled dull orange, elongate cells. A few thick-walled (1 um), straight, 6-8 um diam. hairs projecting from the peridium. Putative mycorrhizal hosts and phenology: Fruiting from mid May to late June at 1300-1400 m elevation in southwestern Utah. Two of the collections were hypogeous under shrub live oak (Q. turbinella) in the Interior Chaparral (133.3) vegetation-type (Brown, 1994). One collection was hypogeous under Rocky Mountain pifion pine (Pinus edulis Engelm.) in. the Great Basin Conifer Woodland (122.4) vegetation-type, one of the most common types in the Intermountain Region (Brown, 1994). Material examined: USA, Utah, Kane Co., Fogel F4969. Washington Co., Fogel F4887 (Holotype), F4981 (all MICH). Discussion: Pachyphloeus lateritius is easily mistaken for a Balsamia due to the reddish brown color and warted peridium. Under the microscope, the globose spores easily distinguish this taxon from Balsamia. Three species of Pachyphloeus are known from North America (Gilkey, 1954; Castellano et al., 1989). Two of them (P. virescens and P. melano- xanthus) have green to greenish-black ascoma. P. citrinus has orange to brown ascoma. The spores of the latter have low, truncate papillae clearly distinct from the echinulate spores of P. lateritius. P. citrinus also has larger diam. paraphyses (8-13 vs. 4 um) and smaller globose spores (8-13 vs. 18-27 um). CAZIA QUERICOLA Fogel & States Figs. 5-10 Sporocarpia usque ad 35 x 30 x 28 mm, cerebriformum, prope album, sub- tomentum. Gleba album ad pallide violaceo-grise. Asci 120-170 x 14-30 um, sporis 8, irregulatim cylindrici vel saccati, plerumque flexi, hyalini, non-amyloidei. Sporae 11-14 x 10-12 ym, subglobosae, hyalinae, minute verrucose reticulatae, ornamentum <0.5 um altum. Holotypus: USA, Utah: Washington Co., 22 May 1995, Fogel F4909 (MICR). Etymology: cola (L.) dweller, hence dweller in Quercus forests. Stereothecia up to 35 x 30 x 28 mm high, craniform with several basal lobes; white (ISCC-NBS 263) bruising brownish orange (ISCC-NBS 54); subtomentose, larger stereothecium with cracks from rapid expansion; attachment not evident. Gleba dense, very firm but sectioning easily, fertile Figs. 5-9. Cazia quericola. Fig. 5. Sectioned sporocarp, 1.5X, Fogel F4878 (MICH). Fig 6. Asci mounted in KOH, note crozier, phase contrast, 1400X, Fogel F4909 (MICH). Fig. 7. Uniseriate and biseriate asci and septate paraphyses mounted in KOH, note bending of asci, phase contrast, X166, Fogel F4909. Fig 8. Spores mounted in Melzer’s reagent, 1600X, Fogel F4909. Fig. 9. Isodiametric cells in trama, squash mount in Melzer's reagent, 1600X, Fogel F4909. 87 88 areas white (ISCC-NBS 263) to pale reddish purple (ISCC-NBS 244), empty or stuffed with white hyphae dried. Trama white, drying moderate to dark orange yellow (ISCC-NBS 71-72), radiating outward from basal patch, 300- 350 um wide, composed of hyaline, thin-walled, subparallel hyphae; 6-10 um wide at septa, cells inflated to 18 um. Color changes on application of chemicals not noted. Attachment not obvious. Odor faintly Tuber-like. Spores globose to mostly subglobose 11-14 x (9-)10-12 um averaging 12.2 + 0.8-0.9 x 10.9 + 0.9 um to 11.6 + 0.5, Length/Width ratio 1.06-1.12, hyaline, uniguttulate, immature spores dextrinoid, incompletely warty- reticulate, warts <0.5 um high, non-amyloid, wall 1-1.5 um thick including ornamentation. Asci in pockets of thin-walled, 40-65 um diam. cells; asci cylindrical to saccate, mostly bent, 120-170 x 14-30 um, immature asci dextrinoid; 8-spored, uniseriate to mostly irregularly biseriate; base tapered, obscurely croziered. Paraphyses 6-8 um diam, septate, terminal cells 20-25 um long, tips swollen to 10 um. Peridium 175-425 um thick, outer layer of periclinal, dextrinoid, 5-8 um diam hyphae, some cells swollen to 15 um diam; interrupted occasionally by 20-30 um isodiametric cells. Cells smaller near the surface and more parenchymous Putative mycorrhizal hosts and phenology: Fruiting mid to late May at 1400 m elevation in southwestern Utah. Hypogeous under shrub live oak (Q. turbinella) in the Interior Chaparral (133.3) vegetation-type (Brown, 1994). Material examined: USA, Utah, Washington Co., Fogel F4878, F4909 (Holotype) (all MICH). Discussion: Fruiting in the same stand of Q. turbinella as Pachyphloeus lateritius. The color of the gleba may depend on maturity. The immature gleba was pale reddish purple, the mature gleba was white. The gleba of C. flexiascus is described by Trappe (1989) as "light purplish gray (ISCC-NBS 232)". Cazia may be associated with oaks as both C. quericola and C. flexiascus were collected under Quercus species. Differentiated from C. flexiascus by having smaller subglobose, rather than ellipsoid, spores (11-14 x 10-12 um vs. "13-16 x 10-12 um"; L/W 1.1 vs. 1.3). The paraphyses are also narrower (6-8 im, apices swelling to 10 um vs. "7-10 tm", apices swelling to "15 um") and peridium thicker (175-425 vs. "100-150 tm"). ACKNOWLEDGMENTS This study was supported in part by grants from the National Science Foundation (BSR 90-24572, DEB 94-00871). The generous assistance of Clark T. Rogerson is also gratefully acknowledged. The authors also wish to 89 thank Dr. Bradley Kropp (Utah State University) for his comments on the manuscript. LITERATURE CITED Brown, D. E. (1994). Biotic Communities: Southwestern United States and Northwestern Mexico. Univ. Utah Press, Salt Lake City, Utah. Castellano, M. A., J. M. Trappe, Z. Maser, and C. Maser. (1989). Key to spores of the genera of hypogeous fungi of North Temperate forests with special reference to animal mycophagy. Mad River Press, Eureka, California. Cronquist, A., A. H. Holmgren, N. H. Holmgren, and J. L. Reveal. (1972). Intermountain flora; vascular plants of the Intermountain West, U.S.A. Vol. 1. Hafner Press, New York. Fogel, R. (1994). Materials for a hypogeous mycoflora of the Great Basin and adjacent cordille- ras of the Western United States. Il: Two subemergent species: Cortinarius sax- amontanous sp. nov. and C. magnivellatus, plus comments on their evolution. Mycologia 86:795-801. Fogel, R. and G. Pacioni. (1989). Materials for a hypogeous mycoflora of the Great Basin and adjacent Cordilleras of the western United States. Mem. New York Bot. Gard. 49: | 19- 128. Fogel, R. and J. States. (2002). Materials for a hypogeous mycoflora of the Great Basin and adjacent cordilleras of the Western United States. VII: A new truffle-like Peziza (Ascomycota, Pezizales). Mycotaxon. 81: 75-82. Gilkey, H. M. (1954). Tuberales. N. Amer. FI., Ser. II., Part 1. pp. 1-29. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. (1990). Index Herbariorum Part I: The Herbaria of the world. 8" ed. Regnum Veg. 120: 1-693. Kelly, K. L. and D. B. Judd. (1965). The ISCC-NBS method of designating color and dictionary of color names. Natl. Bur. Standards, Circ. No. 553, U. S. Government Printing Office, Washington, D.C. Smith, A. H., and S. M. Zeller. (1966). A preliminary account of the North American species of Rhizopogon. Mem. New York Bot. Gard. 14: 1-177. Trappe, J. M. (1989). Cazia flexiascus gen. et sp. nov., a hypogeous fungus in the Helvel- laceae. Mem. New York Bot. Gard. 49: 336-338. Van Devender, T. R. (1990). Late Quaternary vegetation and climate of the Sonoran Desert, United States and Mexico. In: Packrat middens: The last 40,000 years of biotic change. Ed. by Betancourt, J. L., T. R. Van Devender and P. S. Martin. Univ. Arizona Press, Tucson. pp. 134-165. Wells, P. V. (1983). Paleobiogeography of montane islands in the Great Basin since the last glaciopluvial. Ecol. Monogr. 53: 341-382. 7 art = RIE 1 A ae 4 Y Ae - ef Gee Oras» a a REPAY GRRL SR ea Ahi oe he Ae uy ; ee ome s is eee Le ND a Sea es 4 Sue i py in’ ud Ay, Wy 4 seu. A. » oe bay i wn TAP as nlf ne Feat a tha, a, ae vee, Ce eat hg a L wets ; | 4 , 2) fh mY F i. 7 A aa GO uteeas5 ¢ ay oe, Stir MM ; Te ean he 7 SPER A FEM AME \o ges Gre te Ar ry 1 ' MY COTAXON Volume LXXXtI, pp. 91-94 January-March 2002 A NEW INONOTUS (BASIDIOMYCOTA, HYMENOCHAETACEAE) FROM HAWAII Robert L. Gilbertson Department of Plant Pathology, University of Arizona, Tucson, AZ 85721 and Leif Ryvarden Department of Botany, Biol. Inst., University of Oslo, Oslo, Norway ABSTRACT Inonotus hemmesii is described as a new species causing a white heartrot of living hardwoods in the Hawaiian Islands. It is characterized by large pileate basidiocarps, frequent setal hyphae in the tube trama, rare hymenial setae, and pigmented, ovoid to ellipsoid basidiospores. This paper reports a previously undescribed species in the genus Jnonotus P. Karst. It is associated with a white heartrot of koa and some introduced hardwoods in the Hawaiian Islands. Current records are from the islands of Hawai‘i, Moloka‘i, and O‘ahu, and it is particularly common on manele (Sapindus saponarius L.) on the island of Hawai‘i. INONOTUS HEMMESITI Gilb. & Ryvarden, sp. nov. Fig. 1 Fructificatio annua, pileata, sessile vel effuso-reflexa; systema hypharum monomiticum; hyphae setarum frequenter adsunt in trama; setae hymeniarum rara; basidia late ellipsoideae, 4-sterigmatibus, 12-15 x 8-10 um; basidiosporae ovoideae vel ellipsoideae, pallido-brunnea, crassitunicatae, 7.5-9 x 5.5-7 um; putrido alba. Holotype: RLG 22943, on Delonix regia, Kaunakakai, island of Moloka‘i, Hawai‘i, Jan. 7, 2000, in herb. BPI, Beltsville, MD. Etymology: named in honor of Dr. Don E. Hemmes, University of Hawai‘i at Hilo, who has contributed greatly to knowledge of Hawaiian fungi. Basidiocarps annual, pileate, sessile to effused-reflexed, up to 30 x 20 x 15 cm; pilear oe Q 49 40> Fig. 1. Microscopic characters of Inonotus hemmesii (RLG 22943). a, contextual hyphae; b, tramal hyphae; c, tramal setal hyphae; d, hymenial setae; e, basidia, f, basidiospores. 95 surface brown, shallowly sulcate, zonate, finely tomentose at first, becoming glabrous to crustose and finally blackened and rimose with age; pore surface yellowish brown at first, becoming darker brown with age, the pores angular, regular, 3-4 per mm, dissepiment edges finely granular under 30x lens; context tough-corky, with a thin tan upper layer up to 2 mm thick and a dark chocolate brown lower layer up to 6 cm thick; tube layer distinct, yellowish-brown, up to 4 cm thick. Hyphal system monomitic; contextual generative hyphae pale yellowish to darker yellowish brown in KOH, thin- to firm-walled, simple-septate, of two types; predominant type long, rarely branched, 7-10 um in diam, these interlaced with narrower, much- branched and contorted hyphae 3-5.5 wm in diam; tramal hyphae of one type, pale golden brown, parallel, with occasional branching, thin- to firm-walled, simple-septate, 3-5.5 pm in diam. Setal hyphae frequent in trama, absent in context, becoming thick-walled, dark reddish brown in KOH, 7-10(-12) wm in diam and up to 170 ym long, tapering to a point, not projecting into hymenium. Hymenial setae rare, absent in many sections, ventricose, thick-walled, pointed, dark reddish brown in KOH, 20-25 x 7.5-10 ym. Basidia broadly ellipsoid, 4-sterigmate, 12-15 x 8-10 um, simple-septate at the base. Basidiospores invariably present in large numbers, even in old specimens, ovoid to ellipsoid, pale golden brown in KOH, smooth, becoming thick-walled, negative in Melzer’s reagent, 7-9 x 5.5-7 um. Specimens examined: RLG 17634, on koa (Acacia koa A. Gray), Keanakolu Rd., North Hilo Dist., Hawai‘i, Aug. 29, 1991; RLG 18037, 18046, on manele (Sapindus saponarius L.), Kipuka Ki, Hawai‘i Volcanoes National Park (HVNP), Ka‘t Dist., Hawai‘i, Sept. 22, 1991; RLG 18364, 18369, on koa, Manoa Cliffs Trail, Honolulu Dist., O‘ahu, Oct. 6, 1991; RLG 18728, on manele, Kipuka Ki, HVNP, Ka‘t Dist., Hawai‘i, Oct. 24, 1991; RLG 19355, on manele, Kipuka Ki, HVNP, Ka‘u Dist., Hawai‘i, Dec. 5, 1991; RLG 21419, on royal poinciana (Delonix regia (Bojer) Raf.), Kaunakakai, Moloka‘i, Jan. 13, 1996; RLG 22363, on manele, Kipuka Ki, HVNP, Ka‘t Dist., Hawai‘i, Nov. 22, 1998; RLG 22943, on royal poinciana, Kaunakakai, Moloka‘i, Jan. 7, 2000 (TYPE). Inonotus hemmesii is close to I. rodwayi D.A. Reid (Reid, 1957) but differs in having wider, more thick-walled basidiospores. Also, the setal hyphae in the trama of J. hemmesii are longer, consistently imbedded and not projecting into the tubes. Those of I. rodwayi are shorter and often project into the tubes. /nonotus patouillardii (Rick) D.A. Reid is also similar but the setal hyphae of that species are abundant, very conspicuous in all sections, and often project up to 100 wm beyond the hymenium (Gilbertson and Ryvarden, 1986). Setal hyphae of J. hemmesii are frequent but not abundant and may be absent in some sections. Also, the basidiospores of J. patouillardii (4.5-6 x 3.5-4.5 um) are smaller than those of J. hemmesii (7-9 x 5.5-7 pm). 94 ACKNOWLEDGMENTS Mahalo to Don Hemmes for his expertise and the pleasure of his company in field work in the Hawaiian Islands over the past eleven years. LITERATURE CITED Gilbertson, R.L. and L. Ryvarden. 1986. North American Polypores. Vol.1: 1-433. Abortiporus-Lindtneria. Fungiflora. Oslo. Reid, D.A. 1957. New or interesting records of Australian basidiomycetes: III. Kew Bull. 12:127-143. - MY COTAXON Volume LXXXI, pp. 95-112 January-March 2002 PHAEOCOLLYBIA IN WESTERN NORTH AMERICA 3: TWO NEW SPECIES WITH NOTES ON THE PHAEOCOLLYBIA FESTIVA COMPLEX’ LORELEI L. NORVELL’ Pacific Northwest Mycology Service 6720 NW Skyline Boulevard, Portland OR 97229-1309 USA Abstract: Morphological and molecular reevaluation of the genus Phaeocollybia has uncovered the existence of two new forest agarics from British Columbia, Washington, Oregon, and California. Phaeocollybia phaeogaleroides produces small, fragile basidiomes with large, lightly ornamented, ellipsoid basidiospores, abundant clamp connections, and a unique fruiting phenology. First revealed during molecular analyses, Phaeocollybia rifflipes produces small basidiomes with tawny viscid pilei and lilac-colored lamellae. It shares affinities with two western species (Phaeocollybia fallax and Phaeocollybia lilacifolia) with glutinous pilei, lilac lamellae, vertical-monopodial pseudorhizae, limoniform basidiospores, and thin-walled clavate cheilocystidia. The close morphological relationship of three western species to the European Phaeocollybia festiva is discussed. Key words: Agaricales, Basidiomycota, Cortinariaceae, RFLP, temperate rainforest Introduction Over one thousand Phaeocollybia (Basidiomycota, Cortinariaceae) collections have been made from the northern temperate rainforests of the western American coast as part of acontinuing study of the genus begun in 1991. This research has resulted in the discovery of generically significant morphological characters (e. g. universal pellicular sheath, different pseudorhizal morphologies, sarcodimitism, cf. Norvell 1998ab), molecular support for the existence of 25 species (inferred from restriction digests of 160 collections, cf. Norvell 1998a, 2000), new Based in part on material submitted for a doctoral dissertation at the Department of Botany, Box 355325, University of Washington, Seattle WA 98195-5325 USA. * Email: <lorelei@teleport.com> 96 insights into the development and biology of the genus (Norvell 1998ab), and naming of five new species (Norvell 2000, Norvell & Redhead 2000). Two additional new species -- P. phaeogaleroides and P. rifflipes -- are proposed and described, and P. rifflipes is compared to species sharing affinities with P. festiva (Fr.) Heim. Materials and Methods Specimen collection and examination, macrochemical tests, DNA extractions, PCR amplification of the ITS1+5.8S+ITS2 rDNA region (hereafter referred to as the “ITS region”), and phenetic and phylogenetic analyses were conducted using protocols previously summarized in Norvell (1998ab, 2000). Descriptions are based on data taken from both type and other well-documented collections. Generic or developmental terms (e.g. tibiiform diverticula, sarcodimitic tissues, pellicular veil, vertical-monpodial) are fully explained in Norvell (1998ab). General non-standardized color names in lower case are accompanied by italicized color references. Ridgway (1912) colors are capitalized and abbreviated, with a slash (/) or parentheses separating modifiers of the same base color (e.g. Mikado /Mars /Verona Brown = Mikado Brown, Mars Brown, Verona Brown; (Pale) Pinkish Cinnamon = Pale Pinkish Cinnamon and Pinkish Cinnamon); bracketed Munsell (1976) alpha-numeric color ranges (e.g. [2.5Y 6- 8/1-4]) follow Ridgway (1912) references. Measurements of anatomical characters were taken from tissues rehydrated in 6% aqueous KOH unless otherwise noted, and basidiospores from the stipe apex were measured when spore prints were lacking. Mean basidiospore dimensions with standard deviations are accompanied by ranges enclosed in parentheses. “L+ll/cm” is the number of lamellae + lamellulae per cm at the “edge” of the pileus and at the “midpoint” between edge and stipe. “Aerial stipe” refers to the portion of the stipe above the ground; “origin” refers to the pseudorhizal origin at the very base of the basidiome. Herbaria housing cited collections are abbreviated in accordance with Holmgren et al. (1990), with “PNW” representing the Pacific Northwest Mycology Service herbarium. RLFP codes are denoted by subscripts, and asterisks (*) flag Northwest Forest Plan Strategy 3 collections (USDA-USDI 1994, Norvell 1998c) that are of significance to US forest management policy. Separate collection dates are not listed for the date-based numbered Norvell collections e.g. LLN1921104-2,,,,, = Norvell November 4, 1992 collection 2, RFLP code riu/0). Taxonomic Descriptions Phaeocollybia phaeogaleroides Norvell sp. nov. Figures 1-2 Pileus 13-50 mm latus, e convexo campanulatus et umbonatus, glaber, viscidulus, hygrophanus, e fulvo ferrugineus; caro tenue, pallida; odor et sapor lenis; lamellae pallidae vel luteae, fulvo- brunnescentes e sporis; stipes e farcto subcavus, 1.5-4 mm crassus, 60-90 mm longus supra terram usque ad 180mm longus cum pseudorhiza attenuata, glaber, apice lamellis concolorus et subapice ferruginascentes. Sporae magnae, 9-12 X 5-6.2 jum, ellipsoideae, verruculosae Ficure 1. Phaeocollybia phaeogaleroides. Fic 1a. LLN1990413-01 (PNW) in situ in 50-year old mixed forest in Benton County, Oregon. Note breaks on fragile stipe caused by removal of debris around specimen. Scale bar = 5 mm. Ha 1B. Clamp connections on hyphae in pileus suprapellis in collection from Drift Creek, Oregon (LLN1961101-01, WTU). Scale bar = 10 microns. Fic Ic. Small basidiome (LLN1990421-04, PNW) from Green Peak, Oregon with Douglas-fir branch and moss (Eurhynchium oreganum). Fic Ip. Detail of mycenoid holotype specimens (Ex199-064, OSC & PNW) from Alsea Falls, Oregon, showing moist striate pilei. Scale bar = 5 mm. 98 Ficurer 2. Phaeocollybia phaeogaleroides. Ric 24. Longitudinal section of mature expanded basidiome with dark, moist cuticle (left, adapted from LLN1961106-01 [WTU], collected near Drift Creek, Oregon) contrasts with slightly less mature specimen with fading, hygrophanous pileus (right, adapted from ROuterbridge298 [DAOM], collected. on Vancouver Island, British Columbia). Scale bar = 5mm. LLN1951106-01 [WTU]. Fics 2p-p. Scale = 10 microns. Hc 28. Basidiospores, basidium, and basidiole. Under low magnification, the large, punctate- roughened basidiospores appear almost smooth while the apical callus might be mistaken for a germ pore. Hc 2c. Thin-walled cylindrical to clavate cheilocystidia with clamp connections. Kc 2p. Tibiiform diverticula emanating from hyphae of the sitipitipellis. 99 subtiliter, cum apice glabro et rotundato vel parum acuto, cum apiculo excentrico glabro et cum plaga obscura; ixocutis cum suprapelle aliquanto, subgelatinosa, hyalina et subpelle cum pigmento subluteo; cheilocystidia cylindracea vel subcapitata, leptoderma, hyalina vel sublutea, 3-5 jum lata; paracystidia filiforma. Hyphis in suprapelle, cheilocystidiae et basidiae fibulati. Diverticula tibiiforma longissima, ad 50 ym longa. Typus ad terram in musco in sylvis Pinacearum mixtarum: R. L. Exeter Ex199-64, December 7, 1999, Finley Hill BLM Reserve Forest, Mary’s Peak RNA, Benton County, Oregon, USA (Holotypus OSC; Isotypi WTU, PNW). Etymology: from morphologies shared with the genus Phaeogalera (general aspect, basidiospores). BRIEF SUMMARY — Basidiomes small, fragile, rooting; pilei thin-fleshed, conic to broadly campanulate with low umbo, subviscid, striate, hygrophanous, dark chestnut brown but soon fading to honey-tan; young lamellae pallid to yellow- tan; stipes thin, fragile, cartilaginous, polished, tan to dark brown; pseudorhizae vertical-monopodial with yellowish origins; basidiospores large (~10 X 5.8 ym), ellipsoid with rounded to very slightly pointed (not beaked) apices, faintly verruculose-roughened; cheilocystidia cylindrical to slender subcapitate with slightly thickened and slightly refractive walls; pileipellis bilaminate, with a colorless suprapellis of clamped narrow hyphae overlying a pale orange-yellow subpellis; clamp connections abundant, medallion-like; tibiiform diverticula unusually long (up to 50 ym). All tissues negative in syringaldazine. EXPANDED TECHNICAL DESCRIPTION — Pileus 7-50 mm diam, when young conic-convex with prominent to obscure obtuse umbo, when mature broadly campanulate with/without acute/low umbo and uplifted margin with cuticle extending beyond lamellae at edge; disc sometimes radially rugulose, surface glabrous, moist to subviscid, hygrophanous, when moist long pellucid striate and dark red-brown (Clay Color [2.5 YR 4-5/6, 5YR 4/4-6, 1OYR 3-4/2-4), when dry soon fading to honey-tan overall (Warm Buff, Yellow Ochre [10YR 5-6/4-6]). Context: thin (2 mm at the disc), pallid to pale tan. Odor not distinctive to very slightly raphanoid. Taste not distinctive. Lamellae free to narrowly sinuate attached, subdistant to close, polydymous with 3-5 tiers of +/- regularly interspersed lamellulae, ventricose, 2-6 mm broad (length:width ratio 3.8), thin, edges even to slightly serrulate, pallid, when young pale honey-tan or pallid (Warm Buff, [10YR 8/4]), when mature pale rusty brown (//0 YR _ 5/4]). Primordial sheath remnants occasionally evident as scattered fibrillose patches on stipe. Stipe central, terete, slender, cartilaginous rigid but fragile (mycenoid), aerial length 60-90 mm, combined length including pseudorhiza up to 180 mm, apex |.5-4 mm diam, more or less equal to ground level, glabrous, dry, slightly polished, young apex honey tan (Warm Buff [2.5Y 5/4, 1OYR 6-7/4-6]), below grading to vinaceous brown (2.5 YR 5/6, 7.5 YR 4/2, IOYR 3/2, IOYR 5/10); stuffed when young, cortex thin (0.5 mm) and cartilaginous, context loosely fibrillose, pallid to pale yellow when young; fistulose in age. Pseudorhiza vertical-monopodial, 1/2 - 2/3 overall stipe length, fragile, 100 slightly enlarged just below ground level before gradually tapering to pale yellow ([2.5 YR 6-8/4-6]) origin. Spore print: reddish brown (Cacao Brown, [2.5YR 4/4-6]]). Basidiospores 10 X5.8 +0.6 x 0.4 ym (overall range 9-12 (13) x 5-6.5 yum), terete or slightly compressed, ellipsoidal with eccentric apiculus and rounded to slightly pointed (unbeaked) apex; very finely verruculose roughened, the suprahilar disc an ill-defined area of lowered ornamentation faintly discernible in oil immersion; amber in KOH, yellowish in H,O, inamyloid and nondextrinoid in Meizer's. Cheilocystidia emanating from the lamellar trama, cylindrical to subcapitate, long (up to 80 um), apices either 2-4 ym diam or (occasionally) forming 3-5 um heads above 2-3 pm diam slightly refractive necks, walls slightly thickened, hyaline to pale orange yellow in KOH, septa with medallion-like clamps, + agglutinated in revived material. Pleurocystidia occasional to infrequent, filamentous or sparsely forked, readily collapsing in mounts and hence very obscure, particularly in revived material. Basidia (2-) 4-spored, 28-32 x 8- 10 ym, clavate, hyaline, basally clamped. Pileipellis bilaminate; the 50-100 pm thick, colorless suprapellis composed of cylindrical, narrow (2-4 ym), thin- walled, (sub)gelatinized, hyaline hyphae, clamp connections frequent; the 150- 200 ym thick, pale orange-yellow subpellis composed of 4-8 wm wide gelatinized hyphae. Stipitipellis bilaminate, but not well delimited; the colorless suprapellis hyphae tightly packed, 2-4 wm wide, subgelatinized, hyalinel; the subpellis a pale orange-yellow zone of 10-15 mm wide, heavily gelatinized hyphae. Pseudorhizal pellis rusty brown, hyphae 5-10 jm wide and moderately thick-walled (0.5 wm); this layer covered by a remaining colorless primordial pellis composed of diverticulate, 2-3 4m wide, gelatinized hyphae. All tramal tissues gelatinized and containing occasional oleifers; in the pseudorhiza weakly sarcodimitic with long (50-80 ym), wide (10-15 ym), moderately thick walled (0.5 um), pale amber, rigid, vessel hyphae intermixed with thinner-walled, narrower (2-4 ym), hyaline, flexuous hyphae; in the stipe and pileus composed of uniformly thin-walled, 4-8 um wide, pale amber hyphae; in the lamellae parallel, the relatively narrow central region composed of moderately thick-walled (0.5 um), 3-7 um wide, pale orange-yellow hyphae. Tibiiform diverticula found on primordial sheath remnants, frequent on stipitipellis, abundant on pseudorhizal pellis; unusually long (15-50 ym) and wide (1-2 ym) for the genus; usually capitulate with 2-3 wm diam capitellum and/or apical droplet, lacking basal septum; refractive, hyaline. Clamp connections medallion- like, present on basidial bases, cheilocystidia, and suprapellicular hyphae. Macrochemical reactions: Syringaldazine negative. KOH slightly reddish brown. FeSO4 greenish. Fluorescence absent in fresh material; dried lamellae bright yellow orange. RFLP profile not tested 101 Ecology, range, distribution, phenology: solitary, scattered, or closely gregarious in mossy low lying areas near or under mid-/late-successional conifers such as Pseudotsuga menziesii (Mirbel) Franco, Picea sitchensis (Bong.) Carr., Tsuga heterophylla (Raf.) Sarg. Known from 19 collections made in Oregon and British Columbia. Autumn through early summer (October to July). SPECIMENS EXAMINED: CANADA. BritisH CoLumBiIA. Vancouver Isl. Fairy Lake, Hwy 24. PLor UK Fp 98-8 — solitary on ground in 35yo Pseudotsuga menziesii stand, no understory/vegetation in immediate vicinity, 21.VII.1997 ROuterbridgel09_ (DAOM); Prot #2 — 20.X.1997 ROuterbridge298 with SARedhead (DAOM). UNITED STATES. OreEGoN. Benton Co. HNLEY HILL BLM RESERVE, Mary’s PK RNA — in moss-covered loam in second growth Pseudotsuga menziesii, 7.X11.1999, RLExeter Exl199-64 (HOLOTYPE OSC; Isotypes — WTU, PNW); Mary’s PK RNA (T13S R7W S15) — 24.V.2000, through rotten Eurhynchium oreganum covered log or soil duff in 60yo Pseudotsuga menziesii with Acer circinatum, Gaultheria shallon, RLExeter Ex200-008,009,010 (PNW); GREEN Peak, BLM UNIT, Mip Rp 14-6-17.1 (123.44°W,44.37°N) — at 400m in woody duff & abundant Eurhynchium oreganum in thinned 65 yo Pseudotsuga menziesii with no understory, LLEN/990421-04 with RLExeter (PNW); HULL-OAKES SAWMILL MAINLINE Rpd/101-D spur (123.42°W, 44.35°N) — 200m, in rocky clay covered with Eurhynchium oreganum in 5Oyo Pseudotsuga menziesii with Arbutus menziesii Pursh, Corylus cornuta, Polystichum munitum, Synthiris reniformis, LLN1990413-01 with SARedhead (PNW), LLN2000426-01 with RLExeter (PNW). Lane Co. StUSLAWNF: 1.7MI FROM INDIAN CRK RD JCT ON FSR 2127 (T17S, ROW, S2), INDIANCRK CVS por # 1092048 — 7.XI.2000, in soil, Cazares1380 & 1382 by BPerrone (OSC), Cazares138] by RRipley (OSC); 1.7 MILES FROM 37 Rp Jct ON FSR 3230 (TISS R1OW S24), OLD 37 CVS pLot # 2097048 — 8.X1.2000, in soil, Cazares!1390 by DCDickenson (OSC); 2 Mi FROM Buck CRK BRIDGE ON FSR37 (TI5S ROW S11), 37 Rp CVS pLor # 1098048 — 21.XI.2000, in soil, Cazares1484 & 1485 by DCDickenson (OSC). 15 mi N FLORENCE (124,099°W, 43.983°N) — 20.XI.1980 GWright21-3 (MICH, as P. radicata). Lincoln Co. Drirr CREEK — 6.XI1.1995, LLNI951 106-01 by NSWeber (WTU). StUSLAWNF FSRD 5860, 0.2mI Across CANAL CRK (T14S R1IOW S8), CVS PLOT # 1102044 — 1.XI.2000, in soil, Cazares 1347 & Cazares 1362 by DCDickenson (OSC). COMMENTS — | first discovered Phaeocollybia phaeogaleroides during an examination of a collection labeled as P. radicata (Murrill) Singer in the University of Michigan fungal herbarium. Many of its diagnostic characters — small fragile aspect, solitary to scattered habit, occurrence in moss in coniferous habitats, overall tan coloration, and large ellipsoidal non-dextrinoid basidiospores — recall the genus Phaeogalera for which the species is named. Subsequent collections of fresh material revealed that basidiomes exhibit a much deeper color in situ and may also fruit in loosely gregarious clusters. In the field P. phaeogaleroides might be mistaken for a very small, fragile P. attenuata (Smith) Singer or P. pleurocystidiata Norvell & Redhead. Representatives of both those species, however, are typically far more robust, possess heavily ornamented, prominently beaked basidiospores, and lack clamp connections. P. pleurocystidiata is further differentiated by the possession of secretory, thick-walled, tibiiform cheilocystidia and pleurocystidia. The clamp connections of P. phaeogaleroides are among the most conspicuous of any Phaeocollybia studied by this author, and it is the only species tested thus far with fresh gills that do not fluoresce when viewed under UV. Other diagnostic characters 102 include the unusually large tibiiform diverticula and narrow, thin-walled, strangulated and/or subcapitate cheilocystidia. In the two Canadian specimens (ROuterbridge109 & 298), cystidia-like cells appeared to develop wherever the hymenium was damaged, in itself an unusual phenomenon in the genus. Under low power, basidiospores appear almost smooth, additionally seeming to possess apical pores characterizing the genus Phaeogalera. Under oil immersion, however, the faintly verruculose-roughened ornamentation is visible and the ‘pores’ are more properly seen as small callus deposits forming rounded to slightly pointed refractive apices. Large ellipsoidal punctate-roughened basidiospores, rarely found in the genus, also characterize P. christinae (Fries) Heim. While P. christinae also produces small fragile basidiomes with filamentous/subcapitate thin-walled cheilocystidia, it lacks clamp connections and possesses a more brightly colored pileus, slightly shorter, narrower, more highly ornamented basidiospores, and smaller tibiiform diverticula. P. phaeogaleroides is one of very few western Phaeocollybias known to fruit in both autumn and spring; both vernal collections were made in areas where the vernal P. pleurocystidiata was quite abundant. No molecular research has yet been performed on P. phaeogaleroides. Phaeocollybia rifflipes Norvell sp. nov. FiGuREs 3-5 Pileus (10) 20-40 (55) mm latus, e conico-convexo cum papilla acuta conico- campanulatus, umbonatus, viscidus vel glutinosus, glaber, e fulvo olivaceo-brunneus; caro in disco crassa, super lamellas tenue, pallida; odor lenis vel leviter florali; sapor lenis vel subamarus; lamellae violaceae vel avellaneae, brunnescentes e sporis; stipes 4-9 (12) mm versus apicem, 30-50 mm longus supra terram usque ad 165 mm longus cum pseudorhiza attenuata, versus apicem squamulosus, e farcto cavus, versus apicem avellaneus, versus subapicem aurantiascens. Sporae 7-7.5 x 4-4.5 jam, limoniformes, verruculosae vel verrucosae cum rostro acuto, glabro et cum apiculo excentrico glabro; ixocutis cum suprapelle crassa, gelatinosa, hyalina et subpelle cum pigmento intraparietali; cheilocystidia angusto-clavata vel late clavata (5-7 jam usque ad 10 ym lata), gelatinosa, leptoderma, hyalina. Fibulae desunt in texturis omnibus. Typus ad terram in Pinacearum mixtarum, 20 October, 1993, L. L. Norvell cum S. A. Redhead et G. L. Barron, Barlow Pass, Baker-Snoqualmie National Forest, Snohomish County, Washington, USA (Holotypus LLN1I931020-01 WTU; Isotypus, DAOM). Etymology: from RFLP (Random Fragment Length Polymorphism), the molecular analytical technique that first implied the existence of a previously undiagnosed Phaeocollybia. BRIEF SUMMARY — Basidiomes deeply rooting, scattered to gregarious, small to medium sized; pilei convex-campanulate, glutinous, tawny to dark brown; young lamellae pale violet gray to dull pinkish brown; stipes slender, shiny, drab, fibrillose-stuffed to hollow; pseudorhizae fleshy, vertical-monopodial; basidiospores relatively small (~7 x 4.5 ym), limoniform with an apical beak, verruculose; cheilocystidia thin-walled, clavate; pileipellis bilaminate with a thick gelatinous colorless suprapellis overlying an orange subpellis; clamp connections absent. All tissues negative in syringaldazine. Dried pileus 103 Ficure 3. Phaeocollybia rifflipes. Microscale (for Fics 3a, 3c, 3D) = 10 ym; Macroscale (He 3a) = 1 cm. He 3a. Thin-walled clavate cheilocystidia and limoniform basidiospores from a collection from Larch Mountain, Mt Hood National Forest, Oregon (LLN1921104-05, WTU). Fic 3s. Excavated basidiomes displaying vertical-monopodial pseudorhizae typical of most Phaeocollybia species. (Left adapted from LLN1931020-01, Holotype — WTU, from Barlow Pass, Washington; Right adapted from LLN1921023-08, WTU, Olympic National Park, Washington.) Fic 3c. Limoniform beaked basidiospores and thin-walled cheilocystidia found in Holotype. Fic 3p. Basidia, hyphidia (paracystidia), basidioles and basidiospores. LLN1921023- 08 (WTU). 104 carbonized blackish brown. EXPANDED TECHNICAL DESCRIPTION — Pileus (10) 20-40 (55) mm, initially conic-convex with acute papillate umbo, soon conic-campanulate with low broad umbo and incurved to downturned margin with long-inrolled edge; surface glabrous, viscid to heavily glutinous; color when young tawny to dark orange- brown overall or with darker disc (Verona Brown, Russet, Hazel), tawny to brown margin (Hazel, Tawny, Chestnut /Sayal /Verona Brown), and red-/yellow-brown edge (Buffy /Army /Chestnut /Cinnamon Brown), in age more or less uniformly dark brown (Olive Brown) with a dark green-brown (Dark Olive) edge; when dried carbonized black-brown, noticeably striate. Context relatively thick (1-4 mm) at the disc, thinning abruptly over the lamellae; color pallid or buff when young, in age or after exposure brown or drab; often consumed by insects. Odor not distinctive to slightly floral. Taste mild or (occasionally) faintly bitter. Lamellae free to narrowly attached; subcrowded (L+ll/em —15 @ edge, 12 @ midpoint) with 3-5 tiers of lamellulae irregularly interspersed; _-ventricose, moderately broad (3-5 mm), thin; edges even at first, soon uneven to eroded; color pale violet or bluish gray when young (Dark Vinaceous Gray, Pale Vinaceous Drab), dulling as maturing ((Light Brownish) Drab, Avellaneous, Wood Brown, Vinaceous Buff), in age obscured by dark brown (Verona Brown) spores. Primordial sheath remnants when present evident as (usually) scattered, short, fibrillose patches on stipe apex and/or (occasionally) fibrils overlying lamellae. Stipe central to slightly eccentric, terete or compressed, fleshy, aerial length (when pileus fully expanded) generally 30-50 mm, combined length including pseudorhiza up to 165 mm; apex 4-9 (12) mm diam, either widening slightly to ground level or tapering evenly to pseudorhizal origin; surface smooth and shiny beneath relatively dense fibrillose patches; stuffed, with 1.5-2 mm thick cartilaginous cortex surrounding firm (packed when young) to spongy (in age) pallid context, context easily compressed when water-soaked and in age often insect-infested; color generally with drab to smoky brown tones at apex (Deep Quaker /Cinnamon Drab, Avellaneous, Benzo /Army /Natal /Pecan /Wood Brown), increasingly darkening in hue towards ground level (Dusky Drab, Fawn Color, Army /Liver /Natal /Rood's /Sayal Brown, Tawny Olive). Pseudorhiza vertical monopodial, fleshy-cartilaginous, with dark red-brown (Cameo /Hay's /Liver /Mikado /Van Dyke /Verona Brown) upper portion arising from narrow, curled/nippled, brownish drab origin. Spore print color cinnamon-brown ({7.5YR 6-7/4] = (Light) Pinkish Cinnamon). Basidiospores 7x 4.5 +0.0 x 0.3 um (overall range 7-7.5 x 4.4.5 ym), ovate with small apiculus projecting in face view, strongly limoniform with distinct small apiculus and low to moderate beak in profile; verruculose to verrucose, suprahilar disc an area of lowered ornamentation visible rarely and with difficulty in oil immersion; pale to medium pale amber in KOH, pale straw in H,O. 105 Cheilocystidia abundant and crowded, emanating from the lamellar trama to form a dense gelatinous layer, variable in length (15-45 pm), narrowly to broadly clavate, apices usually 5-7 wm diam above long, 3-4 ym diam pedicels, these elements often intermixed with narrower (2-3 ym), cylindrical and/or wider (up to 10 ym), clavate elements, thin-walled; hyaline or (in age) filled with oily brownish contents. Paracystidia occasional within basidial clusters, filiform, 2 wm wide. Basidia (2-) 4-spored, 26-36 x 6-7.5 mm, clavate, hyaline (brownish in older specimens). Pileipellis a bilaminate ixocutis; the suprapellis extensive (150-350 ym thick) and colorless, hyphae long, branched, narrow (1-3 ym), straight or occasionally curling, hyaline, and gelatinized with the uppermost hyphae uplifted from a radially aligned base within a gelatinous matrix; the subpellis brilliant amber-orange, hyphae thin-walled, inflated (to 5-8- 10 wm from 3-5 pm diam septa), with intraparietal pigments most noticeable at the septa. All tramal tissues gelatinized, colorless; in the pseudorhiza mildly sarcodimitic, with wider, thick-walled, cylindrical, vessel hyphae intermixed with less conspicuous, thin-walled, branched, flexuous hyphae; from stipe to pileus dual hyphal system progressively less noticeable, with flexuous hyphae increasing in number as vessel hyphae become more narrow, thin-walled and distorted; in the /amellae subparallel, the central region of 3-5 jm wide hyphae flanked by a compact subhymenium. Tibiiform diverticula abundant on remnants of the pellicular sheath on pseudorhizal and stipitipellis, 5-12 (20) x 2 x 0.5 ym with/without 1-1.5 jm wide capitellum, usually surrounded by an apical secretory droplet; hyaline. Clamp connections absent in all tissues. Macrochemical reactions: Syringaldazine negative (in LLN/I92/031-02 lamellae slightly pink). KOH turning dull or slightly brown. FeSO4 dull green- brown. Fluorescence: Lamellae intense yellow-white when young, intensity dulling as covered by spores; Context dull orange (stipe). RFLP profile: ITS region of 4 isolates including type = 730 bps. Unique among. tested Phaeocollybias in Hinfl (400-190-140); Pall (475-170-85) and Sall (535-190) also diagnostic. Also cut in EcoRI! and Nde2; uncut in Pvu2, Rsal, and Xhol. Ecology, range, distribution, phenology: solitary to scattered (rarely gregarious) in old-growth coniferous montane (650-1350 m elev) forests (Tsuga heterophylla with Abies spp. Picea sitchensis, Pseudotsuga menziesii and Sequoia sempervirens (D. Don) Endl. also present). Uncommon to rare, known from 14 collections made in coastal and inland Washington, Oregon and northern California. Mid/late autumn (October-December). SPECIMENS EXAMINED: UNITED STATES. Catirornia. Mendocino Co. JACKSON SF NEAR MENDOCINO (123.792°W, 39.308°N) — solitary in humus in mixed woods, 29.XII.1960, HDThiers8634 pp (SFSU, as P. fallax); 14.X1.1967 HDThiers21536 (SFSU, as P. fallax). OREGON. Clackamas Co. ESTACADA , HORNER Rp (122.285°W, 45.225°N) — 17.X.1992 JRoger92.10.17-1 (WTU). Linn Co. S McCuLty BLM UNitA 9S 2E 31 — under Tsuga heterophylla and Pseudotsuga menziesii, 15.X1.1999, RLExeter199-M by MLowery (PNW). Multnomah Co. LARCH MTN sUMMIT, Mt HoopNF (122.089°W, 45.531°N) — in late successional (120 yo) Abies procera, Tsuga heterophylla (no understory), LLN1921104-02,,,19¢ -O3:u9 -O5 ing With GTNorvell, SARedhead, PWTrimm (PNW, WTU), LLN1931024.- 05 with 106 FicurE 4. Phaeocollybia rifflipes. LEFT. Holotype (LLN1931020-01, WTU) in situ, Barlow Pass, Snohomish County, Washington. Note appressed fibrillose patches on stipe apex. Scale = 1 cm. RicHt. Young basidiome from Olympic National Park, Washington. LLN1921023-08 (WTU). FiGuRE 5. Basidiospores produced by western North American representatives of the Phaeocollybia festiva complex. Scale = 10 microns. Tor. Phaeocoilybia lilacifolia(Holotype_ -- AHSmith39976, MICH 11625). Photo at right reveals generically characteristic tibiiform diverticula found on vestiges of pellicular sheath on apical stipitipellis. Borrom. Comparison between larger, darker basidiospores from Phaeocollybia fallax (left, LLN1951018-23, WTU) and smaller, paler basidiospores from Phaeocollybia rifflipes (right, Holotype -- LLN1931020- 01, WTU). 107 GTNorvell (PNW); LARCH MTN RoabD, 3 MI BELOW SUMMIT (122.094°W, 45.540°N) — in 2nd growth, dense mid-successional Tsuga, Picea, Abies, LLN1921111-02,-05,-06 with OLNorvell, PWTrimm (PNW, WTU). WasHINGTON. Clallam Co. OLympic NP. RUGGED RIDGE TRAIL (124.172°W, 47.932°N) - old growth Tsuga, Abies, Pseudotsuga, LLNI921023-08 (WTU); Sot Duc, > UPPER VALLEY (123.8632°W,47.9623°N) - in ancient Picea, Tsuga, Abies, LLN1921031-02 by GLBarron (DAOM). Snohomish Co. Mr BAKER-SNOQUALMIE NF, BARLOW Pass (121.437°W, 48.034°N) — in ancient (>450 yo) Abies, Tsuga, Pseudotsuga forest, LLN1931020-01,,,; with GLBarron, SARedhead (WTU — HOLOTYPE; DAOM — isotype). COMMENTS — Phaeocollybia rifflipes, which shares morphological similarities with P. fallax AHSmith and P lilacifolia AHSmith, was first recognized as a distinct species during RFLP analyses of 160 Phaeocollybia collections. The ITS region amplified from 17 specimens provisionally identified as P. fallax or P. lilacifolia produced five different enzyme profiles. One “fallax” isolate from Washington produced such different banding patterns that its ITS region (then coded fal3, later recoded to rif3) was originally thought to have been an amplified contaminant. When its banding patterns were later compared to all other Phaeocollybia isolates, however, three 'unknowns' from Oregon (unk8, unk 9, & unk10, later recoded to riu8, riu9, & riulO respectively) were found to possess the identical profile. Subsequent comparison of morphological characters revealed that all four collections represented the same undescribed species. Phaeocollybia rifflipes, named here for the RFLP-facilitated discovery of acryptic species, differs from P. lilacifolia in its smaller size, more brightly colored (tawny instead of dark black brown) young pileus, narrower, duller lamellae, lack of disagreeable odor, smaller basidiospores, filiform paracystidia, and brilliant orange subpellis. Phaeocollybia fallax is differentiated by an olive green young pileus, intense violet lamellae, frequently hollow stipes, larger basidiospores, and gel-secreting, subcapitate cheilocystidia. The Phaeocollybia festiva “complex” Phaeocollybia rifflipes is a member of a difficult complex of agarics centered around P. festiva, originally described from Sweden (Fries 1838). Species in this complex — P. festiva and the North American endemics P. fallax, P. lilacifolia, and P. rifflipes -- produce small to moderately sized, relatively fragile, relatively odorless/tasteless basidiomes with conic-campanulate glutinous pilei, medium to large (7-9 (10) x 4-5 wm), roughened, limoniform basidiospores, thin-walled +/- clavate cheilocystidia emanating from the lamellar trama, and highly gelatinized tissues lacking clamp connections (NB: Smith's listing of clamp connections for P. fallax in his 1975 field guide was a lapsus calamis). Phaeocollybia festiva (with P. christinae and P. lugubris (Fries) Heim) is one of the three most frequently discussed Phaeocollybias in the literature. Fries, who in his Epicrisis (1838) first treated the species within the tribe Naucoria in Agaricus, described P. festiva as having a green-brown glutinous pileus that dried to a dirty gold (isabelline), pale olive colored lamellae, an apically compressed (neither 108 equal nor ventricose) stipe often covered with dark fibrils, and a raphanoid odor. Later authors expanded the original description to accommodate observed variability. Thus Kiihner & Romagnesi (1953) noted that while the pileus was generally olive to olive brown, it could also be chestnut or red-brown (‘aussi bai ou roux’). Moser (1983, 1995 pers. com.; cf. also Moser & Jiilich 1994), Gulden (1983), and Jacobsson & Stridvall (1983) all remark on the violet lamellae found in occasional specimens, and Gulden (1983, 1992) mentions the drab to violet gray coloration found on some stipe apices (otherwise characterized as being olive-colored). A wide range of spore lengths (from 7-10 ym) is listed for the species, and examinations of Gulden771/81 (O) from Bredehaugen, Norway show that this range can be found within a single specimen. Descriptions of the spore ornamentation range from subdued and marbled (Kiihner & Romagnesi 1953; Reid 1972) through minutely warty (Horak 1977) to strongly or heavily ornamented (Laber 1982; Gulden 1983; Jacobsson & Stridvall 1983). Examination of Gulden771/81 reveals marbled spores with very low indistinct warts stopping well below the apical beak. Also observed in the same collection is a suprahilar disc of lowered ornamentation (reported as “plages” for the species by Bon 1992 and Watling & Gregory 1993, which were noted as lacking by Horak 1977). The shape is variously described as being ovoid (Horak 1977) to limoniform. Field identification of species within the complex is often complicated by mutable pigments found in the pilei and lamellae. Within a single species, pileus colors can be green, brown, orange or some shade in-between, while the base color of the young lamellae -- yellowish buff, tan, pink, drab, violet -- is soon obscured by brown spores. Also within a given species, stipes may vary from firmly stuffed, to stuffed with fibrillose pith, to hollow. Differentiation of species within the complex using a rather undistinguished assemblage of microscopical characters is problematical as well. In some species (notably P. festiva), basidiospore size can be highly variable, while in others the size range appears to be more restricted. Cheilocystidial morphology can be filamentous, cylindrical, narrowly or broadly clavate, or subcapitate subglobose pedicellate. The cheilocystidia are generally quite abundant and crowded, and often there is a mix of all forms within a single specimen. Comparisons of North American material earlier identified as P. festiva as well as recent collections made during the current study to the Norwegian Gulden771/81 suggest that P. festiva does not occur in western North America. Smith (1937) originally reported Naucoria festiva from the Pacific Northwest but later (Smith 1957b) reassigned his collections to P. fallax. Although Smith et al. (1979) reintroduced P. festiva in their keys to North American gilled mushrooms, there is no published documentation or correctly identified material of that species (at MICH or elsewhere) from North America. It would seem that inclusion of P. festiva in the 1979 key is erroneous and that the species should be considered strictly extralimital. 109 In North America, Phaeocollybias with olive-green pilei and intense bluish violet lamellae are instantly identifiable as P. fallax, originally also characterized by a sharply conic-umbonate glutinous pileus, dull drab to greenish stipe apex above a hollow polished orange stipe, and a moderately small stature and fragile aspect. The violet lamellae, helpful in separating P. fallax in the field from two other green-capped species — P. olivacea AHSmith and P. pseudofestiva AHSmith — rapidly dull and darken as they mature. The olive-green cuticle, which usually darkens with age or exposure, can also be obscured in young specimens by a bright orange-brown to dark brown glutin, thereby causing confusion with the tawny capped P. rifflipes and dark brown capped P. lilacifolia. Character variations in P. fallax noted by this author should be mentioned here. While Smith (1957b) describes P. fallax stipes as “tubular-hollow”, stipes stuffed with relatively firm (i.e. not fibrillose) context have been observed in several specimens otherwise clearly identifiable as P. fallax. Likewise, while most specimens sampled by this author did taste “mild”, numerous older specimens, (including all November 1992 collections from California’s Humboldt Redwoods State Park) were found to have pilei with bitter-tasting cuticles and/or glutin. Additionally, although some “fallax” collections were found to possess the “basidia-like” cheilocystidia originally described and illustrated by Smith, a much greater proportion (including some type specimens) were found to be dominated by distinctly subcapitate cheilocystidia, which Smith considered more characteristic of P. lilacifolia. it would appear that a more reliable diagnostic character is the fact that in KOH, “fallax” cheilocystidia characteristically exude a clear heavy gel that eventually spreads out into a dense gelatinous matrix; evidence of this matrix can also be seen in squash mounts in which the abundant cheilocystidia stick together in a dense agglutinated “barrier”, particularly in older or poorly preserved specimens. With additional molecular sampling, some of the observed variability, such as basidiospore length differences, may be shown to be associated with other cryptospecies hiding underthe name of P. fallax. Of the 122 “fallax” collections examined in the current study, most were characterized by basidiospores measuring ~9 x 5 wm; 26 collections, however, contained specimens with basidiospores measuring ~8 x 5 ym. Because these differing spore lengths are not tied to other morphological, anatomical, or molecular differences, the small-spored collections are retained in P. fallax. More problematical are collections from Oregon’s Wildcat Mountain near Mt. Hood (in particular LLN/941]028-04*, -05*,-08*,-17* [WTU, PNW] and LZENI95/0/8-/2*,-23* |WTU, PNW]). In the field, these were all clearly identifiable as P. fallax, despite having less intensely violet lamellae and lacking striations and incurved edges on the pileus. In view of the presence of secretory cheilocystidia and green pigments and the absence of supporting molecular data, these collections have been retained in P. fallax, despite sharing a similar basidiospore morphology with P. rifflipes. Phaeocollyba lilacifolia, like P. rifflipes, lacks green pigments. This largest representative of the P. festiva complex is distinguished from P. rifflipes by its 110 more robust stature, darker pileus, larger basidiospores, and narrower cheilocystidia and from P. fallax by its dark brown slightly broader pileus, larger size, narrower cheilocystidia, and slightly disagreeable odor. Although Horak (1973) initially illustrated clear differences between basidiospore size, ornamentation, and extent of “plage” in his drawings of P. fallax and P. lilacifolia, he eventually placed the latter species in synonymy with P. fallax (Horak 1977). At that time Horak did not discuss either basidiospore or pigmentation differences, although he later addressed the mutability of both green and lilac pigments in Pyrrhoglossum and Phaeocollybia, genera which he considered to be closely allied (Horak 1989). As noted above, recent collections of numerous “fallax” basidiomes with both green and brown colors on their pilei underscore the difficulty of relying solely on pileus pigmentation for species differentiation in this difficult group. At this date, molecular data analyses are still too fragmentary and inconclusive to test the two species hypotheses. However, given the more robust stature of P. lilacifolia noted by Smith (1957ab), the different average spore lengths observed in the types [8 wm and 9 yum respectively for AHSmith39976, MICH 11625* (P. lilacifolia), and AHSmith79252*, MICH (P. fallax)|, and pigmentation differences [also cited by Bandala (1994) as a reason for rejecting Horak’s synonymy], P. fallax and P. lilacifolia should be retained independent taxa, at least for the time being. A final nomenclatural note: It should be noted that in his monograph, Smith (1957b) discussed P. lilacifolia, a species he had previously described in another paper published the same year (Smith 1957a), before P. fallax. From this, an argument can be made that should the two species be considered synonymous, the name P. lilacifolia has priority. Acknowledgments I thank Joe Ammirati for guidance, financial support, and laboratory facilities during my doctoral studies at the University of Washington. Scott Redhead provided general support, field assistance, and numerous insightful discussions. Oregon BLM Botanist Ron Exeter supplied the holotype of P. phaeogaleroides and many well-annotated Phaeocollybias. Todd Norvell provided encouragement and field assistance. George Barron, Judy Roger, and Nancy Smith Weber generously donated collections, and specimens were kindly loaned to me by Efren Cazares, Dennis Desjardin and Harry D Thiers (SFSU), Robert Fogel (MICH), Gro Gulden (O), and Renata Outerbridge. Luca Comai, Amy Denton, David Hibbett, Francois Lutzoni, Yajuan Liu, George Mueller, Karen Nakasone, Michelle Seidl, John Stiller, and Gene Takimoto provided guidance or supplies during molecular work and data analyses. Agriculture & Agri-Food Canada (via Scott Redhead and Elizabeth Fox) granted permission to publish photographs of P. phaeogaleroides (Fig la) and P. rifflipes (Figs 3a, 3b). Field support provided by the Olympic National Resource Center (Forks, WA) and financial assistance provided by the 111 Daniel E Stuntz Memorial Foundation, Oregon-Washington Reciprocity Award, and the Mycological Society of America (Graduate Fellowship) are gratefully acknowledged. Thanks go to Scott Redhead and Ron Exeter for presubmission reviews. Literature Cited Bandala VM. 1994. Contribucién al estudio monografico del género Phaeocollybia (Fungi, Basidiomycotina, Agaricales) en México. MSc thesis. Facultad de Ciencias, UNAM, Mexico City, Mexico. Bon M. 1992. Clé monographique des espéces Galero-naucorioides. Genre Phaeocollybia Heim. Documents mycologiques 21(84): 44-47. Fries EM. 1836-38. Epicrisis Systematis Mycologici, Synopsis Hymenomycetum. Uppsala: Typographia Academica. Gulden G. 1983. Slekten topphatt -- Phaeocollybia | Norge. Bleckksoppen 30: 3-17. Gulden G. 1992. Phaeocollybia. In Nordic Macromycetes, Vol. 2: Polyporales, Boletales, Agaricales, Russulales (eds. L. Hansen & H. Knudsen). pp. 133-134. Copenhagen: Nordsvamp. Holmgren PK, Holmgren NH, Barnett LC. 1990. Index herbariorum: Part I: Herbaria of the World, 86th ed. Bronx: New York Botanical Garden. Horak E. 1973. Fungi Agaricini Novazelandiae I-V: IV, Phaeocollybia Heim (1931). Beih. Nova Hedwigia. 43: 183-191. Horak E. 1977. Further additions towards a monograph of Phaeocollybia. Sydowia 29: 28-70. Horak E. 1989. New and additional data concerning Pyrrhoglossum and eccentric or stipitate taxa of Gymnopilus (Agaricales). Op. Bot. 100: 115-129. Jacobsson §S, Stridvall L. 1983. Slaktet Phaeocollybia in Sverige. Windahlia 12-13: 57-79. Kiihner R, Romagnesi H. 1953. Flore analytique des champignons supérieurs. Paris: Masson et Cie. Laber D. 1982. Die europaischen Arten der Gattung Phaeocollybia (Wurzelschnitzlinge) und ihr Vorkommen im siidlichen Schwarzwald. Zeitschrift fiir Mykologie. 48(1): 89-98. Moser M. 1983. Keys to Agarics and Boleti (Polyporales, Boletales, Agaricales, Russulales). Tonbridge: Roger Phillips & Whitefriars Press. Moser M, Jiilich W. 1994. Farbatlas der Basidiomyceten. Stuttgart: Gustav Fischer. [Munsell] Munsell Color. 1976. Munsell Book of Color. Baltimore: MacBeth. Norvell LL. 1998a. The Biology and Taxonomy of Pacific Northwest Species of Phaeocollybia (Agaricales, Cortinariaceae). PhD Dissertation, University of Washington, Seattle. 391 pp. Norvell LL. 1998b. Observations on the development, morphology and biology of Phaeocollybia. Mycological Research 102(5): 615-630. Norvell LL. 1998c. ROD: Strategy 3 Fungal species Evaluation (11 Gilled Basidiomycete Strategy 3 Species). Unpublished report on file at USFS -- Forest Sci Lab, Corvallis. 1480 pp. Norvell LL. 2000. Phaeocollybia in Western North America 1: The Phaeocollybia kauffmanii complex. Canadian Journal of Botany 78: 1055-1076. Norvell LL, Redhead, SA. 2000. Phaeocollybia in Western North America 2: The vernal P. pleurocystidiata sp. nov. and P. carmanahensis reconsidered. Mycologia 92: 984- Oot Reid DA. 1972. Fungorum Rariorum Icones Coloratae VI. pp. 1-64. Pl. 41-48. Lehre: J. Cramer. Ridgway R. 1912. Color standards and color nomenclature. Washington, D. C.: Published by the author. . Smith AH. 1937. New and unusual Agarics from the western United States. Mycologia 29: 45- 59. Smith AH. 1957a. Additional new or unusual North American agarics. Beih. Sydowia. |: 40-61. Smith AH. 1957b. A contribution toward a monograph of Phaeocollybia. Brittonia 9: 195-217. Smith AH. 1975. A Field Guide to Western Mushrooms. Univ. Mich Press: Ann Arbor. Smith AH, SmithHV, Weber NS. 1979. How to Know the Gilled Mushrooms. Dubuque: Wm. C. Brown. ty [USDA-USDI] USDA-Forest Service, USDI-Bureau of Land Management. April 1994. Record of decision [ROD] for amendments to Forest Service and Bureau of Land Management planning documents within the range of the Northern Spotted Owl & Standards and guidelines for late-successional and old-growth forest related species within the range of the Northern Spotted Owl. US Govt Printing Office. d Boleti. 7 / Cortinariaceae p.p. Watling R, Gregory N. 1993. British fungus flora: Agarics an Phaeocollybia, — Phaeogalera, [Galerina, | Gymnopilus, _ Leucocortinarius, Phaeolepiota, Phaeomarasmius, Pleuroflammula, Rocites & Stagnicola]. Edinburgh: Royal Botanic Garden. MY COTAXON Volume LXXXI, pp. 113-176 January-March 2002 PRELIMINARY CHECKLIST OF AGARICALES FROM CROATIA. I. FAMILIES PLEUROTACEAE AND TRICHOLOMATACEAE. ZDENKO TKALCEC & ARMIN MESIC Laboratory of Biocoenotics, Department of Molecular Genetics Ruder BoSkovié Institute Bijeni¢ka cesta 54, 10000 Zagreb, Croatia E-mail: zdenko.tkalcec@zg.tel.hr E-mail: armin.mesic@zg.hinet.hr ABSTRACT The first part of the preliminary checklist of Agaricales from Croatia is given. It comprises the families Pleurotaceae and Tricholomataceae. 252 species (three of them in a broad sense) belonging to 48 genera are presented. Three genera and 35 species are reported for the first time from Croatia. For each species, the published and unpublished sources of data are given, as well as the collections in which the material is deposited. Key words: Basidiomycotina, biodiversity, mycoflora, bibliography. INTRODUCTION The fungi of Croatia are poorly known. Some parts of Croatia have never been mycologically explored. The Agaricales are no exception as few mycologists have researched the order. The first data on this taxonomic group were published by Stjepan Schulzer (1857). He was the only Croatian myco- logist in the second half of the 19 century. It was not until the second half of the 20 century that more intensive research was carried out. Significant con- tributions to the research on the Agaricales during that period were made by Milica Torti¢é, Ivan Focht, Neven Matotec and the authors of this paper. Although it has been a century and a half since the beginning of mycological research in Croatia, no survey of species of Agaricales has been made until now. As the first step to a recently started comprehensive research of this 114 taxonomic group in Croatia, we are preparing a checklist of species of the order Agaricales from Croatia including all published and most unpublished sources of data known to us. This checklist should be considered only pre- liminary since only some data have been revised and that comprehensive research of this taxonomic group in Croatia has just begun. The checklist will be divided into five articles of which this is the first. The proposed articles will treat the families in the following order: I. - Pleurotaceae, Tricholomata- ceae; II. - Agaricaceae, Amanitaceae, Cortinariaceae, Hygrophoraceae; III. - Boletaceae, Gomphidiaceae, Paxillaceae; IV. - Bolbitiaceae, Coprinaceae, Entolomataceae, Pluteaceae; V. - Crepidotaceae, Russulaceae, Strophariaceae. Geographically, Croatia connects southern and central Europe. It is situated between 42°22' and 46°33' northern latitude, and between 13°29' and 19°27' eastern longitude. The land area of Croatia is 56,610 km’. The altitude varies from 0 to 1,831 m. Croatia consists of three main geographic entities: the Pannonian lowland zone, the Karst mountainous zone of the Dinarides and the Mediterranean zone of the coast and islands of the Adriatic Sea. The main types of climate are moderate continental, moun- tainous and mediterranean. The average annual temperature ranges from 3.0 to 16.5 °C. The average annual rainfall ranges from 260 to 3,700 mm. The area of Croatia covered by forest is 37% of its land area. In the Pannonian lowland zone the dominant trees are Quercus robur, Q. petraea and Carpinus betulus. In the Karst mountainous zone the dominant trees are Fagus sylvatica, Abies alba and Picea abies. In the Mediterranean zone the dominant trees are Quercus ilex, QO. pubescens, Carpinus orientalis, Ostrya carpinifolia, Pinus halepensis and P. nigra. MATERIAL, METHODS AND PRESENTATION This checklist includes, except for a few cases, all the published records of species, subspecies and varieties of the order Agaricales (sensu lato) from Croatia, known to us. Records from daily and weekly newspapers, records obviously erroneously identified, and records from Hafner (1997) due to false data, have been excluded. This checklist also includes, except for a few cases, all the unpublished records known and available to us. Two manuscripts by Stjepan Schulzer have not been included (one is kept at the Library of the Hungarian Academy of Sciences in Budapest, and the other at the National and University Library in Zagreb), as well as notes by Ivan Focht, because it is not possible to interpret their data in a satisfactory way without thorough revision. We have also excluded the unpublished records whose identification the original author doubted. The checklist includes data published or collected until 1 January 2000. The material known to us is deposited in the following herbaria and fungaria. 115 BEO - Herbarium, Botany Department, Natural History Museum of Serbia, Beograd, Yugoslavia. BP - Herbarium, Botanical Department, Hungarian Natural History Museum, Budapest, Hungary. BRA - Herbarium, Botany Department, Natural History Museum, Slovak National Museum, Bratislava, Slovakia. BRNM - Herbarium, Department of Botany, Moravian Museum, Brno, Czech Republic. Candusso - Private fungarium of Massimo Candusso, Alassio, Italy. CNF - Croatian National Fungarium, Croatian Mycological Society, Zagreb, Croatia. H - Herbarium, Botanical Museum, University of Helsinki, Helsinki, Finland. Hilber - Private fungarium of Oswald Hilber, Tegernheim b. Regens- burg, Germany. L - Rijksherbarium, Leiden, the Netherlands. Mrvo8 - Private fungarium of DuSan Mrvo8, Zagreb, Croatia. POZM - Herbarium, Department of Plant Ecology and Environment Protection, Adam Mickiewicz University, Poznan, Poland. PRM - Herbarium, Mycological Department, National Museum in Prague, Praha, Czech Republic. Along with each presented species we specify the published and unpublished sources of records for Croatia, and the collections in which the collected material is deposited. In this way we give a more or less complete survey of the published sources of data, as well as a good survey of the unpublished sources of data on the species of the order Agaricales in Croatia. We also specify synonyms and misapplied names which the authors of records of taxa have used, as well as those used in modern taxonomic literature. We have tried to interpret all records on the basis of taxa names and dates of recording in the sense of modern taxonomy. The older the record, the greater the need for revision. So far we have revised only a part of the revisable records. We have revised all our entries in this checklist, as well as a smaller number of records by other authors indicated as such in the text. Species and varieties revised by us, as well as species and varieties recorded for Croatia by authors of taxonomic literature which we have used for the interpretation of records are marked with a black dot. At the end of the data specified for such species we cite the literature (in brackets) on the basis of which we have revised or interpreted the records. The name of each genus is followed by the literature (given in brackets) used for the interpretation, nomenclature and synonyms of records in the genus. At the end of genus we have included those species for which only a wide locality is quoted, but on which basis one may conclude that the species was most probably found in Croatia. Schulzer (1867) corrects numerous mistakes from the paper by Schulzer & 116 al. (1866), so we give such quotes as "Schulzer & al. (1866) + Schulzer (1867)". We have cited abbreviations of authors of taxa according to Kirk & Ansell (1992) and abbreviations of journals in chapter "Literature cited" according to Bridson & Smith (1991). The chapter "Excluded records" comprises: (a) records of species and infraspecific taxa we were unable to find in modern taxonomic literature, (b) records we could not interpret in the sense of modern classification, (c) records which are a nomen nudum, and (d) records which have been incorrectly recorded for Croatia or have been recorded for the territory which does not now belong to Croatia. Genus, subgenus or section listed by the original author is specified in square brackets. When we could not check whether the author of the record had given the correct author citation, we have given it herein according to that author. In this paper we present 252 species (three of them in a broad sense) belonging to 48 genera found on the territory of Croatia, of which we have revised 125 species. Out of the overall number 13 species belong to the family Pleurotaceae and 239 to the family Tricholomataceae. Three genera and 35 species are reported for the first time from Croatia. Marks and abbreviations: * - taxon reported for the first time from Croatia in this paper; ¢ -at least part of records of taxon revised; (1) - number in brackets indicates synonym or misapplied name under which the author of the record has quoted taxon; ( ) - empty brackets indicate that the author of the record has also quoted the taxon under the name we use; - (a) question mark instead of an author of combination means that we could not find the author in the literature available to us; (b) question mark following the source of a published record or abbreviation of a collection means that the author himself doubted his original identification; =? - taxon not found in waratite taxonomic literature; Exs. - collections in which the dried material (exsiccatum) is deposited; HR? - taxon most probably found in the territory of Croatia (locality not precisely quoted); Incl. - including; Lit. - literature; Misappl. - misapplied name; Ref. - references (sources of records of taxon); rev. - revised (when we have revised records of other authors); unp. - unpublished data. 117 SURVEY OF TAXA Familia Pleurotaceae Overeem ex Kiihner et Romagn. Lentinus Fr. : Fr. = Panus Fr. (Lit.: Arnolds & al. 1995, Bas & al. 1990, Bollmann & al. 1996, Dennis & al. 1960, Hansen & Knudsen 1992, Kotlaba & Pouzar 1996, Kreisel 1987, Pegler 1983, Pilat 1935, Pilat 1946, Singer 1986, Watling & Gregory 1989) Lentinus adhaerens (Alb. et Schwein. : Fr.) Fr. = Lentinus pulverulentus (Scop.) Fr. (1) Ref.: Skorié 1928 (1), Tortié 1973 ( )(1), Tortié 1989 b, Tortié 1993, unp. Torti¢ Exs.: CNF °Lentinus conchatus (Bull. : Fr.) J. Schrot. = Panus conchatus (Bull. : Fr.) Fr. (1) = Lentinus bresadolae Schulzer (nom. illeg.) (2) = Lentinus divisus Schulzer (3) = Agaricus pulvinatus Pers. : Fr. (4) = Lentinus torulosus (Pers. : Fr.) Lloyd = Panus torulosus (Pers; Fr.) Fr. (5) Ref.: Saccardo 1887 (2), Schulzer 1879 (3), Schulzer 1885 (2), Schulzer & al. 1866 (5), Torti¢é 1964 (1), Torti¢é 1966 a (1), Tortié 1966 b (1), Torti¢é 1993 (1), Vouk & Pevalek 1916 (4), unp. Matoéec (1), unp. Tkaléec & MeSi¢, unp. Tortié (1) EXsn CONE (Lit.: Hansen & Knudsen 1992, Moser 1983, Watling & Gregory 1989) °Lentinus degener Kalchbr. = Lentinus leontopodius Schulzer (1) Misappl.: Lentinus cyathiformis (Schaeff.) Bres. sensu auct. non Schaeff. (2) (sensu Schaeff. = Pleurotus pulmonarius (Fr. : Fr.) Quél. - Kotlaba & Pouzar 1996) Ref.: Kalchbrenner & Schulzer 1873-1877 (1), Pilat 1946 (2), Tortié 1968 b (2), unp. Tkaléec & MeSi¢, unp. Torti¢ (2), Fries 1874 HR? (1), Saccardo 1887 HR? (1) Exss GNA (Lit.: Bas & al. 1990, Moser 1983, Pegler 1983) Lentinus lepideus (Fr. : Fr.) Fr. = Lentinus queletii Schulzer (1) = Lentinus squamosus (Schaeff. —) Quél. (2) 118 Ref.: Focht 1974, Schulzer 1885 (1), Tortié 1989 b, Vouk & Pevalek 1916 (2), unp. Tortic, Saccardo 1887 HR? (1) EXs GNE | °Lentinus strigosus (Schwein. : Fr.) Fr. = Lentinus lecomtei Fr. : Fr. = Panus lecomtei (Fr. : Fr.) Corner (1) = Lentinus rudis (Fr.) Henn: (2) = Panus rudis Fr. (3) = Pleurotus rudis (Fr.) Pilat (4) = Panus sainsonii (Lév.) Heufl. (5) Ref.: Moesz 1938 (3), Picbauer 1928 (3), Pilat 1925 (3), Pilat 1935 (4), Schulzer 1882 a (5), Torti¢ 1962 (3), Tortié 1964 (3), Torti¢ 1966 a (3), Torti€é 1966 b (3), Tortié 1989 b (3), Tortié 1993 (1), Vouk & Pevalek 1915 (2), unp. Listewska (3), unp. Matoéec (1), unp. Tkaléec & MeSié, unp. Torti¢ (3) Exs.: CNF, POZM (3) (Lit.: Hansen & Knudsen 1992, Moser 1983) ¢Lentinus tigrinus (Bull. : Fr.) Fr. = Panus tigrinus (Bull. : Fr.) Singer (1) Ref.: Bozac 1982 (1), Jelié & Torti¢é 1973 (1), Schulzer 1882 a, Schulzer & al. 1866, Skorié 1938, Tortié 1964 (1), Tortié 1989 b (1), unp. Matoéec (1), unp. Tkaléec & Me8ié, unp. Torti¢ ( )(1) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983, Watling & Gregory 1989) Phyllotopsis (J.-E. Gilbert et Donk) ex Singer (Lit.: Bas & al. 1990, Hansen & Knudsen 1992, Kreisel 1987, Pilat 1935, Singer 1986, Watling & Gregory 1989) Phyllotopsis nidulans (Pers. : Fr.) Singer = Agaricus nidulans Pers. : Fr. (1) = Pleurotus nidulans (Pers. : Fr.) P. Kumm. (2) Ref.: Focht 1974, Schulzer & al. 1866 (1), Torti¢ 1968 b, unp. Torti¢ ( )(2), Schulzer 1857 HR? (1) Exs.:.CNF Pleurotus (Fr.) P. Kumm. (Lit.: Arnolds & al. 1995, Bas & al. 1990, Bollmann & al. 1996, Hansen & Knudsen 1992, Hilber 1982, Konrad & Maublanc 1924- -1937, Kreisel 1987, Krisai-Greilhuber & Voglmayr 1995, Pilat 1935, Singer 1986, Watling & Gregory 1989) ®Pleurotus calyptratus (Lindblad) Sacc. Ref.: Hilber 1982, Jeli¢é & Torti¢é 1973, unp. Tkaléec & MeSié, unp. Tortic 119 Exs.: CNF, Hilber (Lit.: Hilber 1982) °Pleurotus cornucopiae (Paulet ex Pers.) Rolland = Agaricus sapidus Schulzer (1) = Panus sapidus Schulzer (nom. nud.) (2) = Pleurotus sapidus (Schulzer) Sacc. (3) Ref.: Blagai¢é 1931 (3), Hilber 1982, Kalchbrenner & Schulzer 1873- -1877 (1), Schulzer 1881 (1), Schulzer 1882 a (1), Schulzer & al. 1866 (2), unp. Tkaléec & MeSi¢, unp. Torti¢, Fries 1874 HR? (1), Saccardo 1887 HR? (3) Exs.: CNF, Hilber (Lit.: Hilber 1982) °Pleurotus dryinus (Pers. : Fr.) P. Kumm. = Pleurotus corticatus (Fr. : Fr.) P. Kumm. (1) Ref.: Focht 1974, Hilber 1982, KiSpati¢ 1950 (1), unp. Tkaléec & MeSié, unp. Tortic Exs.: CNF, Hilber (Lit.: Hilber 1982) °Pleurotus eryngii (DC. : Fr.) Quél. °var. nebrodensis (Inzenga) Sacc. Ref.: Hilber 1982, unp. Tkaléec & MeSic¢, unp. Torti¢ Exs.: CNF, Hilber (Lit.: Hilber 1982) °Pleurotus ostreatus (Jacq. : Fr.) P. Kumm. = Agaricus ostreatus Jacq. : Fr. (1) = Panus ostreatus (Jacq. : Fr.) ? (2) = Agaricus salignus Pers. : Fr. (3) = Panus salignus (Pers. : Fr.) ? (4) = Pleurotus salignus (Pers. : Fr.) P. Kumm. Ref.: Blagaié 1921 b, Blagai¢ 1931, Focht 1974, Focht 1979, Focht 1981 a, Focht 1983 a, Focht 1986, Kranjéev 1986 b, Lorinser - Janda 1877, Moesz 1938, Schulzer & al. 1866 (2)(4), Skorié 1928 (1), Tortié 1962, Tortié 1964, Torti¢ 1966 a, Torti¢ 1966 b, Tortié 1993, Torti¢é 1994, Urbani 1946, Vouk & Pevalek 1915 (1)(3), Vouk & Pevalek 1916 (1)(3), unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Torti¢é, Schulzer 1857 HR? (1)(3) Exs.: CNF (Lit.: Hilber 1982) Pleurotus ostreatus (Jacq. : Fr.) P. Kumm. sensu lato Ref.: Tortié 1989 b °Pleurotus pulmonarius (Fr. : Fr.) Quél. = Pleurotus ostreatus (Jacq. : Fr.) P. Kumm. var. pulmonarius (Fr. : Fr.) Konrad et Maubl. (1) 120 Ref.: Baréié 1982 (1), Baréié 1996 (1), Focht 1983 a, Lisiewska & Tortié 1990, Tortié 1966 a (1), unp. Lisiewska, unp. Tkaléec & Me8ié, unp. Torti¢ (1) Exs.: CNF, POZM (Lit.: Hilber 1982) Familia Tricholomataceae R. Heim ex Pouzar Armillaria (Fr.) Staude = Armillariella (P. Karst.) P. Karst. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Hansen & Knudsen 1992, Kreisel 1987, Marxmiiller 1992, Pilat 1951 b, Termorshuizen & Arnolds 1987, Volk & Burdsall 1995) *° Armillaria gallica Marxmiller et Romagn. Misappl.: Armillaria lutea Gillet sensu auct. non Gillet Ref.: unp. Tkaléec & Mesié BXxs.sCNE (Lit.: Bas & al. 1995, Hansen & Knudsen 1992) ° Armillaria mellea (Vahl : Fr.) P. Kumm. = Agaricus melleus Vahl : Fr. (1) = Armillariella mellea (Vahl : Fr.) P. Karst. (2) = Clitocybe mellea (Vahl : Fr.) Ricken (3) Ref.: Baréié 1982 (2), Barcié 1996 (2), Blagai¢ 1921 a (3), Blagaié 1931 (3), Focht 1974 (2), Focht 1981 b (2), Forenbacher 1915, Georgevitch 1926, GjuraSin 1898, GlavaS & Halambek 1992 (2), Halambek & al. 19937 (2) Jelicy dec) Lortics 1973 52 ae Rispaticw 1 99 Pe ik ranjcey a1 979, Kranjcev 1986 b (2),.Kranjéev 1991 b (2), Kranjéev 1995 (2), Lange 1974, Lorinser - Janda 1877, Muzic & Bozac 1984 (2), Schulzer & al. 1866 (1), Sarié-Sabado’ 1957, Skorié 1926 (1), Skorié 1928, Tortié _ 1964, Torti¢ 1966 a, Torti¢ 1966 b, Tortié 1968 a (2), Tortié 1978 (2), Torti¢ 1994, Vouk & Pevalek 1915, Vouk & Pevalek 1916, unp. Matoéec, unp. Tkaléec & MeSic, unp. Torti¢ 1970 (2), unp. Tortié, Schulzer 1857 HR? (1) _ Exs.: BEO, CNF (Lit.: Bas & al. 1995, Hansen & Knudsen 1992) Armillaria mellea (Vahl : Fr.) P. Kumm. sensu lato = Armillariella mellea (Vahl : Fr.) P. Karst. sensu lato (1) = Agaricus melleus Vahl : Fr. var. fuscopallidus (Bolton) ? (2) Ref.: Lisiewska & Torti¢ 1990, Schulzer 1867 (2), Torti¢é 1989 b (1), Torti¢é 1993 °*Armillaria ostoyae (Romagn.) Herink Misappl.: Armillaria obscura (Schaeff.) Herink sensu auct. non Schaeff. (sensu Schaeff. = nom. dub. - Termorshuizen & Arnolds 1987) 121 Misappl.: Armillariella obscura (Schaeff.) Romagn. sensu auct. non Schaeff. (comb. inval.) (1) Ref.: Focht 1986 (1), unp. Tkaléec & MeSié Exseac NG (it basic) 99)) *Armillaria tabescens (Scop.) Emel = Armillariella tabescens (Scop.) Singer (1) = Clitocybe tabescens (Scop.) Bres. (2) = Agaricus inarmillatus Schulzer (3) Ref.: Barci¢ 1982 (1), Barcic 1996 (1), Blagaié 1921 a (2), Blagaié 1931 (2), Focht 1979 (2), Focht 1981 a (2), Focht 1983 a (2), Focht 1986 (1), Kranjéev 1995 (2), Torti¢ 1964 (2), Tortié & Lisiewska 1978 (1), unp. Mato€ec, unp. Tkaléec & MeSi¢, unp. Torti¢ 1970 (1), unp. Tortié (2), Schulzer 1883 a HR? (3) Exs CNE (Lit.: Bas & al. 1995, Moser 1983) * Arrhenia Fr. = Leptoglossum P. Karst. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Hansen & Knudsen 1992, Singer 1986) *Arrhenia spathulata (Fr. : Fr.) Redhead = Leptoglossum muscigenum (Bull. : Fr.) P. Karst. (1) Ref.: unp. Tortié (1) Exs.) ONE Asterophora Ditmar : Fr. = Nyctalis Fr. (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Moser 1983, Singer 1986) ° Asterophora lycoperdoides (Bull. : Fr.) Ditmar = Nyctalis asterophora Fr. (1) Ref.: Kranjéev 1991 a, Kranjéev 1995, unp. Matoéec (1), unp. Tkaléec & MeSié, unp. Torti¢é, Schulzer 1857 HR? (1) Exs GNE (Lit.: Hansen & Knudsen 1992, Moser 1983) Asterophora parasitica (Bull. : Fr.) Singer = Nyctalis parasitica (Bull. : Fr.) Fr. Ref.: Focht 1974 Baeospora Singer (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Turnbull 1998) 2 *® Baeospora myosura (Fr. : Fr.) Singer Ref.: unp. Tkaléec & Mesic Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) Baeospora myriadophylla (Peck) Singer Ref.: Torti¢é 1973, Tortié & Jeli¢ 1970, unp. Matocec, unp. Torti¢é Exs.: CNF Calocybe Kiihner ex Donk (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1977, Singer 1986) °Calocybe carnea (Bull. : Fr.) Ktthner ex Donk Ref.: Tortié 1973, unp. Tkaléec & MeSi¢, unp. Tortic Exs:: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) *® Calocybe chrysentera (Bull. : Fr.) Singer Ref.: unp. Tkaléec & Mesic Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) *Calocybe gambosa (Fr. : Fr.) Singer ex Donk = Agaricus gambosus Fr. : Fr. (1) = Tricholoma georgii (Clus. ex L.) Quél. var. gambosa (Fr. : Fr.) ? (2) = Agaricus georgii Clus. ex L. (3) = Calocybe georgii (Clus. ex L.) Kitihner (comb. inval.) (4) = Tricholoma georgii (Clus. ex L.) Quél. (5) Ref.: Blagaié 1923 (2)(5), Blagai¢ 1931 (5), Focht 1979 (4), Focht 1983 b (4), Focht 1986 (5), Muzic & Bozac 1984 (5), Muzic & Bozac 1997 ( )(5), Tortié 1966 a (4), Torti¢ 1966 b (4), Torti¢ 1968 b (4), Vouk & Pevalek 1916 (1)(3), unp. Matocéec (4), unp. Tkaléec & MeSic¢, unp. Torti¢ (5), Focht 1981 a HR? (4) Exsi,CNE | (Lit.: Hansen & Knudsen 1992, Moser 1983) ~ *Calocybe ionides (Bull. : Fr.) Kiihner ex Donk = Agaricus ionides Bull. : Fr. (1) Ref.: Schulzer & al. 1866 (1), Torti¢ 1968 b, Tortié & Lisiewska 1978, unp. Matocéec, unp. Tkaléec & Me8ié, unp. Torti¢ Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) *°@ Calocybe obscurissima (A. Pearson) Moser Ref.: unp. Tkaléec & MeSié Exs.: CNF (Lit.: Moser 1983) 123 Catathelasma Lovejoy (Lit.: Bas & al. 1995, Hansen & Knudsen 1992, Kreisel -1987, Moser 1983) Catathelasma imperiale (Fr. > Quél.) Singer Ref.: Focht 1986, unp. Torti¢ Exs.: CNF Chaetocalathus Singer (Lit.: Antonin & Noordeloos 1997, Bollmann & al. 1996, Matzke & Senn-Irlet 1992, Pilat 1935, Singer 1986) Chaetocalathus craterellus (Durieu et Lév.) Singer = Pleurotus craterellus (Durieu et Lév.) Quél. (1) Ref.: Jaap 1916 (1), Pilat 1935 (1), Torti¢é 1993, unp. Kotlaba (according to unp. Tortic) Exs.: CNF, PRM Chrysomphalina Clémencon (Lit.: Bollmann & al. 1996, Clémen¢gon 1982 a, Hansen & Knudsen 1992, Singer 1986) Chrysomphalina chrysophylla (Fr. : Fr.) Clémengon = Gerronema chrysophyllum (Fr. : Fr.) Singer (1) = Omphalina chrysophylla (Fr. : Fr.) Murrill Ref; Tortic 1973 (1); Tortic 1989'b (1), unp, Tortic (1) Exs,: CNF Clitocybe (Fr.) Staude (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Bollmann & al. 1996, Clémengon 1984 a, Hansen & Knudsen 1992, Harmaja 1969, Kreisel 1987, Ktihner & Romagnesi 1953, Kuyper 1996, Moser 1983, Singer 1978, Vouk & Pevalek 1916) *Clitocybe alexandri (Gillet) Gillet Ref.: BoZac 1984, VrSéaj 1990, unp. Tkaléec & Mesié EXst: GNP (Lit.: Bas & al. 1995, Clémengon 1984 a) Clitocybe cacabus (Fr.) Gillet = Agaricus cacabus Fr. (1) Ref.: Skorié 1928 (1), unp. Torti¢é ExsviG NE. Clitocybe catinus (Fr.) Quél. = Clitocybe infundibuliformis Schaeff. var. catinus (Fr.) Maire (1) Ref.: Focht 1972, Vouk & Pevalek 1915 (1) 124 *Clitocybe clavipes (Pers. : Fr.) P. Kumm. Ref.: Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993, GjuraSin 1898, unp. Matoéec, unp. Tkaléec & MeSi¢, Blagai¢ 1921 b? Exs.: CNF (Lit.: Bas & al. 1995, Clémengon 1984 a) * Clitocybe costata Kihner et Romagn. = Clitocybe infundibuliformis (Schaeff.) Quél. var. costata (Kiihner et Romagn.) ? (1) Ref.: unp. Torti¢ ( )(1) Exs. GNE Clitocybe ericetorum (Bull.) Quél. = Agaricus ericetorum Bull. (1) Ref.: GjuraSin 1898, Vouk & Pevalek 1915 (1) Clitocybe fragrans (With. : Fr.) P. Kumm. = Clitocybe suaveolens (Schumach. : Fr.) P. Kumm. (1) Ref.: GjuraSin 1898 (1), Tortié 1968 b (1), unp. Torti¢ (1), Exs.: CNF ; Clitocybe gallinacea (Scop.) Gillet Ref.: Bozac 1982, Gjurasin 1898 *Clitocybe geotropa (DC. et Lam.) Quel. Ref.: Baréié 1982, Barcicé 1984, Baréié 1996, Bozac 1982, Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993, Focht 1972, Focht 1979, Focht 1983 b, Focht 1986, unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Tortic Exs.: CNF (Lit.: Bas & al. 1995, Clémen¢gon 1984 a, Hansen & Knudsen 1992) *Clitocybe gibba (Pers. : Fr.) P. Kumm. Misappl.: Agaricus infundibuliformis Schaeff. sensu auct. non Schaeff. (1) (sensu Schaeff. = Lepista flaccida (Sowerby : Fr.) Pat. - Bas & al. 1995) Misappl.: Clitocybe infundibuliformis (Schaeff.) Quél. sensu auct. non Schaeff. (2) | Ref.: Baréié 1982, Baréié 1996, Focht 1972 (2), Focht 1974( )(2), Focht 1979 (2), GjuraSin 1898 (2), Lisiewska & Torti¢é 1990, Tortic 1964 (2), Torti¢ 1966 a (2), Tortié 1966 b (2), Torti¢ 1993, Torti¢ & Lisiewska 1978, Vouk & Pevalek 1916 (1), unp. Matocec (2), unp. Tkaléec & Me8ic¢, unp. Tortié ( )(2) EXs GE (Lit.: Bas & al. 1995, Clémencon 1984 a) Clitocybe gyrans (Paulet) Sacc. Ref.: Focht 1974 125 * Clitocybe incilis (Fr.) Quél. Ref.: unp. Matocec Clitocybe inornata (Sowerby : Fr.) Gillet Ref.: Focht 1972, unp. Torti¢ Exs.CNE Clitocybe lignatilis (Pers. : Fr.) P. Karst. = Ossicaulis lignatilis (Pers. : Fr.) Redhead et Ginns = Pleurocybella lignatilis (Pers. : Fr.) Singer (1) = Pleurotus lignatilis (Pers. : Fr.) P. Kumm. (2) Ref.: Torti¢é 1968 b (1), Tortié 1989 b, unp. Tortié ( )(2) EXSe NE Clitocybe metachroa (Fr. : Fr.) P. Kumm. = Clitocybe dicolor (Pers.) Murrill (1) Ref; Baréic 1982°(1); Barcic 1996 (1), Focht 1972, Torti¢. 1973, unp. Torti¢ EXSeeC NE *Clitocybe nebularis (Batsch : Fr.) P. Kumm. = Agaricus nebularis Batsch : Fr. (1) = Lepista nebularis (Batsch : Fr.) Harmaja Ref.: Baréié 1982, Baréié 1996, Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993, Jelié & Torti¢é 1973, Kranjéev 1995, Tortic 1964, Torti¢ 1966 a, Torti¢ 1966 b, unp. Matocec, unp. Tkaléec & MeS8i¢, unp. Tortic, Schulzer & al. 1866 ? (1) Bxs 2CNE (Lit.: Clémengon 1984 a) *Clitocybe obsoleta (Batsch) Quél. Ref.: Focht 1974, unp. Tkaléec & Me8sié Exs.: CNF (Lit.: Clémengon 1984 a) *Clitocybe odora (Bull. : Fr.) P. Kumm. = Agaricus odorus Bull. : Fr. (1) = Agaricus odorus Bull. : Fr. var. anisatus (Pers.) ? (2) Ref.: Focht 1979, Lisiewska & Torti¢é 1990, Schulzer & al. 1866 (1)(2), Torti¢é 1964, Tortié 1966 a, Torti¢é 1966 b, Vouk & Pevalek 1916 (1), unp. Lisiewska, unp. Matocec, unp. Tkaléec & MeSi¢, unp. Torti¢, Schulzer 1857 HR? (1) Exs.: CNF, POZM (Lit.: Bas & al. 1995, Clémengon 1984 a) Clitocybe phaeophthalma (Pers.) Kuyper = Clitocybe fritilliformis (Lasch) Gillet Misappl.: Clitocybe hydrogramma (Bull. : Fr.) P. Kumm. sensu auct. non Bull. (1) 126 Misappl.: Hyporrhodius hydrogrammus (Bull. : Fr.) J. Schr6t. sensu auct. non Bull. (2) Ref.: Bozac 1982 (1), Lisiewska & Tortié 1990 (1), Vouk & Pevalek 1916 (2), unp. Torti¢ (1) EXs<3GINE Clitocybe phyllophila (Pers. : Fr.) P. Kumm. = Agaricus phyllophilus Pers. : Fr. (1) = Clitocybe cerussata (Fr. : Fr.) P. Kumm. (2) = Clitocybe pithyophila (Fr.) Gillet (3) Ref.: Bozac 1982 ( )(2)(3), Focht 1979 (2), Focht 1983 b (2), Lisiewska & Torti¢é 1990 (2), Schulzer & al. 1866 (1), Tortié 1966 a (3), Tortic 1968 b, unp. Lisiewska (2), unp. Tortié (2), Exs.: CNF, POZM (2) Clitocybe pruinosa (Lasch) P. Kumm. = Clitocybe radicellata Gillet Ref.: Focht 1972 Clitocybe rivulosa (Pers. : Fr.) P. Kumm. Misappl.: Clitocybe dealbata (Sowerby : Fr.) P. Kumm. sensu auct. p. p. non Sowerby (sensu Sowerby = nom. dub. - Bas & al. 1995) Ref.: Focht 1979 *Clitocybe sinopica (Fr. : Fr.) P. Kumm. Ref.: Torti¢é 1966 a, Torti¢ 1966 b, Tortié 1968 a, unp. Tkaléec & MeSié, unp. Torti¢ EXSCNE (Lit.: Clémencgon 1984 a) * Clitocybe trulliformis (Fr. : Fr.) P. Karst. Ref.: unp. Tortic Exs, GNE Clitocybe tuba (Fr.) Gillet Ref.: Focht 1974 Clitocybula (Singer) Métrod (Lit.: Bollmann & al. 1996, Moser 1983, Singer 1978, Singer 1986) *Clitocybula lacerata (Scop.) Métrod = Fayodia lacerata (Scop.) Singer (1) Ref.: Torti€é 1966 a (1), Tortié 1966 b (1), unp. Tortié (1) Exs3:CNE *@var. odorata Bon Ref.: unp. Tkaléec & MeSsic Exs.: CNF (Lit.: Bon 1997) PE Collybia (Fr.) Staude = Rhodocollybia Singer (Lit.: Antonin & Noordeloos 1997, Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Clémengon 1981, Fries 1821, Fries 1874, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983) Collybia acervata (Fr. : Fr.) P. Kumm. = Gymnopus acervatus (Fr. : Fr.) Murrill Ref.: Blagai¢ 1921 b *° Collybia aquosa (Bull. : Fr.) P. Kumm. = Gymnopus aquosus (Bull. : Fr.) Antonin et Noordel. = Collybia dryophila (Bull. : Fr.) P. Kumm. var. aquosa (Bull. : Fr.) Quél. Ref.: unp. Tkaléec & MeSic¢ Exs.7CNE (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995) *Collybia butyracea (Bull. : Fr.) P. Kumm. = Agaricus butyraceus Bull. : Fr. (1) = Rhodocollybia butyracea (Bull. : Fr.) Lennox = Collybia butyracea (Bull. : Fr.) P. Kumm. var. asema (Fr. : Fr.) Quél. (2) = Collybia asema (Fr. : Fr.) P. Kumm. (3) Ref.: Lisiewska & Torti¢ 1990, Schulzer 1867 (1), Skorié 1928 (1); Torti¢ 1973, Torti¢é & Lisiewska 1978 (2), unp. Matoéec, unp. Tkaléec & Me8ic, unp. Torti¢ ( )(3) Exs.: CNF (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995, Moser 1983) Collybia cirrhata (Pers.) P. Kumm. = Agaricus cirrhatus (Pers.) ? (1) = Collybia amanitae (Batsch) Kreisel (comb. inval.) Ref.: Stoitzner 1869 (1), Torti¢é & Lisiewska 1978, unp. Lisiewska Exs.: POZM *Collybia confluens (Pers. : Fr.) P. Kumm. = Gymnopus confluens (Pers. : Fr.) Antonin, Halling et Noordel. = Marasmius archyropus (Pers.) Fr. (1) = Marasmius ingratus (Schumach. : Fr.) Quél. (2) Ref.: Blagai¢é 1921 b (2), Lisiewska & Tortié 1990, Tortic 1966 a, Tortic 1966 b, Tortié 1968 a, Tortié 1968 b, Tortié 1993, Vouk & Pevalek 1916 (1), unp. Tkaléec & MeSi¢, unp. Torti¢é Exss .CNE (Lit.: Bas & al. 1995, Clémengon 1981) Collybia cookei (Bres.) J.D. Arnold Ref.: Lisiewska & Torti¢ 1990, unp. Tortic EXss2ONE 128 *Collybia dryophila (Bull. : Fr.) P. Kumm. = Agaricus dryophilus Bull. : Fr. (1) = Gymnopus dryophilus (Bull. : Fr.) Murrill (2) Ref.: Antonin & Noordeloos 1997 (2), Baréié 1982, Barcié 1996, Focht 1974, Schulzer & al. 1866 (1), Torti¢ 1964, Torti¢ 1966 a, Torti¢ 1966 b, Tortié & Lisiewska 1978, Vouk & Pevalek 1916 (1), unp. Kotlaba (according to unp. Tortic), unp. Matocec, unp. Tkaléec & MeSic, unp. Torti¢é, Schulzer 1857 HR? (1) Exs.: CNF, PRM (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995) Collybia erythropus (Pers. : Fr.) P. Kumm. = Gymnopus erythropus (Pers. : Fr.) Antonin, Halling et Noordel. = Marasmius erythropus (Pers. : Fr.) Quél. (1) = Collybia bresadolae (Kiihner et Romagn.) Singer (2) = Collybia marasmioides (Britzelm.) Bresinsky et Stangl (3) Ref.: Barcié 1982 (3), Barcéié 1996 (3), Schulzer & al. 1866 (1), Torti¢ 1968 b, Tortié & Lisiewska 1978 (2), unp. Lisiewska (3), unp. Torti¢ Exs.: CNF, POZM (3) Collybia fuscopurpurea (Pers. : Fr.) P. Kumm. = Gymnopus fuscopurpureus (Pers. : Fr.) Antonin, Halling et Noordel. Ref.: Torti¢ 1968 b, unp. Torti¢ Exs.: CNF *Collybia fusipes (Bull. : Fr.) Quél. = Agaricus fusipes Bull. : Fr. (1) = Gymnopus fusipes (Bull. : Fr.) Gray Ref.: Blagai¢ 1921 b, Schulzer & al. 1866 (1), Tortié 1964, Torti¢é & Lisiewska 1978, Vouk & Pevalek 1916 (1), unp. Lisiewska, unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Torti¢, Schulzer 1857 HR? (1) Exs.: CNF, POZM (Lit.: Bas & al. 1995, Moser 1983) °Collybia hariolorum (Bull. : Fr.) Quél. = Agaricus hariolorum Bull. : Fr. (1) = Gymnopus hariolorum (Bull. : Fr.) Antonin, Halling et Noordel. (2) = Marasmius hariolorum (Bull. : Fr.) Quél. (3) Ref.: Antonin & Noordeloos 1997 (2), Skorié 1928 (1), Tortié 1966 a, Torti¢ 1966 b, Torti¢ 1968 b, unp. Lisiewska, unp. Matocec, unp. Tkaléec & MeSié, unp. Torti¢ ( )(3) Exs.: CNF, H, POZM, PRM (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995) *@Collybia hybrida (Kiihner et Romagn.) Svréek et Kubiéka = Gymnopus hybridus (Kiihner et Romagn.) Antonin et Noordel. Ref.: unp. Tkaléec & MeSié 129 EXS;32ONE (Lit.: Clémengon 1981, Moser 1983) *Collybia impudica (Fr.) Singer = Gymnopus impudicus (Fr.) Antonin, Halling et Noordel. (1) Ref.: Antonin & Noordeloos 1997 (1) Exs.: PRM (Lit.: Antonin & Noordeloos 1997) *Collybia maculata (Alb. et Schwein. : Fr.) P. Kumm. = Rhodocollybia maculata (Alb. et Schwein. : Fr.) Singer Ref.: Barci¢ 1982, Barcié 1996, unp. Tkaléec & MeSié, unp. Tortié Exs.: CNF (Lit.: Bas & al. 1995, Hansen & Knudsen 1992, Moser 1983) *Collybia ocior (Pers.) Vilgalys et O.K. Mill. = Gymnopus ocior (Pers.) Antonin et Noordel. (1) = Collybia exculpta (Fr.) Gillet = Collybia extuberans (Fr.) Quél. (2) = Collybia luteifolia Quél. = Collybia succinea (Fr.) Quél. Ref.: Antonin & Noordeloos 1997 (1), Focht 1974 (2), unp. Tkaléec & MeSic Exs.: BRA, CNF (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995) *Collybia peronata (Bolton : Fr.) P. Kumm. = Agaricus peronatus Bolton : Fr. (1) = Gymnopus peronatus (Bolton : Fr.) Antonin, Halling et Noordel. = Marasmius peronatus (Bolton : Fr.) Fr. (2) Ref.: Blagaié 1921 b (2), Lisiewska & Tortié 1990, Schulzer & al. 1866 (1), Tortié 1968 b, Tortié 1973, Tortié 1993, Torti¢é & Lisiewska 1978, Vouk & Pevalek 1916 (2), unp. Tkaléec & MeSi¢, unp. Tortic EXS SNE (Lit.: Antonin & Noordeloos 1997, Bas & al. 1995) Collybia tuberosa (Bull. : Fr.) P. Kumm. = Marasmius tuberosus (Bull. : Fr.) Schulzer (1) Ref.: Schulzer & al. 1866 (1), Tortié 1973, unp. Torti¢é PxseNT : * Cyphella Fr. : Fr. (Lit.: Bollmann & al. 1996, Kreisel 1987, Singer 1986) *Cyphella digitalis (Alb. et Schwein. : Fr.) Fr. Ref.: unp. Torti¢ Exs Ni 130 Delicatula Fayod (Lit.: Kreisel 1987, Moser 1983, Singer 1986) Delicatula cuspidata (Quél.) Cejp Ref.: Torti¢ 1968 b, unp. Torti¢é Exs.: CNF Dermoloma (J.E. Lange) Singer (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1986, Hansen & Knudsen 1992, Singer 1966, Watling & Turnbull 1998). °Dermoloma cuneifolium (Fr. : Fr.) Singer ex Bon = Dermoloma atrocinereum (Pers) P.D. Orton (1) = Dermoloma fuscobrunneum P.D. Orton Ref.: Tortié 1973 (1), unp. Tkaléec & Mesic EXxs: CONE WPRM;(1) (Lit.: Bas & al. 1995) *e Dermoloma josserandii Dennis et P.D. Orton = Dermoloma hygrophorus Joss. = Dermoloma phaeopodium P.D. Orton = Dermoloma pragensis Kubi¢ka (nom. inval.) Ref.: unp. Tkaléec & MeSié Exs.: CNF (Lit'? Bas’ &ai7 1995) *® Dermoloma pseudocuneifolium Herink ex Bon Ref.: unp. Tkaléec & Mesié Exs2.CNE (Lit.: Bas & al. 1995) Flammulina P. Karst. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Fries 1821, Hansen & Knudsen 1992, Kreisel 1987) °Flammulina velutipes (Curtis : Fr.) Singer = Agaricus velutipes Curtis : Fr. (1) = Collybia velutipes (Curtis : Fr.) P. Kumm. (2) Ref.: Baréic 1982, Barcié 1996, Focht 1979, Focht 1981 a, Focht 1983 a, Jeli¢é & Tortié 1973, Schulzer & al. 1866 (1), Tortié 1962 (2), Tortic 1964, Torti¢ 1966 a, Torti¢ 1966 b, Tortié 1989 b, Vouk & Pevalek 1915 (1), Vouk & Pevalek 1916 (1), unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Torti¢ (_)(2), Schulzer 1857 HR? (1) Exs.-CNF (Lit.: Bas & al. 1995) 131 Gerronema Singer (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Clémencon 1982 a, Dennis & al. 1960, Fries 1836-1838, Hansen & Knudsen 1992, Kreisel 1987, Singer 1975, Singer 1986) Gerronema ericetorum (Pers. : Fr.) Singer = Omphalina ericetorum (Pers. : Fr.) M. Lange (1) = Phytoconis ericetorum (Pers. : Fr.) Redhead et Kuyper Misappl.: Omphalia umbellifera (L. : Fr.) P. Kumm. sensu auct. non L. (2) Misappl.: Omphalina umbellifera (L. : Fr.) Quél. sensu auct. non L. (3) Ref.: Torti¢ 1966 a (3), Tortié 1966 b (3), Tortié 1968 a (1), unp. Tortié (2) Exsi7 CNP *Gerronema grossulum (Pers.) Singer = Camarophyllus grossulus (Pers.) Clémengon = Omphalina grossula (Pers.) Singer (1) = Omphalia abiegna (Berk. et Broome) J.E. Lange (2) = Omphalina abiegna (Berk. et Broome) Singer (3) = Omphalina wynniae (Berk. et Broome) S. Ito (4) Ref.: Torti¢ 1966 a (3), Torti¢ 1966 b (3), Torti¢ 1968 a (4), Torti¢ 1968 b (3), Tortié 1989 b (1), unp. Tkaléec & MeSic¢, unp. Tortié (2)(3) EXs.GNE (Lit.: Clémengon 1982 a, Hansen & Knudsen 1992, Moser 1983) Gerronema umbilicatum (Schaeff.) Singer = Clitocybe umbilicata (Schaeff.) P. Kumm. (1) = Omphalina umbilicata (Schaeff.) Quél. Refs7Focht 1972.(1) Hemimycena Singer (Lit.: Arnolds & al. 1995, Dennis & al. 1960, Hansen & Knudsen 1992, Kreisel 1987, Maas Geesteranus 1992 a, Singer 1986, Watling & Turnbull 1998) Hemimycena cucullata (Pers. : Fr.) Singer = Agaricus cucullatus Pers. : Fr. (1) = Mycena nivea Quél. (2) Ref.: GjuraSin 1898 (2), Vouk & Pevalek 1916 (1) Hemimycena gracilis (Quél.) Singer = Mycena gracilis (Quél.) Ktthner (1) Ref.: Tortié 1968 b (1), unp. Torti¢ (1) Exs:aCNE Hemimycena lactea (Pers. : Fr.) Singer = Mycena lactea (Pers. : Fr.) P. Kumm. (1) = Hemimycena delicatella (Peck) Singer (2) 132 Ref.: Focht 1974 (1), unp. Tortié (2) Exs.:\CNE Hohenbuehelia Schulzer (Lit.: Arnolds & al. 1995, Bas & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Gregory 1989) Hohenbuehelia atrocoerulea (Fr. : Fr.) Singer = Agaricus atrocoeruleus Fr. : Fr. (1) Ref.: Schulzer & al. 1866 (1), unp. Torti¢ Exs CONE ° Hohenbuehelia mastrucata (Fr. : Fr.) Singer Ref.: Stropnik & al. 1988, unp. Tkaléec & Mesié Exs:- CNE (Lit.: Bas & al. 1995, Hansen & Knudsen 1992, Watling & Gregory 1989) ° Hohenbuehelia petaloides (Bull. : Fr.) Schulzer = Pleurotus petaloides (Bull. : Fr.) Quél. (1) = Agaricus geogenius DC. : Fr. (2) = Hohenbuehelia geogenia (DC. : Fr.) Singer (3) = Pleurotus geogenius (DC. : Fr.) Gillet (4) Ref.: Baréié 1982, Barcié 1996, Focht 1972 (3), Focht 1974 (3), Focht 1986 (3), Schulzer & al. 1866, Torti¢é 1989 b, Vouk & Pevalek 1915 (2), VrSéaj 1990 (3), unp. Tkaléec & MeSic, unp. Tortié ( )(1)(3)(4) Exs.: CNF (Lit.: Bas & al. 1995, Watling & Gregory 1989) MOST PROBABLY FOUND IN THE TERRITORY OF CROATIA Hohenbuehelia repanda Huijsman Ref.: Focht 1986 HR? Hydropus Kihner ex Singer (Lit.: Arnolds & al. 1995, Bas & al. 1999, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Turnbull 1998) Hydropus atramentosus (Kalchbr.) Kotl. et Pouzar Ref.: Torti¢ 1973 Exs.: PRM Hydropus marginellus (Pers. : Fr.) Singer Ref.: Lisiewska & Torti¢é 1990, Torti¢é 1966 a, Torti¢ 1966 b, Tortié 1973, Tortié 1989 b, unp. Torti¢é Bxss. CME 133 *e Hydropus subalpinus (Hohn.) Singer Ref.: unp. Tkaléec & Me8Sié¢, unp. Tortié xs e7CINT (Lit.: Bas & al. 1999, Moser 1983, Watling & Turnbull 1998) Hypsizygus Singer (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Moser 1983, Singer 1986, Watling & Gregory 1989) Hypsizygus circinatus (Fr.) Singer Ref.: Tortié 1989 b, unp. Torti¢ °Hypsizygus ulmarius (Bull. : Fr.) Redhead = Lyophyllum ulmarium (Bull. : Fr.) Ktthner (1) = Panus ulmarius (Bull. : Fr.) Schulzer (2) = Pleurotus ulmarius (Bull. : Fr.) P. Kumm. (3) Ref.: Schulzer & al. 1866 (2), Torti¢ 1964 (1), Tortié 1989 b (1), unp. Tkaléec & MeSi¢, unp. Tortié (1)(3) Exse CONF (Lit.: Moser 1983, Pilat 1935, Watling & Gregory 1989) Laccaria Berk. et Broome (Lit.: Arnolds & al. 1995, Bas & al. 1995, Clémengon 1984 b, Fries 1821, Hansen & Knudsen 1992, Kreisel 1987, Mueller & Vellinga 1986) *Laccaria amethystina (Huds. —) Cooke = Clitocybe laccata (Scop. : Fr.) P. Kumm. var. amethystina (Huds. > Cooke) ? (1) = Laccaria laccata (Scop. : Fr.) Berk. et Broome var. amethystina (Huds. — Cooke) Rea (2) = Laccaria amethystea (Bull. Gray) Murrill (3) Ref.: Baréié 1982, Barcié 1996, Blagaié 1921 b (1), GjuraSin 1898 (2), Lisiewska & Tortié 1990 (3), Torti¢ 1964 (3), Tortié 1966 a (3), Tortic 1966 b (3), Torti¢é 1968 a (3), Tortié & Lisiewska 1978, unp. Matoéec, unp. Tkaléec & Me8ic¢, unp. Tortié ( )(3) Exs.: BEO, CNF (Lit.: Bas & al. 1995, Clémengon 1984 b) *Laccaria bicolor (Maire) P.D. Orton Ref.: unp. Matocec ExsGink °Laccaria laccata (Scop. : Fr.) Berk. et Broome = Agaricus laccatus Scop. : Fr. (1) = Russuliopsis laccata (Scop. : Fr.) J. Schrot. (2) = Laccaria tetraspora Singer (3) 134 Ref.: Baréié 1982, Barcic 1996, KiSpati¢ 1950 (2), Jeli¢ & Tortié 1973, Lisiewska & Tortié 1990, Schulzer 1867 (1), Skorié 1928 (1), Tortié 1964, Tortié 1966 a, Tortié 1966 b, Torticé 1993, Tortié & Lisiewska 1978, Vouk & Pevalek 1916 (1), unp. Matoéec ( )(3), unp. Tkaléec & Me8ic¢, unp. Tortic, Schulzer 1857 HR? (1) Exst.ONF (Lit.: Bas & al. 1995, Clémencgon 1984 b) Laccaria proxima (Boud.) Pat. Ref.: Lisiewska & Torti¢ 1990 Lachnella Fr. (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Singer 1986) Lachnella alboviolascens (Alb. et Schwein. : Fr.) Fr. = Cyphella alboviolascens (Alb. et Schwein. : Fr.) Crouan (1) Ref.: Jaap 1916 (1) Lachnella villosa (Pers. : Fr.) Gillet = Cyphella villosa (Pers. : Fr.) Crouan (1) Ref.: Jaap 1916 (1), Picbauer 1928 (1) Lepista (Fr.) W.G. Sm. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Bon 1983, Bon 1997, Dennis & al. 1960, Hansen & Knudsen 1992, Konrad & Maublanc 1924-1937, Kreisel 1987, Moser 1983, Pilat 1951 b, Singer 1986) *° Lepista caespitosa (Bres.) Singer Ref.: unp. Tkaléec & MeSié Exss,G NG (Lit.: Moser 1983) *Lepista flaccida (Sowerby : Fr.) Pat. = Agaricus flaccidus Sowerby : Fr. (1) = Clitocybe flaccida (Sowerby : Fr.) P. Kumm. (2) = Agaricus infundibuliformis Schaeff. = Clitocybe infundibuliformis (Schaeff.) Quél. non sensu auct. = Agaricus inversus Scop. (3) = Clitocybe inversa (Scop.) Quél. (4) = Lepista inversa (Scop.) Pat. Ref.: Focht 1972 (4), Focht 1974 (2), Focht 1979 (2), Gjurasin 1898 (4), Muzic & BoZac 1997 (4), Schulzer & al. 1866 (1), Skorié 1928 (3), Torti¢ 1964 (4), Torti¢ 1966 a (4), Tortié 1966 b (4), Tortié & Lisiewska 1978 (4), unp. Matocec (4), unp. Tkaléec & MeSié, unp. Tortié (4) Exs.: CNF (Lit.: Bas & al. 1995, Hansen & Knudsen 1992, Moser 1983) 135 *° LT epista gilva (Pers. : Fr.) Pat. = Clitocybe gilva (Pers. : Fr.) P. Kumm. (1) Ref.: unp. Tkaléec & Me8ic, unp. Torti¢ (1) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) °Lepista glaucocana (Bres.) Singer = Tricholoma nudum (Bull. : Fr.) P. Kumm. var. glaucocanum (Bres.) L. Maire (1) | Ref.: Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993, unp. Matocec, unp. Tkaléec & MeSic, unp. Tortié ( )(1) Exs:)CNE (Lit.: Bon 1983, Moser 1983) Lepista irina (Fr.) H.E. Bigelow = Tricholoma irinum (Fr.) P. Kumm. (1) Ref.: Bozac 1982 (1), Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993 °Lepista nuda (Bull. : Fr.) Cooke = Agaricus nudus Bull. : Fr. (1) = Rhodopaxillus nudus (Bull. : Fr.) Maire (2) = Tricholoma nudum (Bull. : Fr.) P. Kumm. (3) Ref.: Baréié 1989, Baréié 1996, Blagai¢ 1921 b (3), Blagaié 1931 (3), Focht 1972 (2), Focht 1979 (2), Focht 1986, GjuraSin 1898 (3), Jelic & Tortié 1973, Torti¢ 1964, Tortié 1966 a, Torti¢é 1966 b, Torti¢ & Lisiewska 1978, Vouk & Pevalek 1916 (1), unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Torti¢ ( )(2)(3) Exs.: CNF (Lit.: Bon 1983, Moser 1983) °Lepista saeva (Fr.) P.D. Orton = Agaricus bicolor Pers. (1) Misappl.: Lepista personata (Fr. : Fr.) Cooke sensu auct. non Fr. (2) (sensu Fr. = Lepista nuda (Bull. : Fr.) Cooke - Dennis & al. 1960) Ref.: Vouk & Pevalek 1916 (1), Vrséaj 1990 (2), unp. Tkaléec & Me8ic EXsaGNr . (Lit.: Bas & al. 1995, Moser 1983) Lepista sordida (Schumach. : Fr.) Singer = Rhodopaxillus sordidus (Schumach. : Fr.) Maire (1) Ref.: Tortié 1968 b, unp. Torti¢ (1) Exss ONE Leucopaxillus Boursier (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Bon 1991, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Schulzer 1867, Schulzer & al. 1866, Singer 1986) 136 Leucopaxillus candidus (Bres.) Singer = Agaricus candidus (Bres.) Skorié (1) = Clitocybe candida Bres. (2) Ref.: Bozac 1984 (2), Bozac 1986 (2), Bozac 1989 (2), Bozac 1993 (2), Schulzer 1885 (2), Skorié 1928 (1), Vrséaj 1990 Leucopaxillus cerealis (Lasch) Singer = Leucopaxillus lentus (H. Post) Singer et A.H. Sm. (1) Ref.: Lisiewska & Tortié 1990 (1), unp. Tortié (1) Exs.: CNF °Leucopaxillus compactus (Fr.) Neuhoff = Leucopaxillus tricolor (Peck) Kiihner Ref.: VrSéaj 1990, unp. Tkaléec & MeSié Exs:;CNE (Lit.: Hansen & Knudsen 1992, Moser 1983) *Leucopaxillus gentianeus (Quél.) Kotl. Misappl.: Clitocybe amara (Alb. et Schwein. : Fr.) P. Kumm. sensu auct. non Alb. et Schwein. (1) Misappl.: Leucopaxillus amarus (Alb. et Schwein. : Fr.) Kiihner sensu auct. non Alb. et Schwein. (2) Ref.: Tortié 1964 (2), Tortié 1966 a (2), Torti¢é 1966 b (2), Torti¢ 1968 a (2), Torti¢ & Jeli¢ 1970 (2), unp. Lisiewska, unp. Tkaléec & MeSié, unp. Tortié (1)(2) Exs.: CNF, POZM (Lit.: Hansen & Knudsen 1992, Moser 1983) Leucopaxillus giganteus (Sibth. : Fr.) Singer = Clitocybe gigantea (Sibth. : Fr.) Quél. (1) = Paxillus giganteus (Sibth. : Fr.) Fr. (2) Ref.: Bartié 1982 (1), Baréié 1996 (1), Blagai¢é 1931 (2), Focht 1979 (1) °*Leucopaxillus macrocephalus (Schulzer) Bohus = Agaricus macrocephalus Schulzer (1) = Porpoloma macrocephalum (Schulzer) Bon = Agaricus megacephalus Schulzer (nom. nud.) (2) - as Agaricus macrorhizus Lasch (nom. illeg.) (3) - as Tricholoma macrorhizum (Lasch) —> Sacc. (4) Ref.: Kalchbrenner & Schulzer 1873-1877 (1), Schulzer & al. 1866 (2) + Schulzer 1867 (nom. nud.) (1), Fries 1874 HR? (3), Saccardo 1887 HR? (4), unp. Tkaléec & MeSi¢ Exsi3CNE (Lit.: Bon 1991, Moser 1983) Annotation: Fries (1874) and Saccardo (1887) quote Schulzer's records (they do not precisely quote the locality) regarding Agaricus macro- cephalus a synonym of A. macrorhizus. Bon (1991) considers that A. macrocephalus and A. macrorhizus are two different species of genus Porpoloma. Lory Leucopaxillus paradoxus (Costantin et L.M. Dufour) Boursier = Clitocybe paradoxa Costantin et L.M. Dufour (1) Ref.: Torti¢é 1964, VrSéaj 1990, unp. Lisiewska, unp. Tortié ( )(1) Exs.: CNF, POZM Lyophyllum P. Karst. = Tephrocybe Donk (Lit.: Arnolds & al. 1995, Bollmann & al. 1996, Clémencon 1986, Dennis & al. 1960, Hansen & Knudsen 1992, Kreisel 1987, Kiihner & Romagnesi 1953, Moser 1983, Pilat 1951 b, Singer 1986) Lyophyllum atratum (Fr. : Fr.) Singer = Agaricus atratus Fr. : Fr. (1) = Tephrocybe atrata (Fr. : Fr.) Donk Ref.: Skorié 1928 (1) Lyophyllum connatum (Schumach. : Fr.) Singer Ref.: Focht 1972, Focht 1979, Tortié 1966 a, Tortié 1966 b, Tortié 1968 b, unp. Matocec, unp. Tortic Exss NE °*Lyophyllum deliberatum (Britzelm.) Kreisel = Lyophyllum infumatum (Bres.) Kithner (1) Ref.: Torti¢é 1968 b (1), Torti¢ & Lisiewska 1978 (1), unp. Tkaléec & Me8ic¢, unp. Torti¢ (1) Exs.: CNF (Lit.: Clémengon 1986) Lyophyllum fumosum (Pers. : Fr.) P.D. Orton sensu lato Incl.: Lyophyllum aggregatum (Schaeff.) Kithner (1) Lyophyllum cartilagineum (Bull.) ? (2) Lyophyllum cinerascens (Bull.) Konrad et Maubl. (3) Lyophyllum conglobatum (Vittad.) Moser (4) = Clitocybe conglobata (Vittad.) Bres. (5) = Tricholoma conglobatum (Vittad.) Ricken (6) = Lyophyllum aggregatum (Schaeff.) Kithner subsp. conglobatum (Vittad.) ? (7) Lyophyllum decastes (Fr. : Fr.) Singer (8) Lyophyllum fumosum (Pers. : Fr.) P.D. Orton (9) = Agaricus fumosus Pers. : Fr. (10) Lyophyllum loricatum (Fr.) Kthner (11) Lyophyllum tumulosum (Kalchbr.) ? (12) = Tricholoma tumulosum (Kalchbr.) Barla (13) = Agaricus tumulosus Kalchbr. (14) = Lyophyllum aggregatum (Schaeff.) Ktihner sensu lato (15) 138 Ref.: Barcic 1982 (8), Barci¢ 1996 (8), Blagai¢ 1931 (6)(13), Bozac 1984 (12), Bozac 1989 (2), Bozac 1993 (2), Focht 1972 (7), Focht 1979 (1)(2)(3), Focht 1981 b (11), Focht 1983 a (11), Focht 1986 (8)(15), Schulzer 1880 b (14), Skorié 1928 (10), Tortié 1964 (4), Tortié 1966 a (15), Tortié 1966 b (1), Tortié & Lisiewska 1978 (9), unp. Lisiewska (9), unp. Torti¢ (5)(6)(11) Exs.: CNF, POZM (9) Annotation: Most modern authors distinguish three species here, but disagreements exist about species concepts, synonymy and nomenclature (see Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983). Lyophyllum inolens (Fr.) Singer = Agaricus inolens Fr. (1) = Tephrocybe inolens (Fr.) Moser Ref.: Vouk & Pevalek 1916 (1) *°Lyophyllum macrosporum Singer Ref.: unp. Tkaléec & Me8Si¢ Exs7 ONE (Lit.: Clémengon 1986) Lyophyllum paelochroum Clémencgon Misappl.: Lyophyllum immundum (Berk.) Kithner sensu auct. non Berk. (1) (sensu Berk. = nom. dub. - Clémengon 1986) Ref.: Torti¢ 1966 a (1), unp. Tortié (1) Bxsa_CNE *Lyophyllum rancidum (Fr. : Fr.) Singer = Tephrocybe rancida (Fr. : Fr.) Donk (1) Ref.: Jelié & Tortié 1973 (1), unp. Tkaléec & MeSié, unp. Tortié (1) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) *@Lyophyllum rhopalopodium Clémencon Ref.: unp. Tkaléec & MeSic . Exs:,CNE (Lit.: Clémengon 1986) *Lyophyllum semitale (Fr. : Fr.) Kiihner Ref.: Bozac 1984, Focht 1972, unp. Tkaléec & MeSi¢ EXSesOINP) (Lit.: Clémen¢gon 1986) *Lyophyllum transforme (Britzelm.) Singer = Lyophyllum trigonosporum (Bres.) Kihner (1) Ref.: Torti¢é 1968 b (1), unp. Tkaléec & MeSié, unp. Tortié (1) Exs.GNE (Lit.: Clémengon 1986) 3° *Macrocystidia Joss. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983) *® Macrocystidia cucumis (Pers. : Fr.) Joss. Ref.: unp. Mato¢éec, unp. Tkaléec & Me8ic¢, unp. Torti¢é Exs:; CNE (Lit.: Bas & al. 1995, Moser 1983) Marasmiellus Murrill (Lit.: Antonin & Noordeloos 1993, Antonin & Noordeloos 1997, Arnolds & al. 1995, Bas & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Schulzer 1867, Schulzer & al. 1866, Singer 1986) * Marasmiellus candidus (Bolton) Singer = Marasmiellus albus-corticis (Secr.) ex Singer (1) Ref.: unp. Tortié (1) EXSONE Marasmiellus omphaliformis (Kiihner) Noordel. = Marasmius omphaliformis Kihner (1) Ref.: Focht 1974 (1) Marasmiellus ramealis (Bull. : Fr.) Singer = Agaricus ramealis Bull. : Fr. (1) = Marasmius ramealis (Bull. : Fr.) Fr. (2) = Marasmiellus amadelphus (Bull. : Fr.) Moser = Marasmius exalbidus Schulzer (nom. nud.) (3) Ref.: Schulzer & al. 1866 (3) + Schulzer 1867 (1), Torti€é 1968 b (2), Torti¢ 1989 b, Torti¢é & Lisiewska 1978, unp. Torti¢ (2) ExXs,GNE *Marasmiellus tricolor (Alb. et Schwein. : Fr.) Singer Ref.: unp. Tortié Exs:) GNF Marasmius Fr. = Setulipes Antonin (Lit: Antonin & Noordeloos 1993, Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Clémengon 1982 b, Fries 1821, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986) °*Marasmius alliaceus (Jacq. : Fr.) Fr. = Marasmius alliaceus (Jacq. : Fr.) Fr. var. subtilis J.E. Lange (nom. inval.) (1) 140 Ref.: Antonin & Noordeloos 1993, Blagai¢é 1931, Lange 1974, Lisiewska 1993, Lisiewska & Tortié 1990, Tortié 1966 a, Tortié 1966 b, Tortié 1968 a, Torti¢ 1968 b, Tortié 1989 b, Tortié 1993, Vouk & Pevalek 1915, unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Tortié ( )(1) Exs.: BEO, BRNM, CNF (Lit.: Bas & al. 1995, Clémencon 1982 c) ®Marasmius androsaceus (L. : Fr.) Fr. = Setulipes androsaceus (L. : Fr.) Antonin (1) Ref.: Antonin & Noordeloos 1993 (1), Lisiewska & Torti¢é 1990, Moesz 1938, Skorié 1928, Tortié 1966 a, Tortié 1966 b, Tortié 1993, Vouk & Pevalek 1916, unp. Matocec, unp. Tkaléec & MeSic¢, unp. Tortic Exs.: BP, CNF (Lit.: Bas & al. 1995, Clémencon 1982 c) Marasmius bulliardii Quél. Ref.: Jelié & Torti¢é 1973, Lisiewska & Torti¢é 1990, Torti¢ 1968 b, unp. Lisiewska, unp. Torti¢é Exs.: CNF, POZM Marasmius cohaerens (Pers. : Fr.) Cooke et Quél. = Agaricus cohaerens Pers. : Fr. (1) Ref.: Skorié 1928 (1), unp. Tortié, Schulzer 1857 HR? (1) ExsiCNE Marasmius corbariensis (Roum.) Singer = Marasmius hygrometricus (Brig.) Sacc. (1) Ref.: Saccardo 1887 (1) *®@ Marasmius curreyi Berk. et Broome Misappl.: Marasmius graminum (Lib.) Berk. sensu auct. recent. non Lib. (1) Ref.: unp. Tkaléec & Me8i¢, unp. Tortié (1) Exs- 7G NF (Lit.: Bas & al. 1995, Clémengon 1982 b) Marasmius epiphyllus (Pers. : Fr.) Fr. = Agaricus epiphyllus Pers. : Fr. (1) Ref.: Focht 1974, Schulzer & al. 1866, Schulzer 1857 HR? (1) °Marasmius oreades (Bolton : Fr.) Fr. = Agaricus oreades Bolton: Fr. (1) Ref.: Blagaié 1921 a, Blagai¢é 1931, Bozac 1982, Focht 1972, Focht 1974, Focht 1979, GjuraSin 1898, Lorinser - Janda 1877, Schulzer & al. 1866 (1), Torti¢ 1966 a, Tortié 1966 b, Tortié 1968 b, unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Torti¢é Exs.: CNF (Lit.: Bas & al. 1995, Clémengon 1982 b) 141 Marasmius querceus Britzelm. Misappl.: Marasmius prasiosmus (Fr. : Fr.) Fr. sensu Fr. 1836-1838 et auct. non Fr. 1818 (1) Ref.: Blagai¢é 1921 b (1), GjuraSin 1898 (1), unp. Tortié (1) BXSa UND *° Marasmius quercophilus Pouzar = Setulipes quercophilus (Pouzar) Antonin Misappl.: Marasmius splachnoides (Hornem. : Fr.) Fr. sensu auct. non Hornem. Ref.: unp. Tkaléec & MeSi¢ EXSe:G NE (Lit.: Bas & al. 1995, Clémengon 1982 b) °Marasmius rotula (Scop. : Fr.) Fr. = Agaricus rotula Scop. : Fr. (1) Ref.: Blagai¢é 1921 b, Lisitewska & Torti¢ 1990, Schulzer 1882 a (1), Schulzer & al. 1866, Skorié 1928, Tortié 1966 a, Tortié 1966 b, Tortié 1968 b, Torti¢é 1973, Tortié 1979, Tortié 1989 b, Torti¢ 1993, Tortié & Lisiewska 1978, Vouk & Pevalek 1916, unp. Matocec, unp. Tkaléec & Me8ic, unp. Tortic, Schulzer 1857 HR? (1) Bxs: ONE (Lit.: Bas & al. 1995, Clémengon 1982 b) ®*Marasmius scorodonius (Fr. : Fr.) Fr. = Marasmius alliatus (Schaeff.) J. Schroét. (1) Ref.: Blagai¢é 1921 b, Blagai¢ 1931, Focht 1974, Lisiewska & Torti¢ 1990, Skorié 1928 (1), Tortié 1966 a, Tortié 1966 b, Tortié 1968 b, Vouk & Pevalek 1916 (1), unp. Tkaléec & MeSi¢, unp. Torti¢é Exs.., CNE (Lit.: Bas & al. 1995, Clémencgon 1982 b) °Marasmius torquescens Quél. Misappl.: Marasmius lupuletorum (Weinm.) Bres. sensu auct. non Weinm. (1) (sensu Weinm. = nom. dub. - Clémengon 1982 b) Ref.: Lisiewska & Torti¢ 1990 (1), Tortié 1993 (1), unp. Lisiewska (1), unp. Tkaléec & Me8ic, unp. Torti¢ (1) Exs.: CNF, POZM (1) (Lit.: Bas & al. 1995, Clémencgon 1982 b) *Marasmius undatus (Berk.) Fr. = Marasmius chordalis Fr. (1) Ref.: unp. Torti¢ (1) Exs,: CNE *Marasmius wynnei Berk. et Broome = Marasmius bresadolae Schulzer (1) 142 Ref.: Lisiewska & Torti¢ 1990, Saccardo 1887 (1), Schulzer 1885 (1), Torti¢ 1968 b, unp. Tkaléec & MeSic, unp. Tortic Exs.: CNF (Lit.: Bas & al. 1995, Clémengon 1982 b) Megacollybia Kotl. et Pouzar (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Watling & Turnbull 1998) °Megacollybia platyphylla (Pers. : Fr.) Kotl. et Pouzar = Collybia platyphylla (Pers. : Fr.) P. Kumm. (1) = Oudemansiella platyphylla (Pers. : Fr.) Moser (2) Ref.: Jelié & Torti¢é. 1973 (2), Lisiewska & Tortié 1990, Tortié 1966 a (2), Tortié 1966 b (2), Torti¢ 1968 b (2), Tortié 1989 b (2), Tortic 1993, Torti¢ & Lisiewska 1978 (2), unp. Lindtner (1) (according to unp. Tortic), unp. Matocéec (1), unp. Tkaléec & MeSsic¢, unp. Tortié (1)(2) Exs.: BEO (1)(2), CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) Melanoleuca Pat. (Lit.: Arnolds & al. 1995, Bas & al. 1999, Bollmann & al. 1996, Bon 1991, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Turnbull 1998) Melanoleuca brevipes (Bull. : Fr.) Pat. = Agaricus brevipes Bull. : Fr. (1) Ref.: Schulzer 1867 (1), unp. Torti¢ ExsacCNE *Melanoleuca cognata (Fr.) Konrad et Maubl. Ref.: Bozac 1984, Torti¢ 1973, unp. Tkaléec & MeSic, unp. Tortic EXs.C NE (Lit.: Bas & al. 1999) Melanoleuca grammopodia (Bull. : Fr.) Pat. Ref.: Torti¢ 1964, unp. Torti¢ Melanoleuca melaleuca (Pers. : Fr.) Murrill sensu lato Incl.: Melanoleuca melaleuca (Pers. : Fr.) Murrill (1) Melanoleuca polioleuca (Fr.) Kithner et Maire Melanoleuca vulgaris (Pat.) Pat. (2) Ref.: Barcié 1982 (1), Barcié 1996 (1), Focht 1972 (1), Focht 1974 (2), Torti¢ 1973 (1), unp. Tortié ( )(1) Exsa2GNE Annotation: These three species belong to the species complex about which different opinions exist (see Arnolds & al. 1995, Bon 1991, Hansen & Knudsen 1992, Kreisel 1987, Watling & Turnbull 1998). 143 Melanoleuca schumacheri (Fr. : Fr.) Singer = Agaricus schumacheri Fr. : Fr. (1) Ref.: Vouk & Pevalek 1915 (1) Melanoleuca Strictipes (P. Karst.) Murrill Ref.: Baréié 1982, Baréié 1996 Micromphale (Nees) Gray (Lit.: Antonin & Noordeloos 1997, Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Hansen & Knudsen 1992, Kreisel 1987) °Micromphale brassicolens (Romagn.) P.D. Orton °var. brassicolens = Gymnopus brassicolens (Romagn.) Antonin et Noordel. var. brassicolens (1) = Marasmius brassicolens Romagn. (2) Ref.: Antonin & Noordeloos 1997 (1), unp. Tkaléec & Me8Sic¢, unp. Tortié (2) Exs.: CNF, PRM (Lit.: Antonin & Noordeloos 1997) °Micromphale foetidum (Sowerby : Fr.) Singer = Marasmiellus foetidus (Sowerby : Fr.) Antonin, Halling et Noordel. = Marasmius foetidus (Sowerby : Fr.) Fr. (1) Ref.: Focht 1974 (1), Torti¢é 1968 b (1), Tortié 1989 b, unp. Tkaltéec & MeSic¢, unp. Torti¢ (1) Exs.: CNF (Lit.: Antonin & Noordeloos 1997) °Micromphale perforans (Hoffm. : Fr.) Gray = Marasmiellus perforans (Hoffm. : Fr.) Antonin, Halling et Noordel. = Marasmius perforans (Hoffm. : Fr.) Fr. (1) Ref.: Blagaié 1931 (1), Lisiewska & Torti¢é 1990, Stoitzner 1869 (1), Tortié 1966 a (1), Tortié 1966 b (1), Tortié 1968 b (1), unp. Tkaléec & MeSic¢, unp. Torti¢ (1) Exs.: CNF (Lit.: Antonin & Noordeloos 1997) Mycena (Pers.) Roussel (Lit.: Arnolds & al. 1995, Hansen & Knudsen 1992, Kreisel 1987, Lisiewska 1987, Maas Geesteranus 1992 a, Maas Geesteranus 1992 b, Moser 1983, Singer 1986) Mycena acicula (Schaeff.) P. Kumm. = Agaricus acicula Schaeff. (1) Ref.: Schulzer & al. 1866 (1), Schulzer 1857 HR? (1) 144 Mycena amicta (Fr. : Fr.) Quél. Ref.: Torti¢ 1973, Torti¢é 1989 b, unp. Tortié Exs: CNEGh Mycena atroalba (Bolton : Fr.) Gray Ref.: Lisiewska & Tortié 1990, unp. Torti¢ Mycena aurantiomarginata (Fr. : Fr.) Quél. Ref.: Torti¢ 1973, unp. Torti¢ Exs.: CNF Mycena capillaris (Schumach. : Fr.) P. Kumm. Ref.: Vouk & Pevalek 1915 Mycena crocata (Schrad. : Fr.) P. Kumm. = Agaricus crocatus Schrad. : Fr. (1) Ref.: Blagai¢é 1921 b, Lange 1974, Lisiewska 1993, Lisiewska & Tortié 1990, Skorié 1928 (1), Tortié 1968 b, Tortié 1973, Tortié 1989 b, unp. Lisiewska, unp. Torti¢ Exs.: CNF, POZM, PRM Mycena epipterygia (Scop. : Fr.) Gray = Agaricus epipterygius Scop. : Fr. (1) = Mycena citrinella (Pers. : Fr.) P. Kumm. (2) = Mycena viscosa Maire (3) Ref.: GjuraSin 1898 (2), Tortié 1964, Tortié 1966 a ( )(3), Tortié 1966 b ( )(3), Torti€é 1968 b (3), Tortié 1989 b (3), Tortié 1994 (3), Tortié & Lisiewska 1978, Vouk & Pevalek 1915 (1), unp. Tortié ( )(3) Exs.: CNF Mycena erubescens Hohn. Ref.: Torti¢é 1973, unp. Torti¢ Exs.: CNF, PRM Mycena fagetorum (Fr.) Gillet = Agaricus marcescens Schulzer (1) Ref.: Schulzer 1878 (1) Mycena filopes (Bull. : Fr.) P. Kumm. = Agaricus filopes Bull. : Fr. (1) = Marasmius filopes (Bull. : Fr.) Schulzer (2) = Mycena iodiolens S. Lundell (3) Misappl.: Mycena vitilis (Fr.) Quél. sensu Kiihner non Fr. Ref.: Lisiewska & Tortié 1990, Schulzer 1882 a (1), Schulzer & al. 1866 (2), Vouk & Pevalek 1915 (1), Vouk & Pevalek 1916 (1), unp. Tortié ( )(3) Exs.: CNF Mycena flavoalba (Fr.) Quél. Ref.: Lisiewska & Tortié 1990, Tortié 1973, unp. Lisiewska, unp. Tortié Exs.: CNF, POZM, PRM 145 Mycena galericulata (Scop. : Fr.) Gray = Agaricus galericulatus Scop. : Fr. (1) = Marasmius galericulatus (Scop. : Fr.) Schulzer (2) = Agaricus galeriformis Schulzer (3) = Agaricus pseudoclypeatus Bolton var. radicans Schulzer (4) = Agaricus pseudoclypeatus Bolton var. slavonicus Schulzer (5) = Agaricus rugatoplicatus Schulzer (6) = Agaricus rugatulosus Schulzer (7) = Agaricus rugosus Fr. var. stipite-elongato Schulzer (8) = Mycena sparsa (Bres. et Schulzer) Sacc. (9) = Mycena galericulata (Scop. : Fr.) Gray var. sparsa Bres. et Schulzer (nom. illeg.) (10) = Mycena sudora (Fr.) Gillet (11) Ref.: GjuraSin 1898 (11), Jeli¢ & Tortié 1973, Picbauer 1928, Pilat 1925, Schulzer 1878 (8), Schulzer 1879 (4), Schulzer 1880 a (5)(6)(7), Schulzer 1881 (3), Schulzer 1882 a (1), Schulzer 1885 (10), Schulzer & al. 1866 (2), Tortié 1966 a, Tortié 1968 b, Tortié 1973, Tortié 1989 b, Tortié 1993, Tortié & Lisiewska 1978, unp. Torti¢é, Saccardo 1887 HR? (9), Schulzer 1857 HR? (1) Exs.: CNF °Mycena galopus (Pers. : Fr.) P. Kumm. ("galopoda") Ref.: Baréié 1982, Barcié 1996, Jelié & Torti¢ 1973, Lisiewska & Torti¢ 1990, Tortié 1973, Torti¢ & Lisiewska 1978, unp. Tkaléec & MeSic¢, unp. Torti¢ Exs.: CNF (Lit.: Maas Geesteranus 1992 b, Moser 1983) Mycena haematopus (Pers. : Fr.) P. Kumm. ("haematopoda") Ref.: Tortié 1964, Tortié 1966 a, Torti¢ 1966 b, Torti¢é 1979, Tortic 1989 b, Tortié 1993, unp. Torti¢ Exs.: CNF *Mycena hiemalis (Osbeck) Quél. Ref.: unp. Torti¢é Exs.: CNF Mycena inclinata (Fr.) Quél. = Agaricus amoenipes Schulzer var. hirtipes Schuler (1) Ref.: Jelié & Tortié 1973, Schulzer 1878 (1), Tortié 1968 b, Tortic & Lisiewska 1978, unp. Lisiewska, unp. Torti¢ Exs.: CNF, POZM Mycena latifolia (Peck) A.H. Sm. Ref.: Torti¢ 1973, unp. Torti¢é bxse Ny Mycena leptocephala (Pers. : Fr.) Gillet = Mycena chlorinella (J.E. Lange) Singer (1) Ref.: Lisiewska & Tortié 1990 (1), unp. Torti¢ (1) 146 Mycena maculata P. Karst. Ref.: Tortié 1968 b, unp. Torti¢é Exs CN iy. Mycena metata (Fr. : Fr.) P. Kumm. = Mycena phyllogena (Pers.) Singer (1) Misappl.: Mycena tenella (Schumach.) Quél. sensu Ricken, Kihner non Schumach. (2) (sensu Schumach. = nom. dub. - Maas Geesteranus 1992 a) Ref.: Tortié 1973 (1), unp. Torti¢ (2) Exs-CNE Mycena niveipes (Murrill) Murrill Ref.: Torti¢é 1968 b, unp. Torti¢ Exs*s CNP °Mycena pelianthina (Fr. : Fr.) Quél. Ref.: Baréié 1982, Baréié 1996, Lange 1974, Lisiewska & Torti¢ 1990, Tortié 1973, unp. Lisiewska, unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Torti¢ Exs.: CNF, POZM (Lit.: Hansen & Knudsen 1992, Maas Geesteranus 1992 b) Mycena polygramma (Bull. : Fr.) Gray = Agaricus polygrammus Bull. : Fr. (1) = Marasmius polygrammus (Bull. : Fr.) Schulzer (2) Ref.: Schulzer 1857 (1), Schulzer & al. 1866 (2), Torti¢ & Lisiewska 1978, Vouk & Pevalek 1915 (1), Vouk & Pevalek 1916 (1), unp. Tortic¢ Exs.: CNF °Mycena pura (Pers. : Fr.) P. Kumm. = Agaricus purus Pers. : Fr. (1) Ref.: Baréié 1989, Baréi¢ 1996, Focht 1974, GjuraSin 1898, Lisiewska & Tortié 1990, Schulzer & al. 1866 (1), Tortié 1964, Torti¢ 1966 a, Torti¢ 1966 b, Tortié 1994, Torti¢é & Lisiewska 1978, unp. Matocec, unp. Tkaléec & MeSi¢, unp. Torti¢ Exs.: CNF (Lit.: Hansen & Knudsen 1992, Maas Geesteranus 1992 b) °Mycena renati Queél. = Mycena flavipes Quél. (nom. illeg.) (1) Ref.: Blagai¢é 1921 b, Lisiewska & Torti¢é 1990, Torti¢ 1973, Tortié 1989 b, unp. Lisiewska, unp. Tkaléec & MeSic¢, unp. Torti¢ ( )(1) Exs.: CNF, POZM | (Lit.: Maas Geesteranus 1992 b) *° Mycena rorida (Fr. : Fr.) Quél. Ref.: unp. Tkaléec & MeSi¢, unp. Torti¢é Exs CONE (Lit.: Hansen & Knudsen 1992, Moser 1983) 147 Mycena rosea (Bull. >) Gramberg = Agaricus roseus Bull. (nom. illeg.) (1) = Agaricus purus Pers. : Fr. var. roseus (Bull. +) Pers. (2) Ref.: Vouk & Pevalek 1916 (1), unp. Tortié, Schulzer 1857 HR? (2) ¢Mycena rosella (Fr. : Fr.) P. Kumm. = Agaricus roseus Pers. (nom. illeg.) Ref.: Torti¢é 1973, unp. Tkaléec & MeSié, unp. Tortié EXS7 CINE (Lit.: Hansen & Knudsen 1992, Moser 1983) Mycena rubromarginata (Fr. : Fr.) P. Kumm. Ref.: Lisiewska & Torti¢ 1990, Tortié 1989 b, unp. Tortié EXs.,. ONE Mycena sanguinolenta (Alb. et Schwein. : Fr.) P. Kumm. Ref.: KiSpati¢ 1950, Lisiewska & Tortié 1990, unp. Tortié Exs.: CNF °*Mycena seynii Quél. Ref.: Focht 1974, unp. Matoéec, unp. Tkaléec & MeSi¢ Exs.: CNF (Lit.: Maas Geesteranus 1992 b, Moser 1983) *Mycena speirea (Fr. : Fr.) Gillet Ref.: unp. Torti¢ Exs.: CNF Mycena stipata Maas Geest. et Schwébel Misappl.: Mycena alcalina (Fr. : Fr.) P. Kumm. sensu auct. non Fr. (1) (sensu Fr. = nom. dub. - Maas Geesteranus 1992 a) Ref.: Focht 1974 (1), Tortié 1966 a (1), Torti¢ 1966 b (1), Tortié 1989 b (1), Torti¢é 1993 (1), unp. Torti¢ (1) Exs GNF Mycena stylobates (Pers. : Fr.) P. Kumm. = Marasmius torquatus (Fr. : Fr.) Fr. (1) Ref.: Lisiewska & Tortié 1990, Skorié 1928 (1), unp. Tortié Exs 3CNE Mycena viridimarginata P. Karst. Ref.: Torti¢ 1973, unp. Lisiewska, unp. Torti¢ Exs.: CNF, POZM Mycena vitilis (Fr.) Quél. Ref.: Focht 1972, Tortié & Lisiewska 1978, unp. Torti¢ *Mycena vulgaris (Pers. : Fr.) P. Kumm. Ref.: unp. Torti¢é Exs.: CNF 148 Mycena zephirus (Fr. : Fr.) P. Kumm. Ref.: Tortié 1968 b, unp. Torti¢ EXs7 ONE Omphalina Quél. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Clémengon 1982 a, Dennis & al. 1960, Hansen & Knudsen 1992, Kreisel 1987, Singer 1986) | Omphalina epichysium (Pers. : Fr.) Quél. = Omphalia epichysium (Pers. : Fr.) P. Kumm. (1) Ref.: Torti¢é 1966 a, Tortié 1966 b, Torti¢ 1968 b, Torti¢ 1989 b, unp. Tortié ( )(1) Exs.: CNF Omphalina griseopallida (Desm. : Fr.) Quél. = Leptoglossum griseopallidum (Desm. : Fr.) Moser (comb. inval.) = Phaeotellus griseopallidus (Desm. : Fr.) Kihner et Lamoure (comb. inval.) Ref.: Focht 1974 Oudemansiella Speg. (Lit.: Arnolds & al. 1995, Bas & al. 1999, Hansen & Knudsen 1992, Kreisel 1987, Watling & Turnbull 1998) °OQudemansiella mucida (Schrad. : Fr.) Hohn. = Armillaria mucida (Schrad. : Fr.) P. Kumm. (1) = Lepiota mucida (Schrad. : Fr.) ? (2) = Mucidula mucida (Schrad. : Fr.) Pat. (3) Ref.: Kranjéev 1979 (3), Kranjéev 1986 a, Lange 1974, Muzic & BoZac 1997, Skorié 1928 (1), Tortié 1964, Tortié 1966 a, Tortié 1966 b, Tortié 1968 a, Torti¢é 1979, Torti¢ 1989 b, Vouk & Pevalek 1915 (2), unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Torti¢é 1970, unp. Tortié ( )(3) Exs.: BEO, CNF (Lit.: Bas & al. 1999, Moser 1983) Panellus P. Karst. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Fries 1828, Fries 1836- -1838, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Pilat 1935, Pilat 1951 b, Singer 1986, Watling & Turnbull 1998) *Panellus mitis (Pers. : Fr.) Singer Ref.: Tortié 1973, Torti¢ 1989 b, unp. Tkaléec & MeSié, unp. Torti¢é Exs.: CNF, PRM (Lit.: Hansen & Knudsen 1992, Moser 1983, Watling & Turnbull 1998) 149 Panellus ringens (Fr. : Fr.) Romagn. = Lentinus ringens (Fr. : Fr.) Fr. (1) = Panus ringens (Fr. : Fr.) Fr. (2) Ref.: Stoitzner 1869 (1), Torti¢é 1968 b, unp. Tortié ( )(2) Panellus serotinus (Pers. : Fr.) Kiihner = Hohenbuehelia serotina (Pers. : Fr.) Singer (1) = Pleurotus serotinus (Pers. : Fr.) P. Kumm. (2) Ref.: Torti¢ 1966 a (1), Torti¢ 1966 b (1), Tortié 1968 a, Tortié 1979, Torti¢ 1989 b, unp. Tortié ( )(1)(2) Exs.aCNE *Panellus stipticus (Bull. : Fr.) P. Karst. = Agaricus stipticus Bull. : Fr. (1) = Lentinus stipticus (Bull. : Fr.) J. Schrot. (2) = Panus stipticus (Bull. : Fr.) Fr. (3) = Panus semipetiolatus (Schaeff.) Sacc. (4) Ref.: Barcié 1989, Baréi¢ 1996, Blagaié 1921 b (3), Jelié & Tortié 1973, Lisiewska & Torti¢ 1990, Lorinser - Janda 1877 (3), Moesz 1938 (4), Picbauer 1928 (3), Pilat 1925 (3), Schulzer 1882 a (3), Schulzer & al. 1866 (3), Skorié 1928 (2), Tortié 1964, Tortié 1966 a, Tortié 1966 b, Torti¢é 1979, Torti¢é 1989 b, Tortié 1993, Tortié 1994, Tortié & Lisiewska 1978, Vouk & Pevalek 1915 (2), unp. Matoéec, unp. Tkaléec & Me8ié, . unp. Torti¢ ( )(3), Schulzer 1857 HR? (1) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983, Watling & Turnbull 1998) °Panellus violaceofulvus (Batsch : Fr.) Singer Ref.: Tortié 1989 b, unp. Tkaléec & MeSi¢, unp. Torti¢é EXS INE (Lit.: Hansen & Knudsen 1992, Moser 1983) Pleurocybella Singer (Lit.: Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Gregory 1989) ¢Pleurocybella porrigens (Pers. : Fr.) Singer = Agaricus porrigens Pers. : Fr. (1) = Phyllotus porrigens (Pers. : Fr.) P. Karst. = Pleurotellus porrigens (Pers. : Fr.) Kithner et Romagn. (comb. inval.) (2) = Pleurotus porrigens (Pers. : Fr.) P. Kumm. (3) Ref.: Skorié 1928 (1), Tortié 1966 a, Tortié 1966 b, Tortié 1968 a (2), Tortié 1989 b, Tortié 1994, unp. Tkaléec & MeSié, unp. Tortié ( )(3) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Watling & Gregory 1989) 150 Pseudoclitocybe (Singer) Singer (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bollmann & al. 1996, Bon 1997, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Watling & Turnbull 1998) ° Pseudoclitocybe cyathiformis (Bull. : Fr.) Singer = Agaricus cyathiformis Bull. : Fr. (1) = Cantharellula cyathiformis (Bull. : Fr.) Singer (2) = Clitocybe cyathiformis (Bull. : Fr.) P. Kumm. (3) Ref.: Focht 1974 (2), Gjura3in 1898 (3), Jelié & Tortié 1973, Skori¢ 1928 (1), Tortié. 1964 (2), Torti¢ 1973, Torti¢ 1989 b, Vouk & Pevalek 1915 (1), unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Tortic ( )(3), Schulzer 1857 HR? (1) Exs.: CNF (Lit.: Moser 1983, Watling & Turnbull 1998) Pseudoclitocybe expallens (Pers. : Fr.) Moser = Agaricus expallens Pers. : Fr. (1) Ref.: Schulzer & al. 1866 (1) Resupinatus Nees (Lit.: Arnolds & al. 1995, Bas & al. 1995, Domanski 1964, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Gregory 1989) Resupinatus applicatus (Batsch : Fr.) Gray = Agaricus applicatus Batsch : Fr. (1) Ref.: Schulzer 1857 (1) EXsncGNEs, Resupinatus vetlinianus (Domanski) Moser = Pleurotus vetlinianus Domanski (1) Ref.: Torti¢ 1989 a (1), unp. Tortié (1) Exs.: CNF, PRM Rickenella Raithelh. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Clémencgon 1982 a, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983) °Rickenella fibula (Bull. : Fr.) Raithelh. = Agaricus fibula Bull. : Fr. (1) = Gerronema fibula (Bull. : Fr.) Singer (2) = Mycena fibula (Bull. : Fr.) Kiihner (3) = Omphalia fibula (Bull. : Fr.) P. Kumm. (4) = Omphalina fibula (Bull. : Fr.) Quél. Ref.: Focht 1974 (4), Jeli¢ & Torti¢é 1973 (2), Lisiewska & Torti¢ 1990, Torti¢ 1966 a (3), Torti¢é 1966 b (3), Torti¢ 1968 b (3), Tortié 1993, 151 Torti¢é & Lisiewska 1978 (2), Vouk & Pevalek 1915 (1), Vouk & Pevalek 1916 (1), unp. Matoéec, unp. Tkaléec & Me3ié, unp. Tortié (3) Exs.: GNF (Lit.: Bon 1997, Clémengon 1982 a) Ripartites P. Karst. (Lit.: Arnolds & al. 1995, Bas & al. 1995, Bon 1997, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986) *Ripartites tricholoma (Alb. et Schwein. : Fr.) P. Karst. Ref.: Focht 1974, unp. Matoéec, unp. Tkaléec & MeSic Exs-aCNE (Lit.: Bas & al. 1995, Moser 1983, Hansen & Knudsen 1992) Strobilurus Singer (Lit.: Arnolds & al. 1995, Bas & al. 1999, Dennis & al. 1960, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Turnbull 1998) °Strobilurus esculentus (Wulfen : Fr.) Singer = Pseudohiatula esculenta (Wulfen : Fr.) Singer (1) Ref.: Torti¢ 1973, unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Torti¢ ()(1) Exsero NE (Lit.: Bas & al. 1999, Moser 1983) *° Strobilurus stephanocystis (Kihner et Romagn. ex Hora) Singer Ref.: unp. Matoéec, unp. Tkaléec & MeSi¢, unp. Torti¢é Exs.: CNF (Lit.: Bas & al. 1999, Moser 1983) °Strobilurus tenacellus (Pers. : Fr.) Singer = Pseudohiatula tenacella (Pers. : Fr.) Métrod (1) Ref.: Focht 1972 (1), unp. Matoéec, unp. Tkaléec & Mesié, unp. Tortic Exs.: CNF (Lit.: Bas & al. 1999, Moser 1983) Tricholoma (Fr.) Staude (Lit.: Arnolds & al. 1995, Bas & al. 1999, Bon 1984 a, Bon 1984 b, Bon 1991, Breitenbach & Kranzlin 1991, Christensen & Noordeloos 1999, Dennis & al. 1960, Galli 1999, Hansen & Knudsen 1992, Konrad & Maublanc 1924-1937, Kreisel 1987, Kytévuori 1989, Moser 1983, Riva 1988, Singer 1986) *Tricholoma acerbum (Bull. : Fr.) Quél. Ref.: Tortié 1964, Torti¢ & Lisiewska 1978, unp. Tkaléec & Me8i¢, unp. Torti¢ 12 xsee GN (Lit.: Hansen & Knudsen 1992, Riva 1988) Tricholoma albobrunneum (Pers. : Fr.) P. Kumm. sensu lato Incl.: Tricholoma albobrunneum (Pers. : Fr.) P. Kumm. (1) Tricholoma batschii Gulden ex Mo. Chr. et Noordel. = Tricholoma subannulatum (Batsch) Bres. (comb. illeg.) (2) Tricholoma striatum (Schaeff.) Sacc. = Tricholoma albobrunneum (Pers. : Fr.) P. Kumm. var. striatum (Schaeff.) ? (3) Ref.: Baréié 1982 (1), Barcié 1996 (1), Focht 1972 (3), Focht 1974 (2), Focht 1979 (1), Torti¢ 1964 (1), unp. Tortic¢ (1) Exsi GNE Annotation: Modern authors disagree about interpretation and nomen- clature of Tricholoma albobrunneum and T. striatum (see Arnolds & al. 1995, Christensen & Noordeloos 1999, Bon 1991, Hansen & Knudsen 1992, Kreisel 1987, Riva 1988). Focht (1974) considers 7. albobru- nneum a synonym of 7. subannulatum. Tricholoma album (Schaeff. : Fr.) P. Kumm. Ref.: Baréié 1982, Barcié 1996, unp. Torti¢é Tricholoma argyraceum (Bull.) P. Kumm. = Tricholoma inocybeoides A. Pearson Ref.: Focht 1983 b Tricholoma atrosquamosum (Cheval.) Sacc. Ref.: Torti¢é 1966 a, Torti¢ 1966 b, Torti¢é 1968 b, unp. Matocec, unp. Torti¢ Exs.: CNF °Tricholoma aurantium (Schaeff. : Fr.) Ricken Ref.: Torti¢é 1966 a, Torti¢é 1966 b, unp. Tkaléec & MeSic, unp. Torti¢ Exs.: CNF (Lit.: Bas & al. 1999, Galli 1999, Hansen & Knudsen 1992) Tricholoma batschii Gulden ex Mo. Chr. et Noordel. = Tricholoma subannulatum (Batsch) Bres. (comb. illeg.) non 7. subannulatum (Peck) Zeller (1) = Tricholoma albobrunneum (Pers. : Fr.) P. Kumm. var. subannulatum (Batsch) Lucand (2) Misappl.: Tricholoma fracticum (Britzelm.) Kreisel sensu auct. non Britzelm. (3) (sensu Britzelm. = nom. dub. - Christensen & Noordeloos 1999) Ref.: Focht 1972 (2), VrSéaj 1990 (3), unp. Torti¢ (1) ExseiGNE Tricholoma bresadolanum Clémencon Ref.: Bozac 1984, Bozac 1986, Bozac 1989, Bozac 1993 Lys °Tricholoma bufonium (Pers. : Fr.) Gillet = Tricholoma sulphureum (Bull. : Fr.) P. Kumm. var. bufonium (Pers. : Fr.) Quél. (1) Ref.: Bozac 1984, Focht 1979 (1), Tortié 1966 a, Torti¢é 1966 b, unp. Tkaléec & MeSic, unp. Tortic Exs.: CNF (Lit.: Bas & al. 1999, Riva 1988) °Tricholoma caligatum (Viv.) Ricken Ref.: Torti¢é 1973, unp. Tkaléec & MeSic¢, unp. Tortic¢ Exs.: CNF (Lit.: Kyt6vuori 1989, Riva 1988) *® Tricholoma cingulatum (Almfelt : Fr.) Jacobasch Ref.: unp. Tkaléec & MeSi¢c Exs.; CNF (Lit.: Bas & al. 1999, Hansen & Knudsen 1992, Riva 1988) °Tricholoma columbetta (Fr. : Fr.) P. Kumm. Ref.: Focht 1979, Lisiewska & Tortié 1990, Torti¢ 1966 a, Torti¢ 1966 b, Torti¢é 1968 b, unp. Lisiewska, unp. Matotec, unp. Tkaléec & MeSic, unp. Torti¢ Exs.: CNF, POZM (Lit.: Hansen & Knudsen 1992, Riva 1988) * Tricholoma coryphaeum (Fr.) Gillet = Tricholoma sejunctum (Sowerby : Fr.) Quél. var. coryphaeum (Fr.) Konrad et Maubl. (1) Ref.: unp. Lisiewska (1) Exs.: POZM (1) °Tricholoma equestre (L. : Fr.) P. Kumm. = Tricholoma auratum (Paulet) Gillet (1) = Agaricus flavovirens Pers. : Fr. (2) = Tricholoma flavovirens (Pers. : Fr.) S. Lundell et Nannf. (3) Ref.: BoZac 1989, Bozac 1993, Focht 1979, Focht 1981 b (1)(3), Torti¢ 1966 a, Tortié 1966 b, Tortié 1968 b (3), unp. Matocec, unp. Tkaléec & MeSié, unp. Tortié, Schulzer 1857 HR? (2) ExsiaGNE (Lit.: Bas & al. 1999, Moser 1983, Riva 1988) *Tricholoma fulvum (DC. : Fr.) Sacc. = Tricholoma flavobrunneum (Fr.) P. Kumm. (1) = Tricholoma pseudonictitans Bon Ref.: Tortié 1966 a (1), Tortié 1966 b (1), Tortié 1968 a (1), unp. Tkaléec & MeSié, unp. Tortic¢ (1) Exs.: CNF (Lit.: Bas & al. 1999, Bon 1991, Moser 1983) 154 * Tricholoma imbricatum (Fr. : Fr.) P. Kumm. Ref.: unp. Torti¢ EXS.aGNE Tricholoma lascivum (Fr. : Fr.) Gillet = Agaricus lascivus Fr. : Fr. (1) Ref.: Torti¢é 1964, unp. Lisiewska, unp. Tortic, Vouk & Pevalek 1916 ? (1) Exs.: CNF, POZM °Tricholoma luridum (Schaeff. : Fr.) Quél. = Agaricus luridus Schaeff. : Fr. (1) Ref.: Vouk & Pevalek 1915 (1), Vouk & Pevalek 1916 (1), unp. Tkaléec & MeSic Exs.: CNF (Lit.: Bas & al. 1999, Galli 1999) Tricholoma myomyces (Pers. : Fr.) J.E. Lange = Agaricus myomyces Pers. : Fr. (1) Ref.: Schulzer & al. 1866 (1), VrSéaj 1990, Schulzer 1857 HR? (1) °Tricholoma orirubens Quél. Ref.: Torti¢ 1966 a, Torti¢ 1966 b, Torti¢ 1968 b, unp. Tkaléec & MeSicé, unp. Tortic Exs/:.,CNE (Lit.: Bon 1991, Galli 1999) °Tricholoma pardinum (Pers.) Quél. = Tricholoma pardalotum Herink et Kotl. Misappl.: Tricholoma tigrinum (Schaeff.) P. Kumm. sensu auct. non Schaeff. (1) Ref.: Focht 1979 (1), Focht 1983 b (1), Torti¢ 1966 a, Tortié 1966 b, unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Torti¢ ExStCNE (Lit.: Bas & al. 1999, Moser 1983, Riva 1988) Tricholoma populinum J.E. Lange Ref.: Vrséaj 1990, unp. Torti¢ Exo GNP ¢Tricholoma portentosum (Fr. : Fr.) Quél. Ref.: Baréié 1982, Barci¢ 1984, Barcié 1996, Focht 1979, Focht 1986, Torti¢é 1966 a, Torti¢ 1966 b, unp. Matoéec, unp. Tkaléec & MeSié, unp. Torti¢ Exs.: CNF (Lit.: Bas & al. 1999, Hansen & Knudsen 1992, Riva 1988) Tricholoma psammopus (Kalchbr.) Quél. = Agaricus psammopus Kalchbr. (1) Ref.: Focht 1972, Fries 1874 HR? (1), Saccardo 1887 HR? 1D ¢Tricholoma saponaceum (Fr. : Fr.) P. Kumm. = Agaricus saponaceus Fr. : Fr. (1) Ref.: Lisiewska & Torti¢ 1990, Schulzer & al. 1866 (1), Tortié 1966 a, Torti¢ 1966 b, Torti¢ 1968 b, Tortié & Lisiewska 1978, unp. Lisiewska, unp. Matoéec, unp. Torti¢é Exs.: CNF, POZM *var. Saponaceum Ref.: unp. Tkaléec & MeSié (Lit.: Bas & al. 1999, Bon 1991, Riva 1988) *evar. boudieri Bigeard et H. Guill. Misappl.: Tricholoma saponaceum (Fr. : Fr.) P. Kumm. var. lavedanum Rolland sensu auct. non Rolland Ref.: unp. Tkaléec & MeSi¢é Exs. CNF (Lit.: Bas & al. 1999, Bon 1991, Riva 1988) *®var. squamosum (Cooke) Rea Ref.: unp. Tkaléec & MeSsi¢é xs’ GNP (Lit.: Bas & al. 1999, Galli 1999) °Tricholoma scalpturatum (Fr.) Quél. Ref.: Focht 1972, Focht 1979, Torti¢ 1964, Tortié & Lisiewska 1978, unp. Tkaléec & MeSi¢, unp. Torti¢, Focht 1986 HR? ExsiGNE (Lit.: Bas & al. 1999, Bon 1991, Moser 1983) °Tricholoma sciodes (Pers.) C. Martin Ref.: Tortié 1968 b, Tortié & Lisiewska 1978, unp. Tkaléec & MeSic, unp. Tortic Exs::-CNF (Lit.: Bas & al. 1999, Riva 1988) °Tricholoma sejunctum (Sowerby : Fr.) Quél. = Tricholoma viridilutescens Moser Ref.: KiSpatié 1950, MuZic & Bozac 1984, Muzic & Bozac 1997, Torti¢ 1964, Tortié & Lisiewska 1978, unp. Matocéec, unp. Tkaléec & Me8i¢, unp. Torti¢ Exs:4GNF (Lit.: Bas & al. 1999, Bon 1991, Galli 1999) °Tricholoma squarrulosum Bres. Ref.: unp. Tkaléec & MeSié Exs; ONE (Lit.: Bon 1991, Galli 1999, Hansen & Knudsen 1992) Tricholoma sudum (Fr.) Queél. Ref.: GjuraSin 1898 156 °Tricholoma sulphureum (Bull. : Fr.) P. Kumm. = Agaricus sulphureus Bull. : Fr. (1) Ref.: Baréi¢é 1982, Baréié 1996, Focht 1979, Lisiewska & Torti¢ 1990, Schulzer & al. 1866 (1), Tortié 1964, Tortié 1966 a, Torti€é 1966 b, Torti¢é & Lisiewska 1978, unp. Matoéec, unp. Torti¢ Exs.: BEO, CNF °var. sulphureum Ref.: unp. Tkaléec & Mesi¢ Exs,;GNE (Lit.: Bon 1991) *®var. coronarium (Pers.) Niiesch Ref.: unp. Tkaléec & MeSié Exs.: CNF (Lit.: Bon 1991) Tricholoma terreum (Schaeff. : Fr.) P. Kumm. = Agaricus terreus Schaeff. : Fr. (1) Ref.: Baréié 1982, Baréié 1984, Barcic 1996, Bozac 1989, Bozac 1993, Focht 1972, Focht 1979, Focht 1981 b, Focht 1983 b, Focht 1986, Muzic & Bozac 1984, Muzic & Bozac 1997, Skorié 1928 (1), Tortié 1964, Torti¢ 1966 a, Vouk & Pevalek 1916 (1), unp. Matoéec, unp. Torti¢ Exs.: CNF Tricholoma ustale (Fr. : Fr.) P. Kumm. Ref.: Torti¢é & Lisiewska 1978, unp. Lisiewska, unp. Torti¢é Exs.: POZM *® Tricholoma vaccinum (Schaeff. : Fr.) P. Kumm. Ref.: unp. Matoéec, unp. Tkaléec & Me8ic, unp. Torti¢ Exss CNF (Lit.: Bas & al. 1999, Galli 1999, Riva 1988) Tricholomopsis Singer (Lit.: Arnolds & al. 1995, Bas & al. 1999, Bon 1991, Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Pilat _ 1951 b, Singer 1986) *Tricholomopsis decora (Fr. : Fr.) Singer = Pleurotus decorus (Fr. : Fr.) Sacc. (1) Ref.: Lisiewska & Tortié 1990, Tortié 1966 a, Torti¢ 1966 b, Tortié 1968 a, Torti¢ 1989 b, unp. Lisiewska, unp. Tkaléec & MeSi¢, unp. Torti¢ ( )(1) Exs.: CNF, POZM (Lit.: Bon 1991, Hansen & Knudsen 1992, Moser 1983) °Tricholomopsis rutilans (Schaeff. : Fr.) Singer = Pleurotus rutilans (Schaeff. : Fr.) Pilat (1) = Tricholoma rutilans (Schaeff. : Fr.) P. Kumm. (2) ibe Ref.: Kranjéev 1995, Lisiewska & Tortié 1990, Tortié 1964, Tortié 1966 a, Torti¢ 1966 b, Tortic 1989 b, Torti¢ 1994, unp. Matoéec, unp. Tkaléec & MeSic¢, unp. Tortic ( )(1)(2) Exs:: CNE (Lit.: Bon 1991, Moser 1983) Xeromphalia Kihner et Maire ("Xeromphalina") (Lit.: Hansen & Knudsen 1992, Kreisel 1987, Moser 1983, Singer 1986, Watling & Turnbull 1998) ¢Xeromphalia campanella (Batsch : Fr.) Kiihner et Maire Ref.: Lisiewska & Torti¢ 1990, Tortié 1966 a, Tortié 1966 b, Tortié 1989 b, unp. Tkaléec & Me8ié, unp. Torti¢é Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983) Xeromphalia caulicinalis (Fr.) Kthner et Maire = Marasmius caulicinalis Fr. (1) Ref.: Vouk & Pevalek 1916 (1) Xerula Maire (Lit.: Arnolds & al. 1995, Bas & al. 1999, Bollmann & al. 1996, Breitenbach & Kranzlin 1991, Fries 1821, Hansen & Knudsen 1992, Kreisel 1987, Maas Geesteranus 1992 a, Moser 1983, Watling & Turnbull 1998) *® Xerula melanotricha Dorfelt = Oudemansiella melanotricha (Dorfelt) Moser Ref.: unp. Tkaléec & Mesié Exs.: CNF (Lit.: Breitenbach & Kranzlin 1991, Moser 1983) *Xerula pudens (Pers.) Singer = Agaricus longipes Bull. (nom. illeg.) (1) = Collybia longipes (Bull. —) Quél. (2) = Marasmius longipes (Bull. > Quél.) Quél. (3) = Oudemansiella longipes (Bull. + Quél.) Moser (4) = Xerula longipes (Bull. + Quél.) Maire Misappl.: Oudemansiella badia (Quél.) Moser (comb. inval.) sensu Moser non Quél. (5) Ref.: Blagaié 1921 b (2), GjuraSin 1898 (2), KiSpati¢é 1950 (2), Lisiewska & Tortié 1990 (4), Schulzer 1882 a (1), Schulzer & al. 1866 (1), Torti¢ 1964 (4), Tortié 1966 a (5), Tortié 1966 b (5), Vouk & Pevalek 1915 (1), unp. Lisiewska (5), unp. Matoéec (4), unp. Tkaléec & Me8i¢, unp. Torti¢ (3) Exs.: CNF, POZM (5) (Lit.: Bas & al. 1999, Breitenbach & Kranzlin 1991, Moser 1983) 158 Xerula pudens (Pers.) Singer sensu lato = Oudemansiella longipes (Bull. + Quél.) Moser sensu lato (1) Ref.: Torti¢é 1993, unp. Torti¢ (1) °Xerula radicata (Relhan : Fr.) Dérfelt = Agaricus radicatus Relhan : Fr. (1) = Collybia radicata (Relhan : Fr.) Quél. (2) = Oudemansiella radicata (Relhan : Fr.) Singer (3) Agaricus macrourus Scop. (4) Collybia macroura (Scop.) ? (5) = Agaricus sudoroides Schulzer (6) Ref.: Baréié 1982 (3), Baréié 1996 (3), Blagai¢é 1921 b (5), Bozac 1984 (3), Jeli¢é & Tortié 1973 (3), KiSpati¢ 1950 (2), Lisiewska & Tortié 1990 (3), Schulzer 1880 a (6), Schulzer & al. 1866 (1), Skorié 1928 (4), Tortié 1964 (3), Tortié 1966 a (3), Torti¢é 1966 b (3), Tortié 1989 b (3), Torti¢é 1993, Torti¢é & Lisiewska 1978 (3), Vouk & Pevalek 1915 (4), Vouk & Pevalek 1916 (4), unp. Matoéec (3), unp. Tkaléec & MeS8i¢, unp. Torti¢ (2)(3), Schulzer 1857 HR? (1) Exs.: CNF (Lit.: Hansen & Knudsen 1992, Moser 1983, Watling & Turnbull 1998) EXCLUDED RECORDS Ordo Agaricales (family unknown to us) Agaricus agrorum Schulzer - nom. nud. = Agaricus agricolus Schulzer (nom. nud.) (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Agaricus agrorum Schulzer var. coloratus Schulzer - nom. nud. = Agaricus agricolus Schulzer var. coloratus Schulzer (nom. nud.) (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Agaricus albicolor Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus amethysteoroseus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus anceps Schulzer = ? [Naucoria] Ref.: Schulzer 1877 Agaricus aureus Sowerby = ? Ref.: Schulzer 1857 HR? Agaricus caesius Pers. = ? Ref.: Schulzer & al. 1866 Agaricus candidatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus capitatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 159 Agaricus carneolamellatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus collabescens Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus collapsus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus collapsus Schulzer var. crateriformis Schulzer - nom. nud. Ref.: Schulzer 1867 HR? Agaricus congregatus Rabenh. = ? Ref.: Schulzer 1857 HR? Agaricus cylindripes Schulzer - nom. nud. Ref.: Schulzer 1867 Agaricus disciformis Wettst. = ? Ref.: Skori¢ 1928 Agaricus equinus Pers. = ? Ref.: Schulzer 1857 HR? Agaricus eumorphus Pers. = ? Ref.: Schulzer & al. 1866 ? Agaricus fertilis Pers. : Fr. = ? Ref.: Schulzer & al. 1866 Agaricus flavocinereus Pers. = ? Ref.: Schulzer 1857 HR? Agaricus herpeticus Roques = ? Ref.: Schulzer & al. 1866 Agaricus hiemali-roraceus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus horticolus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus horticolus Schulzer var. glabropilus Schulzer - nom. nud. Ref.: Schulzer 1867 HR? Agaricus ilkae Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus incurvus Pers. Ref.: Schulzer & al. 1866 Agaricus indigestus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus indigestus Schulzer var. juglandis Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus jucundus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus macer Schulzer - nom. nud. Ref.: Schulzer 1867 160 Agaricus mastoideus Krombh. = ? Ref.: Schulzer & al. 1866, Schulzer 1857 HR? Agaricus multicolor Pers. : Fr. = ? Ref.: Schulzer & al. 1866 ? Agaricus nitens Schaeff. = ? Ref.: Schulzer & al. 1866, Schulzer 1857 HR? Agaricus nitens Schaeff. var. albus (Schaeff.) ? = ? Ref.: Schulzer & al. 1866 Agaricus obscurans Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus ocellus Schulzer = ? [Pilosace] Ref.: Schulzer 1877 Agaricus pannosus Fr. : Fr. var. pascuus Schulzer = ? [Naucoria] Ref.: Schulzer 1877 Agaricus pannosus Fr. : Fr. var. rufisporus Schulzer = ? [Naucoria] Ref.: Schulzer 1877 , Agaricus pratorum Schulzer = ? [Naucoria] Ref.: Schulzer 1878 Agaricus pygmaeoides Schulzer = ? [Naucoria] Ref.: Schulzer 1877 Agaricus roseolamellatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus segestrius Fr. : Fr. = ? Ref.: Schulzer & al. 1866 ? Agaricus semiglobosus Schulzer = ? [Naucoria] Ref.: Schulzer 1879 Agaricus seriatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus spadochrous Krombh. = ? Ref.: Schulzer 1857 HR? Agaricus subfissus Schulzer - nom. nud. Ref.: Schulzer 1867 HR? Agaricus subradicatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus subrugatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus subsalsus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus venustus Viv. = ? Ref.: Schulzer & al. 1866, Schulzer 1857 HR? Agaricus vexans Schulzer var. robustior Schulzer = ? [Naucoria] Ref.: Schulzer 1881 161 Agaricus victorinae Schulzer - nom. nud. Ref.: Schulzer 1867 Agaricus vindobonensis Tratt. = ? Ref.: Schulzer 1883 b HR? Agaricus violaceus Pers. = ? Ref.: Schulzer 1857 HR? Pilosace bresadolae Schulzer = ? Ref.: Saccardo 1887, Schulzer 1885 Familia Pleurotaceae Overeem ex Kiihner et Romagn. Agaricus farinodorus Schulzer = ? [Pleurotus] Ref.: Schulzer 1880 a Agaricus squamatus Schulzer = ? [Pleurotus] Ref.: Schulzer 1882 b Lentinus degener Kalchbr. var. elatus Schulzer = ? Ref.: Schulzer 1879 Lentinus degener Kalchbr. var. tomentosus Schulzer = ? Ref.: Schulzer 1879 Lentinus grateolens Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Lentinus leontopodius Schulzer var. insociabilis Schulzer = ? Ref.: Schulzer 1879 Lentinus leontopodius Schulzer var. silvaticus Schulzer = ? Ref.: Schulzer 1879 Panus cibaliensis Schulzer = ? Ref.: Schulzer 1879 Panus salignus (Pers. : Fr.) ? var. anciophyllos Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Panus solitarius Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Panus stipticoides Schulzer = ? Ref.: Schulzer 1879 Panus subinteger Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Familia Tricholomataceae R. Heim ex Pouzar Agaricus alboflavus Schulzer = ? [Clitocybe] Ref.: Schulzer 1876 Agaricus alcalinus Fr. : Fr. - nom. dub. (Maas Geesteranus 1992 a) Ref.: Stoitzner 1869, Vouk & Pevalek 1916 162 Agaricus amadelphoides Schulzer = ? [Collybia] Ref.: Schulzer 1881 Agaricus amaricans Schulzer = ? [Collybia] Ref.: Schulzer 1880 a, Schulzer & al. 1866 Agaricus amicoides Schulzer = ? [Tricholoma] Ref.: Schulzer 1880 a Agaricus androsaceus "Pers." = ? Ref.: Schulzer 1857 HR? Agaricus bellus Pers. : Fr. = ? [Clitocybe] Ref.: Vouk & Pevalek 1916 Agaricus brumalis Fr. : Fr. var. slavonicus Schulz. = ? Ref.: Schulzer 1878 Agaricus candiculus Schulzer = ? [Collybia] Ref.: Schulzer 1878, Schulzer & al. 1866 (nom. nud.) Agaricus candiculus Schulzer var. hortensis Schulzer - nom. nud. Ref.: Schulzer & al. 1866 / Agaricus contortus Bull. : Fr. - nom. dub. (Antonin & Noordeloos 1997) Ref.: Schulzer & al. 1866 Agaricus corticola Pers. : Fr. - nom. ambig. (Maas Geesteranus 1992 a) Ref.: Vouk & Pevalek 1915, Schulzer 1857 HR? Agaricus derodus Schulzer = ? [Armillaria] Ref.: Schulzer 1878 Agaricus deumbonatus Schulzer = ? [Tricholoma] Ref.: Schulzer 1881 Agaricus diaphanus Schulzer - nom. dub. (Maas Geesteranus 1992 a) = Mycena nivea Quél. et Schulzer (nom. illeg.) (1) Ref.: Schulzer 1879, Schulzer 1885 (1), Saccardo 1887 HR? (1) Agaricus electus Schulzer = ? [Collybia] Ref.: Schulzer 1878 Agaricus electus Schulzer var. autumnalis Schulzer = ? [Collybia] Ref.: Schulzer 1879 Agaricus electus Schulzer var. subumbonatus Schulzer = ? [Collybia] Ref.: Schulzer 1878 Agaricus elevatus Weinm. - nom. dub. (Antonin & Noordeloos 1997) Ref.: Vouk & Pevalek 1916 Agaricus erythropus Pers. : Fr. var. slavonica Schulzer = ? (Collybia) Ref.: Schulzer 1880 a Agaricus galericulatus Scop. : Fr. var. "b. amoene albus" Fr. 1821 =? Ref.: Schulzer 1857 HR? Agaricus gracillimus Weinm. Ref.: Vouk & Pevalek 1916 163 Annotation: According to Dennis & al. (1960) this species is doubtful in the original, Weinmann's sense. Bresadola and J. E. Lange have misapplied this name to Hemimycena delectabilis (Peck) Singer (Bollmann & al. 1996, Dennis & al. 1960). We do not know in which sense the author of the record has used this name. Agaricus humicola Schulzer = ? [Clitocybe] Ref.: Schulzer 1878 Agaricus insignis Schulzer = ? [Clitocybe] = Hygrophorus insignis Schulzer (1) Ref.: Schulzer 1876 (1), Schulzer 1880 a Agaricus intornatus Schulzer = ? [Clitocybe] Ref.: Schulzer 1880 a Agaricus lineatus Bull. : Fr. - nom. dub. (Maas Geesteranus 1992 a) Ref.: Vouk & Pevalek 1915 Agaricus longipes Bull. var. inornatus Schulzer = ? Ref.: Schulzer 1878 Agaricus melleus Vahl : Fr. var. sciacephalus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus obliquus Pers. : Fr. var. slavonicus Schulzer = ? [Clitocybe]| Ref.: Schulzer 1878 Agaricus oniscoides Schulzer = ? [Omphalia] Ref.: Schulzer 1878 Agaricus personatus Fr. : Fr. Ref.: Schulzer & al. 1866 Annotation: Agaricus personatus is a synonym of Lepista nuda (Bull. : Fr.) Cooke (Dennis & al. 1960). However, since many authors have misapplied this epithet for L. saeva (Fr.) P.D. Orton, we do not know in which sense the author of the record has used this name. Agaricus platyphylloides Schulzer = ? [Collybia] Ref.: Schulzer 1878 Agaricus platyphyllus Pers. : Fr. var. rhizomatus Schulzer = ? Ref.: Schulzer & al. 1866 Agaricus platyrhizus Schulzer = ? [Collybia] = Tricholoma fallaciosum Sacc. (nom. illeg.) (1) = Tricholoma fallax Quél. et Schulzer (nom. illeg.) (2) Ref.: Saccardo 1887 (1), Schulzer 1878, Schulzer 1885 (2) Agaricus radicatus Relhan : Fr. var. luteolamellatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus radiculiferus Schulzer = ? [Collybia] Ref.: Schulzer 1881 Agaricus sejunctus Sowerby : Fr. var. sapidus Schulzer = ? Ref.: Schulzer 1878 164 Agaricus sulphureus Bull. : Fr. var. inflatipes Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Agaricus tardissimus Schulzer = ? [Omphalia] Ref.: Schulzer 1867 (nom. nud.), Schulzer 1878 Agaricus tardissimus Schulzer var. aciphilus Schulzer - nom. nud. Ref.: Schulzer 1867 Agaricus tumidoideus Schulzer = ? [Tricholoma] Ref.: Schulzer 1879 Agaricus viticola Schulzer - nom. dub. (Maas Geesteranus 1992 a) = Mycena bresadolae Schulzer (nom. illeg.) (1) Ref.: Schulzer 1867 (nom. nud.), Schulzer 1878, Schulzer 1885 (1), Saccardo 1887 HR? Calocybe graveolens (Pers. : Fr.) Singer = Agaricus graveolens Pers. : Fr. (1) = Tricholoma georgii (Clus. ex L.) Quél. var. graveolens (Pers. : Fr.) ? (2) Ref.: Blagaié 1923 (2), Vouk & Pevalek 1916 (1) Annotation: Singer (1977, 1986) considers it an autonomous species, while Dennis & al. (1960) and Kreisel (1987) regard it a synonym of Calocybe gambosa (Fr. : Fr.) Singer ex Donk. If these two species are conspecific, the epithet graveolens has priority. Chaetocalathus craterellus (Durieu et Lév.) Singer Ref.: Antonin & Noordeloos 1997 Exs,7h Annotation: The author of the record specified that the locality (Zelenika near Castelnuovo) belongs to Croatia, but presently the locality belongs to Montenegro (Yugoslavia). Clitocybe bresadolae Schulzer = ? Ref.: Saccardo 1887, Schulzer 1885 Clitocybe brumalis (Fr. : Fr.) P. Kumm. Ref.: Focht 1972 Annotation: This species is interpreted in a few different senses (see Clémengon 1984 a, Kuyper 1996). We do not know in which sense the author of the record has used this name. Clitocybe dealbata (Sowerby : Fr.) P. Kumm. - nom. dub. (Bas & al. 1995) = Agaricus dealbatus Sowerby : Fr. (1) Ref.: Baréi¢é 1982, Baréié 1996, Bozac 1982, Focht 1986, Vouk & Pevalek 1915 (1) Annotation: According to Arnolds & al. (1995) and Bas & al. (1995) this epithet has been misapplied for Clitocybe rivulosa (Pers. : Fr.) P. Kumm. and C. augeana (Mont.) Sacc. Since we do not know in which sense the author of the record has used this name, we exclude it. Clitocybe jandae Schulzer = ? Ref.: Schulzer 1887 165 Clitocybe laccata (Scop. : Fr.) P. Kumm. var. rufocarnea (Fr.) ? = ? Ref.: Blagai¢ 1921 b Clitocybe maxima (G§artn. et G. Mey.) P. Kumm. Ref.: unp. Matoéec, unp. Torti¢ Exs.: CNF Annotation: Bas & al. (1995) and Kuyper (1996) consider it a nomen dubium, and that records of this species in fact represent Clitocybe geotropa (DC. et Lam.) Quél., C. gigas Harmaja or Leucopaxillus giganteus (Sibth. : Fr.) Singer. Without revision it is not possible to know which species the authors of the records for Croatia have found. Collybia accommodans (Schulzer) Sacc. = Agaricus accommodans Schulzer (1) Ref.: Saccardo 1887, Schulzer 1882 a (1) Annotation: Antonin & Noordeloos (1997) consider that it could well be identical with Micromphale brassicolens (Romagn.) P.D. Orton. Collybia dryophila (Bull. : Fr.) P. Kumm. var. minor Bres. = Agaricus radiculans Schulzer (1) Ref.: Saccardo 1887, Schulzer 1878 (1), Schulzer 1885 Annotation: Antonin & Noordeloos (1997) consider that it could be a form of Collybia dryophila (Bull. : Fr.) P. Kumm. Collybia dryophila (Bull. : Fr.) P. Kumm. var. peronata Bres. - nom. dub. (Antonin & Noordeloos 1997) = Agaricus peronatoides Schulzer (1) Ref.: Schulzer 1880 a (1), Schulzer 1885, Saccardo 1887 HR? Collybia dryophila (Bull. : Fr.) P. Kumm. var. vernalis (Schulzer) Schulzer et Bres. = Agaricus vernalis Schulzer (1) Ref.: Schulzer 1880 a (1), Schulzer 1885, Saccardo 1887 HR? Annotation: Antonin & Noordeloos (1997) consider it a nomen dubium; it could represent Collybia nivalis (Luthi et Plomb) Moser. Collybia rigidipes (Schulzer) Sacc. - nom. dub. (Antonin & Noordeloos 1997) = Agaricus rigidipes Schulzer (1) Ref.: Saccardo 1887, Schulzer 1882 a (1) Collybia spuria (Bres.) Sacc. = ? = Collybia platyphylla (Pers. : Fr.) P. Kumm. var. spuria Bres. (1) Ref.: Saccardo 1887, Schulzer 1885 (1) Cyphella erosa Schulzer = ? Ref.: Schulzer 1874 Marasmius aggericola Schulzer - nom. nud. = Marasmius depressus Schulzer (nom. nud.) (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Marasmius amoenipes Schulzer - nom. nud. Ref.: Schulzer & al. 1866 166 Marasmius angulatus (Batsch) Berk. et Broome Ref.: Vouk & Pevalek 1915 Annotation: -Batsch's epithet angulatus refers to Psilocybe angulata (Batsch) Singer (Singer 1986). The combination Marasmius angulatus suggests mis- interpretation of this species, so we do not know which species the authors of the record for Croatia have found. Marasmius auripes Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius avellanae Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius brusinae Schulzer - nom. dub. (Antonin & Noordeloos 1993) Ref.: Saccardo 1887, Schulzer 1886 Marasmius calopus (Pers. : Fr.) Fr. - nom. dub. (Antonin & Noordeloos 1993) Ref.: Schulzer & al. 1866 Marasmius catharinae Schulzer - nom. nud. = Marasmius stellatus Schulzer (nom. nud.) (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Marasmius corticola (Pers. : Fr.) Schulzer var. albidus (Fr. : Fr.) Schulzer - nom. ambig. (Maas Geesteranus 1992 a) Ref.: Schulzer & al. 1866 Marasmius corticola (Pers. : Fr.) Schulzer var. fuscescens (Fr. : Fr.) Schulzer - nom. ambig. (Maas Geesteranus 1992 a) Ref.: Schulzer & al. 1866 Marasmius corticola (Pers. : Fr.) Schulzer var. filipes Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius declinatus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 + Schulzer 1867 Marasmius filopes (Bull. : Fr.) Schulzer var. marcescens Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius galericulatus (Scop. : Fr.) Schulzer var. transiens Schulzer - nom. nud. Ref.: Schulzer 1867 HR? Marasmius leucoumbrinus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius molyoides Fr. - nom. dub. (Antonin & Noordeloos 1993) Ref.: Vouk & Pevalek 1916 Marasmius olens Schulzer - nom. nud. = Marasmius teneripes Schulzer (nom. nud.) (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Marasmius peronatus (Bolton : Fr.) Fr. var. rugulosus (Schulzer) Schulzer et Bres. - nom. dub. (Antonin & Noordeloos 1997) = Agaricus peronatus Bolton : Fr. var. rugulosus Schulzer (1) Ref.: Schulzer 1880 a (1), Schulzer 1885, Saccardo 1887 HR? 167 Marasmius praticolus Schulzer - nom. nud. Ref.: Schulzer & al. 1866 Marasmius queletii Schulzer = ? = Agaricus oreades Bolton : Fr. var. albidus Schulzer (1) Ref.: Schulzer 1879 (1), Schulzer 1885, Saccardo 1887 HR? Marasmius schulzeri Quél. = ? = Agaricus androsaceus "Pers." var. truncicola Schulzer (1) Ref.: Schulzer 1880 a (1), Schulzer 1885, Saccardo 1887 HR? Marasmius tinctipes Schulzer - nom. nud. - as Marasmius androsaceus (L. : Fr.) Fr. (1) Ref.: Schulzer & al. 1866 (1) + Schulzer 1867 Marasmius vukotinovicianus Schulzer Ref.: Schulzer 1887 Annotation: Antonin & Noordeloos (1997) consider it a nomen dubium; it could be identical with Marasmius cohaerens (Pers. : Fr.) Cooke et Quél. Melanoleuca subbrevipes Métrod ex Bon Ref.: Bozac 1984 Annotation: Recent authors interpret this species in different ways. Some authors consider it a synonym of Melanoleuca grammopodia (Bull. : Fr.) Pat. (Arnolds & al. 1995, Bas & al. 1999, Kreisel 1987, Watling & Turnbull 1998), while others consider it an autonomous species (Bon 1991, Hansen & Knudsen 1992, Singer 1986). Mycena arborea (Schulzer) Sacc. - nom. dub. (Maas Geesteranus 1992 a) = Agaricus arboreus Schulzer (1) Ref.: Saccardo 1887, Schulzer 1882 a (1) Mycena corticola (Pers. : Fr.) Gray - nom. ambig. (Maas Geesteranus 1992 a) Ref.: Jaap 1916, unp. Tortic Exs 7 CNE Mycena debilis (Fr.) Quél. - nom. dub. (Maas Geesteranus 1992 a) Ref.: Jaap 1916 Mycena stannea (Fr.) Quél. - nom. dub. (Maas Geesteranus 1992 a) Ref.: GjuraSin 1898 Mycena tintinnabulum (Fr.) Quél. Ref.: Torti¢ 1968 b ? Exs 3GNE Annotation: The author himself doubted his identification. Mycena viscidolutea (Schulzer) Sacc. - nom. dub. (Maas Geesteranus 1992 a) = Agaricus viscidoluteus Schulzer (1) Ref.: Saccardo 1887, Schulzer 1882 a (1) Omphalia scyphiformis (Fr.) Gillet Ref.: GjuraSin 1898 Annotation: Dennis & al. (1960) consider it an autonomous species (Omphalina scyphiformis (Fr.) Quél.), but according to them a modern 168 description is needed. We exclude the record since the species has not been mentioned in more recent literature available to us. Panus craterellus (Durieu et Lév.) ? Ref.: Kalchbrenner & Schulzer 1873-1877 HR? Annotation: According to Pilat (1935), Kalchbrenner & Schulzer (1873- -1877) misapplied this epithet, probably for Pleurotus (today Panellus) violaceofulvus (Batsch : Fr.) Pilat f. delastrei (Mont.) Malk. Paxillus laetipes Schulzer = ? [Lepista] Ref.: Schulzer 1876 Paxillus laetipes Schulzer var. coprophilus Schulzer = ? [Lepista] Ref.: Schulzer 1879 Pseudohiatula conigena (Pers. : Fr.) Moser - nom. ambig. (Dennis & al. 1960, Singer 1986, Watling & Turnbull 1998) Ref.: Focht 1972 Tricholoma amethystinum "Fr." = ? Ref.: GjuraSin 1898 Tricholoma bresadolae Schulzer = ? Ref.: Saccardo 1887, Schulzer 1885 Tricholoma resplendens (Fr.) Quél. Ref.: GjuraSin 1898 Annotation: This species is interpreted in a few different senses (see Bon 1984 b, Dennis & al. 1960, Kreisel 1987). We do not know in which sense the author of the record has used this name. ACKNOWLEDGEMENTS We are grateful to Dr. Milica Torti¢, Prof. Maria Lisiewska and Mr. Neven Mato¢éec for generously allowing us to use the unpublished data from their records; to Dr. Milica Torti¢, Mr. Andrej Piltaver, Prof. Bogdan Cvjetkovi¢ and Mr. Gabriele Cacialli for their assistance with literature; to Prof. Roy Watling and Prof. Wulfard Winterhoff for reviewing the manu- script; to Mrs. Jadranka USnik for helping with the English version of the text; and to the former State Directorate for the Protection of Nature and Environment for financial assistance. We also thank all collectors of mate- rial and all those who have in any way contributed to the writing of this paper. 169 LITERATURE CITED SOURCES OF RECORDED TAXA Published: Antonin, V. & M. E. Noordeloos (1993). 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MYCOTAXON Volume LXXXI, pp. 177-184 January-March 2002 THAXTEROGASTER IS A TAXONOMIC SYNONYM OF CORTINARIUS: NEW COMBINATIONS AND NEW NAMES Ursula Peintner'*, Meinhard Moser? and Rytas Vilgalys' "Institute of Biology, Duke University, Durham, NC 27709, USA “Institute of Microbiology, University of Innsbruck, Technikerstr. 25, 6020 Innsbruck, Austria ursula. peintner@uibk. ac. at Abstract Results of our earlier molecular phylogenetic studies using sequences of the rDNA internal transcribed spacers (ITS) showed that the genus Thaxterogaster is polyphyletic, and nested within Cortinarius: the secotioid Thaxterogaster spp. have multiple origins within the subgenera Myxacium, Phlegmacium and Telamonia. |In order to acknowledge these phylogenetic relationships, and the given morphological transitions, we propose to synonymize Thaxterogaster with Cortinarius. For the 54 currently recognized species of Thaxterogaster as well as for 2 varieties, 41 new combinations and 15 new names are proposed. Key words: Sequestrate Agaricales, Cortinariaceae, nomenclature. Introduction The genus Thaxterogaster Singer (1951) had been erected to accommodate taxa with Cortinarius-like spores, but with basidiomes characterized by a reduced stipe and a basally exposed hymenium with a loculate to lamellate gleba (secotioid basidiomes). The type species is Thaxterogaster magellanicum Singer. In the following years, many new Thaxterogaster spp. have been described (Singer, 1960; Singer and Smith, 1958, 1963; Moser 1964; Horak and Moser, 1965; Thiers and Smith, 1969; Horak, 1973a, 1979; Halling, 1981; Beaton et al., 1985; Calhoun, 1989; Bougher, 1997; Soop, 1998; Trappe and Castellano, 2000). However, already Moser (1964) as well as Horak and Moser (1965) noticed that Thaxterogaster and Cortinarius (Pers.) S.F. Gray are not only very closely related, but that transitions between the respective basidiome morphologies occur, sometimes even between basidiomes of one collection. They regarded Thaxterogaster as polyphyletic because of the existence of links between species, or groups of species of /Jhaxterogaster and different subgenera of Cortinarius. Also the distinctive character of Thaxterogaster spp., 178 namely “lack of active spore discharge’, was observed to be highly variable: the axial spore symmetry can be completely lacking in several Thaxterogaster species. Thus, the delimitation of Thaxterogaster and Cortinarius was mainly based on the presence of a loculate hymenium, and transitions between Cortinarius and Thaxterogaster are obvious (Horak and Moser, 1965). Therefore, Horak and Moser (1965) conclude that if Cortinarius is monophyletic, Thaxterogaster is probably polyphyletic. _ Although the affinities between Cortinarius and Thaxterogaster were soon recognized, the direction of morphological evolution has long been debated (Smith, 1971; Singer, 1975; Thiers, 1984). However, recent molecular phylogenetic studies (Peintner et al., 2001), using sequences of the rDNA internal transcribed spacers (ITS) showed that the reduced basidiome types of Thaxterogaster spp. derived several times from agaric-like Cortinarius precursors. Moreover, the genus Jhaxferogaster is polyphyletic and nested within the Cortinarius clade: species currently classified in the genus Thaxterogaster have multiple origins within the subgenera Myxacium, Phlegmacium and Telamonia. |n order to acknowledge these phylogenetic relationships, and the given morphological transitions between these genera, we propose to synonymize Thaxterogaster with Cortinarius. Regarding the nomenclature of Thaxterogaster, controversy exists about the gender of this genus name: specific epithets were applied in the neuter gender (-um) in most basionyms, but later corrected to masculine (-us). Currently, most generic names derived from “gaster’ (e.g. Thaxterogaster, Phallogaster, Hymenogaster) are used as masculine names. However, the gender of the Greek word “gaster’ (stomach) is feminine. Thus, following the St. Louis code of botanical nomenclature (Greuter et. al, 2000), all generic names derived from “gaster’ should be regarded as feminine. Because epithets should agree grammatically with the generic name, they are to be corrected (Art 23.5, or 32.5, or 60.1). No such corrections have been made in this paper. Correct names, new names and proposed combinations Cortinarius albocanus (E. Horak & M.M. Moser) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster albocanum E. Horak & M.M. Moser 1965, Nova Hedwigia X, 234; ut Thaxterogaster albocanus. Cortinarius anisodorus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster anisodorum E. Horak 1973, Beih. Nova Hedwigia 43:107; ut Thaxterogaster anisodorus. Cortinarius archeuretus (Halling) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster archeuretus Halling 1981, Mycologia 73: 861. Cortinarius basipurpureus (Bougher) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster basipurpureum Bougher 1997, Mycotaxon 63:43-45. Cortinarius brachyspermus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster brevisporum Singer 1960, Persoonia 1:386. non Cortinarius brevisporus M.M. Moser 1975. Cortinarius cabrinii (Raithelh.) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster cabrinii Raithelh. 1983, Metrodiana, Sonderheft 2:20. ee %. ial 179 Cortinarius campbellae (Berk. & Broome ex Zeller & C. W. Dodge) Peintner & M.M. Moser comb. nov. Basionym: Arcangeliella campbellae Berk. & Broome ex Zeller & C. W. Dodge 1935, Ann. Missouri. Bot. Garden 22:366. Synonym: Thaxterogaster campbellae (Zeller & C. W. Dodge) G.W. Beaton, Pegler & T. W. K. Young 1985, Kew Bulletin 40: 173. Hymenogaster campbelli Berkeley & Broome, herb. nom. Cortinarius carneoroseus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster carneoroseum E. Horak 1965, Nova Hedwigia X, 229; ut Thaxterogaster carneoroseus. Cortinarius cartilagineus (G. Cunn.) Peintner & M.M. Moser comb. nov. Basionym: Secotium cartilagineum G. Cunn. 1924, Proc. Linn. Soc. New South Wales 49:115. Synonym: Thaxterogaster cartilagineus (G. Cunn.) M.M. Moser 1965, in E. Horak & M.M. Moser 1965, Nova Hedwigia X, 228. Cortinarius conei (R. Heim) Peintner & M.M. Moser comb. nov. Basionym: Secotium conei R. Heim 1951, Rev. de Myc. 16: 142. Synonym: Thaxterogaster conei (R. Heim) E. Horak & M.M. Moser 1965, Nova Hedwigia X: 239. Cortinarius conoides Peintner & M.M. Moser nom. nov. Basionym: Secotium conicum Hesler 1933, Jour. Elisha Mitchell Soc. 49:153. Synonym: Thaxterogaster conicum (Hesler) Singer & A. H. Sm. 1958, Brittonia 10:214. ut Thaxterogaster conicus. non Cortinarius conicus (Velen.) Rob. Henry 1941. Cortinarius cretaceus (E. Horak) E. Horak (1980) [1979], Fl. criptog. Tierra del Fuego (Buenos Aires) 11(6):464. Basionym: Thaxterogaster cretaceum E. Horak 1965, Nova Hedwigia 10: 236; ut Thaxterogaster cretaceus. Cortinarius cunninghamii (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster cunninghamii E. Horak 1973, Beih. Nova Hedwigia 43: 92. Missapplied name: Secotium leucocephalum auct. non Massee in Cleland 1934-1935, in Toadst. Mush.South Australia:291. Cortinarius daucodes Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster subulatum E. Horak 1965, Nova Hedwigia X, 233. non Cortinarius Subulatus (Rob. Henry) Bidaud, Moénne-Loccoz & Reumaux 1997. Cortinarius dibaphoides (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster dibaphoides E. Horak 1979 (1980), Flora Criptogamica Tierra del Fuego (Buenos Aires) 11(6):33. Cortinarius dombeyi (Singer) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster dombeyi Singer 1960, Persoonia 1:385. Cortinarius epiphaeus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster epiphaeum E. Horak 1973, Beih. Nova Hedwigia 43:100: ut Thaxterogaster epiphaeus. Cortinarius flavidulus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster luteolum E. Horak 1973, Beih. Nova Hedwigia 43:105; ut Thaxterogaster luteo/us. non Cortinarius luteolus M.M. Moser 1975. Cortinarius flavovelus (Grgurinovic) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster flavovelus Grgurinovic 1997, Larger Fungi of South Australia:101. Missapplied name: Secotium ochraceum auct. non Rodway in Cleland 1934-1935, in Toadst. Mush. South Australia:289. Ge Errata Cortinarius fragilis (Zeller & C. W. Dodge) Peintner & M.M. Moser comb. nov. Basionym: Hymenogaster fragilis Zeller & C. W. Dodge in C. W. Dodge & Zeller 1934, Ann. Miss. Bot. Gard. 21:646. Synonym: Thaxterogaster fragile (Zeller & C. W. Dodge) A. H. Sm. & Singer 1963 in Singer and Smith, Madrono 17: 25; ut Thaxterogaster fragilis, ut Thaxterogaster fragilis var. fragilis. Cortinarius fragilis var. cortinariopsis (Singer) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster fragile var. cortinariopsis Singer 1969, Beihefte zur Nova Hedwigia 29: 364. Cortinarius fuegianus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster magellanicum Singer 1951, Mycologia 43:219. non Cortinarius magellanicus Speg. 1887. Cortinarius glutinopallens (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster glutinopallens E. Horak 1965, Nova Hedwigia X, 220. Cortinarius holojanthinus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster violaceum Singer 1951, Mycologia 43:216, ut Thaxterogaster violaceus var. violaceus. non Cortinarius violaceus (it thg) Rh S33: Cortinarius holojanthinus var. citrisporus Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster violaceum var. citrisporum M.M. Moser 1965, Nova Hedwigia X, 223. Cortinarius hygrophilus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster bibulum Singer 1969, Beihefte zur Nova Hedwigia 29: 361: ut Thaxterogaster bibulus. non Cortinarius bibulus Quél. 1881. Cortinarius janthinocaulis Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster violeipes M.M. Moser 1965, Nova Hedwigia X, 230. non Cortinarius violeipes M.M. Moser in M.M. Moser & E. Horak 1975. Cortinarius leucocephalus (Massee) Peintner & M.M. Moser comb. nov. Basionym: Secotium leucocephalum Massee 1891, Grevillea 19:95. ; Synonym: Thaxterogaster leucocephalum (Massee) Singer & A. H. Sm. 1958, Brittonia 10:210; ut Thaxterogaster leucocephalus. . Cortinarius levisporus (Massee & Rodway) Peintner & M.M. Moser comb. nov. Basionym: Hymenogaster levisporus Massee & Rodway in Rodway 1912, Pap. Proc. Roy. Soc. Tasm. 1911:30. Synonyms: Thaxterogaster levisporus (Rodway) G.W. Beaton, Pegler & T. W. K. Young 1985, Kew Bulletin 40: 172. Hymenogaster maideni Rodway 1921, Pap. Proc. Roy. Soc. Tasm. 1920:157. Octaviana levispora Rodway 1924, Pap. Proc. Roy. Soc. Tasm. 1923:157. Cortinarius lubricus (M.M. Moser) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster lubricum M.M. Moser 1965, Nova Hedwigia X, 226. Cortinarius luteirufescens (Bougher) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster /uteirufescens Bougher 1997, Mycotaxon 63:44-47. Cortinarius luteobrunneus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster leoninum E. Horak 1973, Beih. Nova Hedwigia 43:98; ut Thaxterogaster leoninus. non Cortinarius leoninus Velen. 1922. 180 181 Cortinarius luteocarnosus (Raithelh.) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster luteocarnosum Raithelh. (1983) Metrodiana, Sonderheft 2:21; ut Thaxterogaster luteocarnosus. Cortinarius mustella (M.M. Moser) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster mustella M.M. Moser 1965, Nova Hedwigia X, 227. Cortinarius napivelatus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster napivelatum E. Horak 1973, Beih. Nova Hedwigia 43:110; ut Thaxterogaster napivelatus. Cortinarius nivalis (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster nivale E. Horak 1973, Beih. Nova Hedwigia 43:89; ut Thaxterogaster nivalis. Cortinarius nothofagi (E. Horak) E. Horak (1980) [1979], Fl. criptog. Tierra del Fuego (Buenos Aires) 11(6): 452. Basionym: Thaxterogaster nothofagi E. Horak 1965, Nova Hedwigia X, 238. Cortinarius ochraceoazureus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster ochraceoazureum E. Horak 1965, Nova Hedwigia X, 237. Synonyms: Rozites ochraceoazurea (E. Horak) E. Horak (1980); Cortinanus ochrojanthinus E. Horak & M.M. Moser in M.M. Moser & E. Horak 1975: Nova Hedwigia, Beih. 52: 274: Cortinarius gliocyclus E. Horak in Moser & Horak 1975: Nova Hedwigia, Beih. 52: 275. Cortinarius ohauensis (Soop) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster ohauensis Soop 1998, Documents Mycologiques 18 (112):20. Cortinarius olorinus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster olorinum E. Horak 1965, Nova Hedwigia X, 218: ut Thaxterogaster olorinus. Cortinarius orphinus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster brunneus G. W. Beaton, Pegler & T. W. K. Young 1985, Kew Bulletin 40(1): 174. non Cortinarius brunneus (Pers. : Fr.) Fr. 1838. Cortinarius paveleckii (Trappe, Castellano & P. Rawlinson) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster paveleckii Trappe, Castellano & P. Rawlinson 2000, Mycotaxon 75:174-176. Cortinarius peraurantiacus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster aurantiacum E. Horak 1973, Beih. Nova Hedwigia 43:104. non Cortinarius aurantiacus M.M. Moser 1951. Cortinarius pingue (Zeller) Peintner & M.M. Moser comb. nov. Basionym: Secotium pingue Zeller 1941, Mycologia 33:211. Synonym: Thaxterogaster pingue (Zeller) Singer & A. H. Sm. 1958, Brittonia 10:21 1; ut Thaxterogaster pinguis. Cortinarius piriforme (Cleland & G. Cunn.) Peintner & M.M. Moser comb. nov. Basionym: Secotium piriforme Cleland & G. Cunn. 1924, Proc. Linn. Soc. New South Wales 49:115, pl. 15/3. Synonym: Thaxterogaster piriforme (Cleland & G. Cunn.) M.M. Moser 1965, Nova Hedwigia 10: 223; ut Thaxterogaster piriformis. Cortinarius pisciodorus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster pisciodorum E. Horak 1973, Beih. Nova Hedwigia 43:97: ut Thaxterogaster pisciodorus. 182 Cortinarius porphyroideus Peintner & M.M. Moser nom. nov. Basionym: Secotium porphyreum G. Cunn. 1924, Proc. Linn. Soc. New South Wales 49:114. Synonym: Thaxterogaster porphyreum (G. Cunn.) Singer 1958, Brittonia 10:212. ut Thaxterogaster porphyreus. non Cortinarius porphyreus E. Horak 1975, in M.M. Moser & E. Horak 1975. Cortinarius redactus (G. W. Beaton, Pegler & T. W. K. Young) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster redactus G.W. Beaton, Pegler & T. W. K. Young 1985, Kew Bulletin 40(1):177. Cortinarius sarcinochrous Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster carneolum E. Horak 1973, Beih. Nova Hedwigia 43:102; ut Thaxterogaster carneolus. non Cortinarius carneolus M.M. Moser & E. Horak 1975. Cortinarius scabrosus (Cooke & Massee) Peintner & M.M. Moser comb. nov. Basionym: Secotium scabrosum Cooke & Massee 1891, Grevillea 20:35. Synonym: Thaxterogaster scabrosum (Cooke & Massee) A. H. Sm. & Singer 1963, in Singer and A. H. Sm. Madrofo 17:23; ut Thaxterogaster scabrosus. Cortinarius sclerosperma Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster verrucisporum E. Horak 1965, Nova Hedwigia X, 226. non Cortinarius verrucisporus Thiers & A. H. Sm. 1969. Cortinarius sphaerocephalus (E. Horak & M.M. Moser) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster sohaerocephalum E. Horak & M.M. Moser 1965, Nova Hedwigia X, 232; ut Thaxterogaster sphaerocephalus. Cortinarius squamatus (Halling) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster squamatus Halling 1981, Mycologia 73: 863; ut Thaxterogaster squamatum. Cortinarius subalbidus (A. H. Sm.) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster subalbidum A. H. Sm. 1963, in Singer and Smith 1963, Madrofo 17: 24; ut Thaxterogaster subalbidus. Cortinarius thiersianus Peintner & M.M. Moser nom. nov. Basionym: Thaxterogaster thiersii Calhoun 1989, Mycotaxon 34: 65. non Cortinarius thiersii Ammirati & A. H. Sm. 1977. Cortinarius tylostomatoides (Singer) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster tylostomatoides Singer 1969, Beihefte zur Nova Hedwigia 29: 365. Cortinarius tympanicus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster tympanicum E. Horak 1965, Nova Hedwigia X, 224. Cortinarius violaceovolvatus (E. Horak) Peintner & M.M. Moser comb. nov. Basionym: Thaxterogaster violaceovolvatum E. Horak 1973, Beih. Nova Hedwigia 43:95; ut Thaxterogaster violaceovolvatus. Acknowledgments We thank Scott Redhead and Egon Horak for reviewing the manuscript and constructive comments, which certainly helped to improve the publication. This work was supported by an Erwin Schrédinger Auslandsstipendium (J1821- BIO) from the Austrian Science Foundation (FWF) to UP. 183 Literature cited Beaton, G., D. N. Pegler and T. W. K. Young. 1985. Gasteroid Basidiomycota of Victoria State, Australia: 3. Cortinariales. Kew Bulletin 40:167-204. Bougher, N. L. 1997. Three new sequestrate basidiomycetes from Western Australia. Mycotaxon 63: 37-48. Calhoun, C. J. 1989. Thaxterogaster thiersii - a new secotioid species from California. Mycotaxon 34:65-70. Greuter, W., J. McNeill, F. R. Barrie, H.-M. Burdet, V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silav, J. E. Skog, P. Trehane, N. J. Turland and D. L. Hawksworth. 2000. International Code of Botanical Nomenclature. St Louis Code. Electronic version of the original English text. http:/Awww.bgbm.fu- berlin.de/apt/nomenclature/code/SaintLouis/0000St. Luistitle. htm. Grgurinovic, C. A. 1997. Larger Fungi from Australia. The Botanic Gardens of Adelaide and State Herbarium, Adelaide. Halling, R. E. 1981. Thaxter' s Thaxterogasters and other Chilean hypogeous fungi. Mycologia 73:853-868. Heim, R. 1951. Notes sur la flore mycologique des Terres du Pacifique Sud. Ill. Sur les Secotium de Nouvelle-Zelandeet la phylogenie de ce genre. Revue Mycologique 16:129-153. Horak, E. 1973a. Fungi Agaricini Novazealandiae II. Thaxterogaster. Beih. Nova Hedwigia 43:87-114. Horak, E. 1973b. Fungi Agaricini Novazelandae I-VI. Beih. Nova Hedwigia 43. J. Cramer, Vaduz, Switzerland. Horak, E. 1979. Fungi, Basidiomycetes, Agaricales y Gasteromycetes secotioides. Flora Criptogamica de tierre del fuego 9:1-524. Horak, E. and M. Moser. 1965. Fungi Austroamericani XII. Studien zur Gattung Thaxterogaster Sing. Nova Hedwigia 10:211-241. Moser, M. 1964. Das System der Agaricales im Lichte neuer Forschung. Berichte Der Deutschen Botanischen Gesellschaft LXXVII, Sondernummer: 101-109. Peintner, U., N. L. Bougher, MA. Castellano, JM. Moncalvo, M.M. Moser, J. M. Trappe and R. Vilgalys. 2001. Multiple origins of sequestrate fungi related to Cortinarius (Cortinariaceae). American Journal of Botany, in press. Smith A. H. 1971. The origin and evolution of Agaricales. in R.H. Petersen, ed. The evolution of higher Basidiomycetes. University of Tennessee press, Knoxville, Tennessee, 481-504. Singer, R. 1951. Thaxterogaster - a new link between Gasteromycetes and Agaricales. Mycologia 43:215-288. Singer, R. 1960. Three new species of Secotiaceae from Patagonia. Persoonia (atete Singer, R. 1975. The Agaricales in modern Taxonomy. 3 ed. J. Cramer, Vaduz, Switzerland. Singer, R. and A. H. Smith. 1958. Studies in secotiaceous fungi. |. A monograph in the genus Thaxterogaster. Brittonia 10:201-216. Singer, R. and A. H. Smith. 1963. A revision of the genus 7haxterogaster Singer. Madrofio 17:22-26. 184 Soop, K. 1998. Cortinarius pa Undersidan. Jordstjarnan 19:25-37. Thiers, H. D. and A. H. Smith. 1969. Hypogeous Cortinarii. Mycologia 59:526- elon Thiers, H. D. 1984. The secotioid syndrome. Mycologia /6:1-8. Trappe, J. M. and M. A. Castellano. 2000. New sequestrate Ascomycota and Basidiomycota covered by the Northwest Forest Plan. Mycotaxon 75:153- 179. Zeller, S. M. 1941. Further notes on fungi. Mycologia 33,2:196-214. MY COTAXON Volume LXXXI, pp. 185-189 January-March 2002 TREMELLA SPICIFERA SP. NOV., A PARASITE OF MASSARINA ARUNDINACEA VAN RYCKEGEM, G. Ghent University, Dpt. Biology, Lab. Botany, K.L.Ledeganckstraat 35, B-9000 Gent, Belgium VAN DE PUT, K. Herentalsebaan 149, B-2100 Deurne, Belgium ROBERTS, BR: The Herbarium, Royal Botanic Gardens, Kew, Surrey TW9 3AE, UK Abstract Tremella spicifera, a new species parasitic on Massarina arundinacea, is described and illustrated from reeds in a tidal marsh of the river Schelde, Belgium. Keywords Heterobasidiomycetes, Tremellales, mycoparasitism, fungicolous, marsh During a survey of fungi on reeds (Phragmites australis (Cav.) Steud.) some unusual heterobasidiomycetes were found, including Tetragoniomyces uliginosus (P. Karst.) Oberw. & Bandoni (Van de Put et al., 2000) and an unknown Tremella species. One of the collections contained only remarkable, spike-like, clamped, conidiogenous cells bearing conidia formed from a succession of basal clamp-connexions. After incubation in a moist chamber tremellaceous basidia developed and the specimen proved to be identical to earlier collections. These represent a previously unknown Tremella_ species, described as follows: Tremella spicifera sp. nov., Van Ryckegem, Van de Put & P. Roberts Etym.: ‘spike-bearing’, named after the conidiogenous cells which resemble cylindrical grass spikes. Basidiomata 200- 600 pm lata, hyalina, in Massarina arudinacea. Hyphae 2-3 wm latae, usque ad 4-6 pm inflatae, fibulatae. Basidia 10-15 x 6-9 um, 186 septo longitudinali usque ad horizontali. Basidiosporae 4-5 um, globosae, hyalinae, apiculatae, Cellulae conidiogenae lageniformes, basaliter 4-6 um latae, collo usque ad 30 um longo, 1.5 um lato, spicae similes ob fibulas. Conidia 5-8 (12) x 2,5- 4 um, ellipsoidea vel cylindrica, hyalina, aseptata, -guttulata. Holotypus in Belgia, Provincia Flandria orientalis, Kallo, in palude salsa Scaldis lectus, parasitica Massarina arundinacea, 08 V 2000, leg. Van Ryckegem 575 (GENT). Basidiomes 200-600 um diam., hyaline, watery-gelatinous, in patches, growing parasitically on perithecia of Massarina arundinacea. Hyphae 2-3 uum wide, in places inflated up to 4-6 um wide, hyaline, clamped, frequently anastomosing, with thin-, refractive, or gelatinously thickened walls. Basidia 10-15 x 6-9 um, tremelloid, globose to ovoid, clavate to capitate, with longitudinal or sometimes transverse septa. Sterigmata 2-3 (-4), mostly up to 20 x 2 um, exceptionally up to 65 um long. Basidiospores 4-5 1m, globose, with distinct apiculus, hyaline, aguttulate, sometimes producing secondary spores. Conidiogenous cells arising from same hyphae as basidia, sometimes in fascicles, lageniform, 4-6 4m wide at base, neck up to 30 x 1.5 um, retaining when old the collapsed basal clamp-connexions of the conidia. Conidia dikaryotic, 5-8 (-12) x 2.5-4 um, elliptical to cylindrical, hyaline, with many small guttules, formed laterally and terminally on the conidiophores. Ecology: all the collections of the host (and parasite) were collected from brackish tidal marshes. Specimens examined: Belgium, East Flanders, River Schelde, Burcht, tidal marsh, IFBL-coordinates: C4.26.31, 27/03/99, Van Ryckegem 271 (GENT); Belgium, East Flanders, River Schelde, Doel, tidal marsh, IFBL-coordinates: B4.33.43, 27/03/99, Van Ryckegem 291 (GENT), K(M) 82839; Belgium, East Flanders, River Schelde, Kallo, tidal marsh facing Bayer, IFBL- coordinates: B4.54.42, 08/05/00, Van Ryckegem 575 (Holotype, GENT). Discussion This small 7remella species is characterized by its distinctive conidiogenous cells, the small basidiospores, the often thick-walled hyphae, and the host fungus. The formation of dikaryotic conidia in the genus 7remella often precedes and accompanies basidial development (Chen, 1998). Most species form terminal conidia, catenate conidia, or a few successive conidia. The conidiogenous cells in Tremella spicifera resemble those found in Tremella 187 occultifuroidea C.-J. Chen & Oberw. (Chen et al., 1999) and 7. polyporina D.A. Reid (illustrated in Koske, 1972). The former is a parasite of Dacrymyces species, the latter of polypores. Similar cells have also been reported from 7remella caloceraticola Hauerslev, a parasite of Calocera (Hauerslev, 1999), and the anamorphic 7remellina pyrenophila Bandoni, a parasite of Ceratocystis species on palms, whose conidiogenesis was discussed by Bandoni (1986). The lichen parasite, Tremella stevensiana Diederich, forms conidiogenous cells producing multiple young conidia which succesively mature. Clamp-connexions are lacking (Diederich & Christiansen, 1994). Tremella isabellina Lloyd shows a similar type of conidiogenesis (Bandoni, 1958). Outside the Tremellales, two auricularioid species, Occultifur internus (L.S. Olive) Oberw. (Oberwinkler, 1990) and Platygloea abdita Bandoni (Bandoni, 1959), form similar conidiogenous cells, as does Krieglsteinera lasiosphaeriae Pouzar (illustrated in Leessge & Lange, 1998). The very small, globose basidiospores are also exceptional, though not abundantly formed in the material examined. Species producing spores of this size include 7remella polyporina (Reid, 1970), T. colpomaticola Hauerslev, 7. karstenii Hauerslev, T. silvae-dravedae Hauerslev (Hauerslev, 1999), and several lichenicolous species (Diederich, 1996). Spore germination and a yeast stage have not yet been observed in 7remella spicifera. Many 7remella species are mycoparasites of Ascomycetes, but for most of them host-parasite interactions have still to be studied. Haustoria, typical for the genus, were not observed in Tremella spicifera but may well occur. No literature records could be found of any other 7remella species parasitizing Massarina arundinacea (Sowerby: Fr.) Leuchtm. or other members of the Lophiostomataceae (Dothideales). Acknowledgements We would like to thank Dr. A. Verbeken for critically reading the paper and correcting the Latin diagnosis. Research financed with research funds of the Institute for the promotion of Innovation by Science and Technology in Flanders, Belgium. References Bandoni, R.J. (1958). Some tremellaceous fungi in the C. G. Lloyd collection. Lloydia 21: 137- io Bandon, R.J. (1959). An undescribed species of Platygloea from Iowa. Mycologia 51: 94-90. Bandoni, R.J. (1986). On an undescribed basidiomycetous anamorph. Windahlia 16: 53 — 58. Chen, C. (1998). Morphological and molecular studies in the genus 7remella. Biblioth. Mycol. 174: 1-225. Chen, C., Bandoni, F. & Chen, Z. (1999). Tremella occultifuroidea sp. nov., anew mycoparasite 188 of Dacrymyces. Mycoscience 40: 137-143. Diederich, P. (1996). The lichenicolous heterobasidiomycetes. Biblioth. Lichenol. 61: 1-198. Diederich, P. & Christiansen, M.S. (1994). Biatoropsis usnearum Rasanen, and other heterobasidiomycetes on Usnea. Lichenologist 26: 47-66. Hauerslev, K. (1999). New and rare species of heterobasidiomycetes. Mycotaxon 72: 465-486. Koske, R.E. (1972). Two unusual Tremellas from British Columbia. Canad. J. Bot. 50: 2565- 2501: Lessoe, T. & Lange, C. (1998). Krieglsteinera lasiosphaeria — en hojst besynderlig snylter pa Lasiosphaeria ovina. Svampe 37: 42-44. Oberwinkler, F. (1990). New genera of auricularioid heterobasidiomycetes. Rep. Tottori Mycol. Inst. 28: 113-127. Reid, D.A. (1970). New or interesting records of British Hymenomycetes, IV. Trans. Brit. Mycol. Soc. 55: 413-441. Van de Put, K., Van Ryckegem, G. & Antonissen, I. (2000). Eerste vondsten van Tetragoniomyces uliginosus (P. Karst.) Oberw. & Bandoni in Vlaanderen. Sterbeeckia 19: 23-26. P20. ot & S22 Se, an Cag TN pea Pevaue 10 pm Plate 1. a. Several stages of conidiogenous cells and basidia. Note frequent anastomoses. b. Conidia and conidiogenous cells. c. Basidiospores. 189 10 tum Plate 2. a. Conidiogenous form. b. Conidia. c. Basidiospores. ! . myc. al 2 ee ae ; hye sae Hebe} : i) PT ribs WK | At ma) as tN Cone anv suet Rh nen ‘ | , s ae) he ] 5 ' x ne nig ot, bat ‘? ow ‘ ris alls ia ae Pits him 7h bien’) red 8 ti Vad Mibe ya ty i rate eno NL fa iti ihe bi ter Mig ‘ye fh et ; i L =} ‘weet “ Ne boi ; ' 1) | eileen Ray ee ehawers | 4 ; ae wie BA Ay aay taste ai ' i} ie 7 A re ae \ Af 2 a Ae » ny Mi Mares VAS, ie +f ey “i ya ee Adi aay et hao ) pm naa 13 ¢ mae ae y | ‘ , a, ee iy ‘, lasted watt y of fi i" a, 7 a he ae * a inky Bt r lye his cs et ih hy wnat at k : nee “n ' i / ia | ) Seer, Pte ie meg {4 . 7 5 ™ ine hat iy PVE a Polk far igi 1} : i" vw Pe | ’ bi iene A are (eters, ic, PR a ta, ‘i hs i 5 aoe \ Ltt i ete ei, mr ie A i sah aiad at M : | Be nae age a pnt in WR he cfr Wak ee i 1 c [ _ = DP Te» by j ae a, a ~ al f ‘ : 5 it & <= a. Matt! yl ae i i it " + . en | ai hy ‘ ; ' Ay ak? Ma iS As : | Asal oe i Nae Wane) Pi: { 7 : Le ee’ ira <3 i Dri ; } \ A? ee : Te cals 3 I : ae hi mh a a it ' 2 4 s = 7 H | 7 A) sey Ae e Ne a ae i ie : - . ae e ff hg y i Tw, An Mel thd fer ¥ ai | i fy 5 et fi. i A y i] "y il oe | iH] ; pee y ton y i} a ny om. Segara: Os Pes Hy die far He eet Was a a ane ans : + = ’ We de 5 {| ’ ithe ; j i ‘ at f ae ix i Iq if Wi iH i} i tet a er ; , } ¥ res 7 * ark | aie. si ‘i ‘ ; ae at r ‘ es F Se, a 4 : cs ‘ a» eheoxrs 4 ~ it hee, or ep Nae arate Se aa Se PN yh Si ee d v4 y Paks bled ‘Ty oe * n i FAK Teal er’ | | ! : ; ’ , i A AD ‘ ral y i Dey, 4 7 bi ; 7. : cS Te h ie nd pre ad : (; “oJ 4 An. - 7 & ce in hice Sa 2 el a a oe ec rare ah PRT yy ‘ fs sf ee ne ear a { id oa bys 1's a in : 7 Y- bal ey ‘ ¥ 7 : oh Pe Ae ep. - ' fi>@ We v' ® } *. ed At Mri. Le - 1p A ne pt MYCOTAXON Volume LXXXI, pp. 191-194 January-March 2002 THE MATING SYSTEM IN TWO SPECIES OF CREPIDOTUS M. Catherine Aime! and Jessica Ball Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061 'Present address: Department of Plant Sciences, University of Oxford, Oxford OX1 3RB, U.K. Email: cathie.aime@plant-sciences.oxford.ac.uk Abstract Crepidotus aureifolius and C. nephrodes (Agaricales: Crepidotaceae) exhibit a unifactorial (bipolar) mating system. This is the first known report of mating studies conducted in this genus. Key Words: agarics, basidiospores, bipolar mating system Introduction This study is one component of a larger study that will attempt to apply morphological, biological, and phylogenetic species concepts toward a systematic revision of the genus Crepidotus (Fr.) Staude. Crepidotus is comprised of mostly small, pleurotoid, saprophytic agarics, with an estimated 150 species in the genus (Singer 1986). Section Sphaerula Hesler and Smith contains 42 taxa, including C. aureifolius Hesler and Smith and C. nephrodes (Berk. & Curt.) Sacc., many of which are distinguished on the basis of single morphological character differences that may or may not be true indicators of non-interbreeding species. In order to apply a biological species concept within this complex the recovery of single spore isolates and determination of mating types is of paramount importance (Petersen 1995). This paper is the first to report on the mating system in a species of Crepidotus, based on intracollection pairings of single spore isolates in two collections of Crepidotus Section Sphaerula. Materials and Methods Specimens: USA: NORTH CAROLINA, Iredell County, West of Bridge 766, 17-Sept. 1997, coll. A. Stanley, MCA 373 (VPI); VIRGINIA, Giles County, Jefferson National Forest, Cherokee Flats, 14- Sept. 1996, M.C. Aime 189 (VPI). All cultures and mating crosses were grown on malt extract agar (15g Sigma malt extract, 15g Sigma agar, | L distilled HO) and incubated in the dark at 24-26 C. Methods for dilution plating and isolation of monkaryotic isolates follow Aime (1999). Isolates were stored at 4 C for two-three years prior to use. Isolates were then grown out and examined microscopically for the absence of clamp connections before being used in crosses. Confirmed monokaryons were crossed, in all possible pair-wise combinations, on an individual malt agar plate for each cross, and incubated 192 for approximately four weeks. Hyphae from the contact zone on each cross was then mounted in Congo red and examined under oil immersion; compatible matings were inferred by the presence of clamp connections; the absence of clamp connections was interpreted as an incompatible mating. The stability of dikaryons was tested by transferring a small portion of the interface from a compatible cross onto a new malt agar plate and incubating for two months or until growth had covered the plate. A portion of the outermost region of the culture was then examined for clamps as above. In addition, all crosses that initially appeared negative at the four-week examination were re-examined after two to eight weeks of additional incubation at 24-26 C. A total of eight monokaryotic isolates for MCA 373 and six for MCA 189 were recovered which maintained viability throughout the study. These cultures are maintained in the Virginia Tech Culture Collection as VT'CC 3600.2, VTCC 3600.4, VTCC 3600.8, VTCC 3600.12, VTCC 3600.14, VTCC 3600.15, VT'CC 3600.18, and VTCC 3600.39 (MCA 373) and VTCC 3588.1, VIT'CC 3588.3, VT'CC 3588.5, VTCC 3588.8, VTCC 3588.13, and VTCC 3588.18 (MCA 189). Results and Discussion Spore prints from a total of 49 Crepidotus collections representing 27 morphological taxa were plated during a three year period in an attempt to obtain monokaryotic single spore isolates (SSIs) (Aime 1999, Aime and Miller Chap. 4). The same studies found that the recovery of SSIs in Crepidotus was complicated by a dormant period in the temperate North American species and by a very low rate of germinations per plate. Ultimately, only nine of the collections plated in those studies yielded more than ten SSIs; the largest number of isolates (39) was obtained for collection MCA 373, C. aureifolius, which was repeatedly plated to gauge the effect of various abiotic factors on spore dormancy and germination. Furthermore, it was discovered that the viability of monokaryotic isolates was reduced after long term storage at 4 C. Asa result, only two collections of Crepidotus, MCA 373 and MCA 1839, yielded an adequate number of SSIs for inferring mating types. Figure 1 shows the results from intracollection matings of Crepidotus aureifolius (MCA 373). Two mating types were identified indicating the presence of a unifactorial (bipolar) mating system. Assignment of mating types for this collection are shown in Table 1. Figure 2 shows the results from intracollection matings of Crepidotus nephrodes (MCA 189), which also indicate a unifactorial system; assignment of mating types are given in Table 2. Contact zone morphology varied among crosses from almost no observable interaction to a dense hyaline line of mycelium built up at the interaction zone. No correlation could be made between the contact zone morphology and the type of mating interaction. However, there was some tendency for isolates to form certain types of interactions, as has been found in the genus Melanotus (Petersen 1992). 193 |__| 3600.4 _| 3600.8 | 3600.12 | 3600.14 | 3600.15 | 3600.18 | 3600.39 3600.14 3600.15 3600.18 3600.12 + + Fig. 1. Intracollection mating of single spore isolates of Crepidotus aureifolius (MCA 373). + = clamp connections formed; — = no clamp connections formed. | —_——__—{_ 3588.3 | 3588.5 | 3588.8 | 3588.13 | 3588.18 3588.1 3588.3 3588.8 3588.13 Fig. 2. Intracollection mating of single spore isolates of Crepidotus nephrodes (MCA 189). += 3588.5 + clamp connections formed; — = no clamp connections formed. Table 1. Assignment of mating types to Table 2. Assignment of mating types to single spore isolates of C. aureifolius, based single spore isolates of C. nephrodes, based on intracollection crosses. on intracollection crosses. Mating type Isolate number Mating type Isolate number Al By 2 Ne FG PO oa BS Al pies is. A2 1453152939 A2 BSED LOLS er eee eee ee ere ee ee eee 194 Specifically, a nodulose interaction zone was only formed in some of the crosses with isolate 3600.18, while a dense build-up of hyaline mycelium forming a line between isolates was found in three of the seven crosses with isolate 3600.4. The morphology of clamp connections was also diverse and clamp morphology appears to correlate with isolate strain. The majority of dikaryons produced clamps at about 20% of septa. In contrast, one strain (3600.15), when crossed with compatible isolates, produced fewer clamps of reduced size, while compatible crosses formed with isolate 3600.39 produced an abundance of larger clamps. In some compatible pairings, clamp connections were not observed until the cross had been allowed to incubate for an additional two to eight weeks after a contact zone had been established. However, all dikaryons obtained from the interface of compatible pairings and grown out separately remained stable and continued to exhibit clamp connections at the growing margin after several months incubation. Since it has been demonstrated that, statistically, a minimum of 11 SSIs from a single collection should be self-crossed to ensure retrieval of all four mating types in a tetrapolar mating system (Vilgalys 1982), it is possible but unlikely that, had more isolates been available for crosses, two additional compatible mating types would have been uncovered, resulting in a bifactorial system for this fungus. However, the same statistics show that at least three mating types are likely to be uncovered when either six or eight SSIs are crossed pair-wise. We therefore conclude that these fungi possess a unifactorial mating system. Acknowledgments The authors wish to thank Allein Stanley for donation of material used in this study; Orson K. Miller, Jr. for experimental suggestions; and Jack Murphy for reviewing an earlier version of this manuscript, and his many helpful comments and suggestions. This paper is based in part on a dissertation submitted by the first author to Virginia Tech as a requirement for a Ph.D. degree. References Aime, M.C. 1999. Generic Concepts in the Crepidotaceae as Inferred from Nuclear Large Subunit, Ribosomal DNA Sequences, Morphology, and Basidiospore Dormancy Patterns. M.S. Thesis, Virginia Polytechnic Institute and State University, Blacksburg, VA. 127 pp. Aime, M.C. and O.K. Miller, Jr. 2001. Delayed germination of basidiospores in temperate species of Crepidotus (Fr.) Staude. Canadian Journal of Botany. In press. Petersen, R.H. 1992. Further notes on mating systems in Melanotus. Mycotaxon 45:33 1-341. Petersen, R.H. 1996. There’s more to a mushroom than meets the eye: Mating studies i in the Agaricales. Mycologia 87:1-17. Singer, R. 1986. The Agaricales in Modern Taxonomy. 4" ed. J. Cramer, Germany. 981 pp. Vilgalys, R. 1982. A Biosystematic Study of Collybia dryophila (Basidiomycetes: Tricholomataceae). M.S. Thesis, Virginia Polytechnic Institute and State University, Blacksburg, VA. 115 pp. MY COTAXON Volume LXXXI, pp. 195-214 January-March 2002 NOMENCLATURAL REVISIONS IN THE SEQUESTRATE RUSSULOID GENERA James M. Trappe Department of Forest Science, Oregon State University, Corvallis, Oregon, U.S. A. 97331-5752 trappej@ucs.orst.edu Teresa Lebel National Herbarium of Victoria, Royal Botanic Gardens, Birdwood Avenue, South Yarra, Victoria, Australia 3141 teresa.lebel@rbg.vic.gov.au Michael A. Castellano U. S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon 97331 meastellano@fs.fed.us ABSTRACT Type studies of the genera of sequestrate russuloid fungi by Lebel and Trappe plus extensive studies of Australasian taxa led to emended generic descriptions of Cystangium and Gymnomyces and elimination of the genera Martellia and Elasmomyces. Here we bring names of described species into conformity with these changes by new combinations: Martellia spp. are transferred into either Cystangium or Gymnomyces, Elasmomyces spp. into either Cystangium or Macowanites, and some species are transferred from one to another of the retained genera in keeping with the emended concepts. As a result, 19 names of Cystangium, 37 of Gymnomyces, and 34 of Macowanites are accepted. In our studies of holotypes, 8 species previously described in various genera were found instead to relate to Lactarius and are transferred to the genus Zelleromyces. One species previously regarded as sequestrate is transferred to the nonsequestrate genus Russula. Two species formerly assigned to russuloid genera are transferred to the nonrussuloid genus Octaviania. INTRODUCTION Singer and Smith (1960) recognized five sequestrate genera related to Russula, Cystangium, Elasmomyces, Gymnomyces, Macowanites and Martellia, and two related to Lactarius, Arcangeliella and Zelleromyces. They differentiated the genera primarily by presence or absence of 1) a stipe-columella, 2) sphaerocysts in the glebal trama, and 196 3) latex production, based on species known at that time. Since then many more species have been discovered. Australia especially has proven to be a rich source of russuloid sequestrate fungi, and studies of the Australian taxa have revealed important new data on their morphological relationships (Lebel 1998). Lebel and Trappe (2000) studied the type species of each of these genera in detail, and Lebel and Castellano (2002) redescribed the holotypes of all published Australasian russuloid species. Calonge and Martin (2000) sequenced DNA of selected species of sequestrate and epigeous Russulales. Taken together, these studies showed that the characters of stipe-columella and tramal sphaerocysts emphasized by Singer and Smith (1960) were often ambiguous, in that many intergradations occurred within and between specimens, species and genera, and confirmed the relationship of these sequestrate genera to the epigeous Russulaceae as postulated by Singer and Smith (1960), thereby rejecting the separation by Pegler and Young (1979) of Elasmomyces, Gymnomyces, Martellia and Zelleromyces into their artificial family Elasmomycetaceae. Miller et al. (2001) sequenced the nuclear large subunit rDNA of an array of Russulaceae, including both agaricoid and sequestrate species. Their data further confirmed the phylogenetic relationships between the two groups and indicated that the sequestrate taxa arose along several different lineage’s within the Russulaceae. Lebel and Trappe (2000) emended the generic concepts of the sequestrate russuloid fungi to clarify distinctions between genera as follows: Cystangium with or without stipe-columella; peridiopellis with either a single, exposed epithelial layer of inflated, often isodiametric cells or two layers with the epithelium underlying a suprapellis of hyphae and/or cystidia; gleba sublamellate or loculate. Gymnomyces without a stipe but sometimes with a reduced columella; peridiopellis a trichodermium, a turf or of undifferentiated hyphae; gleba loculate. Macowanites with stipe-columella; peridiopellis a trichodermium, a turf or of undifferentiated hyphae; gleba sublamellate or loculate. Martellia and Elasmomyces were eliminated; Martellia species would be reassigned to either Cystangium or Gymnomyces and Elasmomyces species to either Cystangium or Macowanites, in both cases depending on anatomy of the peridiopellis. This revision removed much of the ambiguity inherent in the generic concepts of Singer and Smith (1960), but Lebel and Trappe (2000) regarded it as an interim solution pending development of a comprehensive molecular phylogeny of the Russulaceae. Miller et al. have taken a further step towards that goal. However, much additional research is needed to produce such a phylogeny, which must be based on concordance of multiple gene genealogies (Taylor et al. 2000). Until that happens, researchers on taxonomy, ecology and physiology of these important fungi need a useable nomenclatural system to organize and communicate their research results. In the present paper we recombine previously described russuloid sequestrate species for which we have adequate data either from our examination of holotypes or from type descriptions to bring them into conformity with the generic concepts as emended by Lebel and Trappe (2000). Asa result, 19 names of Cystangium, 37 of Gymnomyces, and 34 of Macowanites are accepted. Taxonomic synonyms are listed when we have confirmed them, but it must be remembered that the recombinations are mostly nomenclatural changes and should not necessarily be regarded as final taxonomic conclusions. 197 Some species originally placed in one or another of these genera proved to belong to the Lactarius-related genera Arcangeliella or Zelleromyces. In our studies of holotypes, 8 species previously described in various genera were found instead to relate to Lactarius and are transferred to the genus Zelleromyces. Names of sequestrate lactarioid species are not otherwise listed, because we have not completed our studies of them. Macowanites americanus Singer & A. H. Sm. was judged to be a nonsequestrate Russula spp. and is renamed R. subloculata to avoid erecting a homonym to Russula americana (Singer) Singer. These new names are included here under the heading “Excluded species.” Species for which type collections have not been examined or type descriptions are inadequate are listed under the heading “Uncertain species.” All references are cited in full in the Literature Cited section to provide a complete bibliography and facilitate literature search and acquisition by any who have an interest in these fungi. All new names and combinations are indicated in boldface. An index to species names, some of which have appeared under as many as five different genera in the past is provided after the Literature Cited. > ACCEPTED NAMES, NEW COMBINATIONS, AND SYNONYMS OF RUSSULA-RELATED SEQUESTRATE SPECIES Cystangium balpineum Grgur., Larger Fungi of South Australia, p. 53. 1997. Cystangium capitis-orae (Dring) T. Lebel comb. nov. BASIONYM: Elasmomyces capitis-orae Dring in Dring & Pegler, Kew Bull. 32: 564-566. 1978. Cystangium depauperatum Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 69-70. 1960. Cystangium echinosporum (Zeller & C. W. Dodge) Trappe, T. Lebel & Castellano, comb. nov. Basionym: Macowanites echinosporus Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 6: 57-58. 1919. = Elasmomyces echinosporus (Zeller & C. W. Dodge) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 22: 370. 1935. Cystangium idahoensis (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia idahoensis Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 43-44. 1960. Cystangium lymanensis (Cazares & Trappe) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Macowanites lymanensis Cazares & Trappe, Mycotaxon 42: 335. 19903 _ Cystangium maculatum (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia maculata Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 28-29. 1960. Cystangium medlockii (Trappe & Castellano) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia medlockii Trappe & Castellano, Mycologia 78: 918-919. 1986. 198 Cystangium megasporum (Rodway) T. Lebel & Castellano, Mycologia 94 (in press) 2002. = Gymnomyces megasporus Rodway, Paps. Proc. Roy. Soc. Tasmania 1925: 168. 1926. = Octaviania megaspora (Rodway) G. Cunn., N. Zeal. J. Sci. Tech. 22: 300B. 1941. Cystangium nothofagi (E. Horak) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces nothofagi E. Horak, Sydowia 17: 211-212. 1964a. Cystangium oregonense (Zeller) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Hydnangium oregonense Zeller, Mycologia 33: 200. 1941. = Martellia oregonensis (Zeller) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 21 1960. Cystangium phymatidosporum G. W. Beaton, Pegler & T. W. K. Young, Kew Bull. 39: 672-674. 1984. Cystangium pineti Singer, Mycol. Helvetica 1: 417-419, 1985. Cystangium rodwayi (Massee) A. H. Sm., Mycologia 54: 633. 1962. = Secotium rodwayi Massee, Kew Bull. Misc. Inf. 1901: 158. 1901. = Elasmomyces rodwayi (Massee) Zeller, Mycologia 40: 642. 1948. Cystangium seminudum (Massee & Rodway) T. Lebel & Castellano, Mycologia 94 (in press) 2002. = Gymnomyces seminudus Massee & Rodway in Massee, Bull. Misc. Inf. Kew 1898: 125. 1898. = Arcangeliella seminuda (Massee & Rodway) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 23: 617. 1936. = Octaviania seminuda (Massee & Rodway) G. Cunn., Trans. Roy. Soc. New Zeal. 67: 408. 1938. = Gymnomyces solidus Rodway, Paps. Proc. Roy. Soc. Tasmania 1920: 157. 1921. Cystangium sessile (Massee & Rodway) Singer & A. H. Sm., Mem. Torrey Bot. Club 21s OF L900: = Secotium sessile Massee & Rodway in Rodway, Paps. Proc. Roy. Soc. Tasmania 1911: 31. 1912. = Elasmomyces sessilis (Massee & Rodway) Rodway, Paps. Proc. Roy. Soc. Tasmania 1924: 8. 1925. Cystangium thaxteri (Singer) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia thaxteri Singer, Nova Hedwigia Beth. 29: 357. 1969.- Cystangium variabilisporum (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia variabilispora Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 27-28. 1960. Cystangium vesiculosum (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia vesiculosa Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 29. 1960. Gymnomyces albus (Harkn.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Hydnangium album Harkn., Proc. Cal. Acad. Sci. III, 1: 251. 1899. = Martellia alba (Harkn.) Singer & Sm., Mem. Torrey Bot. Club 21: 41. 1960. Gymnomyces ammophilus J. M. Vidal & Calonge in Calonge & J. M. Vidal, Bol. Soc. Micol. Madrid 24: 66. 1999. 1s Gymnomyces brunnescens (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia brunnescens Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 37-38. 1960. Gymnomyces californicus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia californica Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 33-34. 1960. Gymnomyces cinnamomeus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 51. 1960. Gymnomyces citrinus (Harkn.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Octaviania citrina Harkn., Proc. Cal. Acad. Sci. HI, 1: 252. 1899. = Hydnangium citrinum (Harkn.) Zeller & C. W. Dodge, Ann. Mo. Bot. Gard. Paps B bs SMU Rey, Gymnomyces compactus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 53-54. 1960. Gymnomyces cremeus (Zeller & C. W. Dodge) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Arcangeliella cremea Zeller & C. W. Dodge, Ann. Mo. Bot. Gard. 22: BOERII3 5: = Martellia cremea (Zeller & C. W. Dodge) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 45. 1960. Gymnomyces dominguezii Moreno-Arroyo, J. Gomez & Calonge, Mycol. Res. 103: 215. 1999) : Gymnomyces eildonensis G. W. Beaton, Pegler & T. W. K. Young, Kew Bull. 39: 680. 1984. Gymnomyces ellipsosporus (Zeller) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Aydnangium ellipsosporum Zeller, Mycologia 31: 13. 1939. = Martellia ellipsospora (Zeller) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 30..1960. Gymnomyces fallax (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia fallax Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 34- 35. 1960. Gymnomyces ferruginascens Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 54-55. 1960. Gymnomyces foetens (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia foetens Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 36- 37. 1960. Gymnomyces fragrans (A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia fragrans A. H. Sm., Mycologia 55: 437. 1963. Gymnomyces fulvisporus (A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia fulvispora A. H. Sm., Mycologia 55: 438. 1963. Gymnomyces gilkeyae (Zeller & C. W. Dodge) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Aydnangium gilkeyae Zeller & C. W. Dodge, Ann. Missouri Bot. Garda 227 belo, = Martellia gilkeyae (Zeller & C. W. Dodge) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 32. 1960. | 200 Gymnomyces ilicis Llistosella & J. M. Vidal, Riv. Micol. 2: 156. 1995. Gymnomyces luteolus (Harkn.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Hydnangium luteolum Harkn., Proc. Cal. Acad. Sci. HI, 1: 251. 1899. Gymnomyces mistiformis (Mattir.) T. Lebel & Trappe, Mycologia 92: 1196. 2000. = Martellia mistiformis Mattir., Malpighia 14: 78-82. 1900. = Hydnangium mistiforme (Mattir.) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard? 2273/25193>. = Octaviania mistiformis (Mattir.) Svrcek in Pilat, Flora C.S.R., p. 192. 1958. Gymnomyces monosporus E. L. Stewart & Trappe, Mycotaxon 2: 209-210. 1975. Gymnomyces monticola (Harkn.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Octaviania monticola Harkn., Proc. Cal. Acad. Sci. II, 1: 254. 1899. = Hydnangium monticola (Harkn.) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard?2273727 1935: = Martellia monticola (Harkn.) Singer & Sm., Mem. Torrey Bot. Club 21: 40. 1960. Gymnomyces nanjingensis (B. Liu & K. Tao) Trappe, T. Lebel & Castellano, comb. nov. Basionym: Martellia nanjingensis B. Liu & K. Tao in K. Tao, M. C. Chang & B. Liu, Acta Mycol. Sinica 12: 103. 1993. Gymnomyces occidentalis (Harkn.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Octaviania occidentalis Harkn., Proc. Cal. Acad. Sci. III, 1: 253. 1899. Gymnomyces pallidus Massee & Rodway in Massee, Bull. Misc. Inf. Kew 1898: 125. 1898. = Octaviania pallida (Massee & Rodway) G. Cunn., Proc. Linn. Soc. New S. Wales 60: 119, 1935. = Hydnangium hinsbyi Rodway, Proc. Roy. Soc. Tasm. 1923: 158. 1924. = Octaviania hinsbyi (Rodway) G. Cunn., Trans. Roy. Soc. N. Zeal. 67: 408. 1938. Gymnomyces parksii Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 55. 1960. Gymnomyces pila (Pat.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Hydnangium pila Pat., Bull. Tri. Soc. Mycol. France 26: 201. 1910. = Martellia mediterranea G. Moreno, R. Galan & Montecchi, Mycotaxon 42: 227- 228. 1991. (See Moreno, et al. 1991). Gymnomyces redolens (G. Cunn.) Pfister, Occ. Pap. Farlow. Herb. 9: 43. 1976. = Octaviania redolens G. Cunn., New Zeal. J. Sci. Tech. 23: 172. 1942. = Stephanospora redolens (G. Cunn.) E. Horak in Oberw. & E. Horak, comb. illeg., Plant Syst. Evol. 131: 163. 1979. = Martellia redolens (G. Cunn.) G. W. Beaton, Pegler & T. W. K. Young, Kew Bull. 39: 682. 1984. Gymnomyces rogersii (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. | BASIONYM: Octavianina rogersii Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 12-13. 1960. 201 Gymnomyces rolfalexii Trappe, T. Lebel & Castellano, nom. nov. BASIONYM: Martellia parksii Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 32- 33, 1960, non Gymnomyces parksii Singer & A. H. Sm., Mem. Torrey Bot. Club Pi 5581960! Gymnomyces roseomaculatus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 50. 1960. Gymnomyces setigerus (Zeller) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Hydnangium setigerum Zeller, Mycologia 31: 14-15. 1939. = Martellia setigera (Zeller) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 39 1960. Gymnomyces subalpinus (A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia subalpina A. H. Sm., Mycologia 55: 439. 1963. Gymnomyces subfulvus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia subfulva Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 35-36. 1960. Gymnomyces subochraceus (A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia subochracea A. H. Sm., Mycologia 55: 440. 1963. Gymnomyces wirrabarensis Grgur., Larger Fungi of South Australia p. 86-88. 1997. Gymnomyces xanthosporus (Hawker) A. H. Sm., Mycologia 54: 635. 1962. Hydnangium carneum var. xanthosporum Hawker, Trans. Brit. Mycol. Soc. 35: 279. L952: Macowanites acris Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 86-86. 1960. Macowanites agaricinus (Kalchbr.) Kalchbr., Hedwigia 15: 115-116. 1876a. = Macowania agaricina Kalchbr. in Berk., nom. illeg., Gardeners Chron. 5 (NS): 785. 1876b. Macowanites albidigleba Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 85-86. 1960. Macowanites carmineus McNabb, New Zeal. J. Bot. 9: 359, 1971. Macowanites chlorinosmus A. H. Sm. & Trappe, Mycologia 55: 423-424. 1963. Macowanites citrinus Singer & A. H. Sm.. Mem. Torrey Bot. Club 21: 69-70. 1960. Macowanites durangensis Guzman, Rev. Mex. Micol. 4: 118-119. 1988. Macowanites fulvescens Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 84-85. 1960. Macowanites fuscoviolaceus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 87-88. 1960. Macowanites galileensis M. M. Moser, Binyam. & Aviz.-Hersh., Trans. Brit. Mycol. SOC. 0825) LALO iy se Macowanites iodiolens A. H. Sm. & V. L. Wells, Mycologia 55: 425-426. 1963. Macowanites krjukowensis (Buchholtz) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 77. 1960. = Secotium krjukowensis Buchholtz, Hedwigia 40: 314. 1901. = Elasmomyces krjukowensis (Buchholtz) Sacc. & D. Sacc., Syll. Fung. 17: 218. 1905) = Bucholtzia krjukowensis (Buchholtz) Lohwag, Osterr. Bot. Zeitschr. 73: 173. 1924. 202 = Arcangeliella krjukowensis (Buchholtz) Zeller & C. W. Dodge, Ann. Missouri Boty Gard? 22236821935: = Hydnangium krjukowense (Buchholtz) Svrcek in Pilat, Flora C.S.R., Gasteromycetes, p. 206. 1958. = Secotium michailowskianum Buchholtz, Hedwigia 40: 315. 1901. = Elasmomyces michailowskianus (Buchholtz) Sacc. & D. Sacc., Syll. Fung. 17: ZNSE 905. / = Arcangeliella krjukowensis var. michailowskiana (Buchholtz) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 32: 368. 1935. Macowanites lilacinus A. H. Sm., Mycologia 55: 426-427. 1963. Macowanites luteiroseus Bougher, Mycotaxon 58: 38-39. 1997. Macowanites luteolus A. H. Sm. & Trappe, Mycologia 55: 427-428. 1963. Macowanites magnus Parks in Zeller & C. W. Dodge, Ann. Missouri Bot. Bard. 22: 369. 1939: = Arcangeliella magna (Parks) Zeller, Mycologia 40: 643. 1948. Macowanites mattirolanus (Cavara) T. Lebel & Trappe, Mycologia 92: 1194. 2000. = Elasmomyces mattirolanus Cavara, Malpighia 11: 426. 1897. = Secotium mattirolanum (Cavara) E. Fischer, Engl. & Prantl, Naturl1. Pflanzenfam. Abt. 1: 301-302. 1900. Macowanites messapicoides Llistosella & J. M. Vidal, Riv. Micol. 38: 150. 1995. Macowanites mexicanus Guzman, Rev. Mex. Micol. 4: 116-117. 1988. Macowanites mollis Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 79-80. 1960. = Elasmomyces mollis (Singer & A. H. Sm.) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: 368. 1979. Macowanites nauseosus A. H. Sm., Mycologia 55: 428-429. 1963. Macowanites odoratus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces odoratus Singer & A. H. Sm., Mem. Torrey Bot. Club 2 Orel 900. Macowanites olidus A. H. Sm., Mycologia 55: 429-430. 1963. Macowanites pilosus (Zeller & C. W. Dodge) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: 4rcangeliella pilosa Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 22230801939) = Elasmomyces pilosus (Zeller & C. W. Dodge) Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 66. 1960. Macowanites pinicola A. H. Sm., Mycologia 55: 430-431. 1963. Macowanites roseipes (Singer & A. H. Sm.) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: 363. 1979. = Elasmomyces roseipes Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 64-65. 1960. Macowanites russuloides (Setch.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces russuloides Setch., J. Mycol. 13: 240. 1907. Macowanites setchellianus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 92-93. 1960. Macowanites stipitatus (H. A. Peters) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces stipitatus H. A. Peters, Mycologia 54: 112. 1962. Macowanites subolivaceus A. H. Sm., Mycologia 55: 432-433. 1963. 203 Macowanites subrosaceus A. H. Sm., Mycologia 55: 433-434. 1963. Macowanites tomentosa (J. W. Cribb) T. Lebel & Castellano, Mycologia 94 (in press) 2002. = Hydnangium tomentosum J. W. Cribb, Pap. Dept. Bot. Univ. Queensland 3: 2511958: = Martellia tomentosa (J. W. Cribb) A. H. Sm., Mycologia 54: 631. 1962. Macowanites vinicolor A. H. Sm., Mycologia 55: 434-435. 1963. Macowanites zellerianus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces zellerianus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 61-62. 1960. EXCLUDED SPECIES (INCLUDING NEW COMBINATIONS) Arcangeliella camphorata (Singer & A. H. Sm.) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: 365. 1979. = Elasmomyces camphoratus Singer & A. H. Sm.. Mem. Torrey Bot. Club 21: 62-63. 1960. Arcangeliella crichtonii (G. W. Beaton, Pegler & T. W. K. Young) T. Lebel & Castellano, Mycologia 94 (in press) 2002. = Cystangium crichtonii G. W. Beaton, Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 86: 181-182. 1986. Arcangeliella hepaticus (G. W. Beaton, Pegler & T. W. K. Young) T. Lebel & Castellano, Mycologia 94 (in press) 2002. = Elasmomyces hepaticus G. W. Beaton, Pegler & T. W. K. Young, Kew Bull. 39: 676-677. 1984b. Arcangeliella texta (J. W. Cribb) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: SOD BLOT: = Secotium sessile var. textum J. W. Cribb, Paps. Dept. Bot. Univ. Queensland 3: 108. 1956. = Elasmomyces texta (J. W. Cribb) A. H. Sm., Mycologia 54: 638. 1962. Hydnangium carneum Wallr. in A. Dietrich, Fl. Boruss. 7: 465. 1839. = Octaviania socialis Harkn., Proc. Cal. Acad. Sci. Ser. 3, 1: 252. 1899. = Arcangeliella socialis (Harkn.) Zeller & C. W. Dodge, Ann. Missouri Bot. Carle et09 e935: = Gymnomyces socialis (Harkn.) Singer & Sm., Mem. Torrey Bot. Club 21: 52. 1960. Octaviania borneensis Petri, Malpighia 14: 128. 1900. = Elasmomyces borneensis (Petri) Zeller & C. W. Dodge. Ann. Missouri Bot. Gard. 22: 370. 1935. = Octavianina borneensis (Petri) Pegler & T. W. K. Young, Trans. Brit. Mycol. b0Cs 72: 58081979. Octaviania lamingtonensis (J. W. Cribb) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Secotium lamingtonense J. W. Cribb, Pap. Dept. Bot. Univ. Queensland 3: 107. 1956. = Elasmomyces lamingtonensis (J. W. Cribb) A. H. Sm., Mycologia 54: 637. 1962. 204 = Octavianina lamingtonensis (J. W. Cribb) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: 380. 1979. Octaviania malaiensis (Corner & Hawker) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Elasmomyces malaiensis Corner & Hawker, Trans. Brit. Mycol. Soc. SONI2Z9IALISS:. = Octavianina malaiensis (Corner & Hawker) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72: 382. 1979. Russula subloculata Trappe, T. Lebel & Castellano, nom. nov. = Macowanites americanus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 88- 89. 1960, non Russula americana (Singer) Singer, Bull. Soc. Mycol. Fr. 55: 264. 1940. Zelleromyces albellus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nov. BASIONYM: Martellia albellus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 38-39. 1960. Zelleromyces alveolatus (Singer & A. H. Sm.) Trappe, T. Lebel & Castellano, comb. nob. BASIONYM: Gymnomyces alveolatus Singer & A. H. Sm., Mem. Torrey Bot. Club 21: 51-52. 1960. Zelleromyces australiensis (Berk.) Pegler & T. W. K. Young, Trans. Brit. Mycol. Soc. 72334 UZ1979, = Hydnangium australiense Berk., Trans. Linn. Soc. London Ser. 2, Bot. 2: 66. 1883. = Octaviania australiensis (Berk.) Berk. in Cooke, Handbook of Australian Fungi, p. 246. 1892. = Arcangeliella australiensis (Berk.) C. W. Dodge, Ann. Missouri Bot. Gard. 18: 463. 1931. = Martellia australiensis (Berk.) A. H. Sm., Mycologia 54: 633. 1962. = Octaviania alveolata Cooke & Massee in Cooke, Grevillea 16: 2. 1887. = Octavianina alveolata (Cooke & Massee) O. Kuntze, Rev. Gen. Plant. 3 (2): 501. 1898. = Hydnangium alveolatum (Cooke & Massee) Rodway, Paps. Proc. Roy. Soc. Tasmania 1919: 112. 1920. = Arcangeliella alveolata (Cooke & Massee) Zeller & C. W. Dodge, Ann. Missouri Bot. Gard. 22:365. 1935. = Martellia alveolata (Cooke & Massee) A. H. Sm., Mycologia 54: 630. 1962. = Hydnangium brisbanensis Berk.& Broom in Cooke, nom. illeg. Handbook of Australian Fungi, p. 247. 1892. = Octaviania brisbanensis (Berk.& Broom) G. Cunn., Proc. Linn. Soc.