t ¢ , J : in P ; ‘ 5 s J . my , Be « é - ’ 4 ¢ ‘ : : i é é oh » Pe re f \ bh fi \ HAY . & , i , le . ‘ . 4 J ‘ y py o ‘ f 4 \, . ' P 1 ALBERT R. MANN LIBRARY AT CORNELL UNIVERSITY CORNELL UNIVERSITY LIBRARY 924 063 069 623 DATE DUE OK. 600 M94 2- Vv, #7 1993 00 MNIA*VN Vi 4-7 m= MY COTAXON AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS Vol. XLVII April-June 1993 New and interesting Polypores from Mexico. ...... Leif Ryvarden and Gast6n Guzman 1 Glomales of Taiwan: III. A Compartative study of spore ontogeny in Sclerocystis ear eae, COneee ol aes rk Gea doo he ee le ewe Chi-Guang Wu —s25 Systematic and biological studies in the Balansieae and related anamorphs. III. Ascospore and macroconidial germination as a taxonomic criterion. Ryan A. Phelps and G. Morgan-Jones 41 Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian Pees Soi i eee M. Muntanola-Cvetkovic and A. G6mez-Bolea 59 Lectotypification of Ophiobolus trichellus (Dothideales, Ascomycetes). Christian Scheuer — 67 Taxomyces andreanae, a proposed new taxon for a bulbilliferous hyphomycete associated with Pacific Yew (Taxus brevifolia). Gary Strobel, Andrea Stierle, Don Stierle and W. W. Hess = 71 Two new South American species of Corticiaceae (Basidiomycetes) with amyloid RRR eke) SS ee Wee Kurt Hjortstam and Leif Ryvarden = 81 Dematiaceous Hyphomycetes on Freycinetia (Pandanaceae). 4. Cryptophiale. E. H. C. McKenzie and A. J. Kuthubutheen = 87 Genetics of a Tuber aestivum population (Ascomycotina, Tuberales). G. Pacioni, G. Frizzi, M. Miranda and C. Visca 93 New species in the lichen family Parmeliaceae (Ascomycotina) from Australia. John A. Elix 10] Taxonomic studies in the genus Mycosphaerella. 2. Notes on some additional Series OeCitisie On DIARsICHCCaG cS ee yy a we ke ks Michael Corlett 131 Development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy and observations on the fine structure of the conidium. J. Muthumary, J. A. Jayachandra and M. Bhagavathy Preetha 147 Hydnangium pila Pat., an older name for Martellia mediterranea Moreno, Galan ee RR CCCN a ee a has haste G. Moreno and R. Galan 157 Computer coding of strain features of the genus Rhizopus. Shung-Chang Jong and Candace McManus 161 Three new hyaloscyphaceous fungi from Norway and Greenland. Sigurd Olsen, John Haines and Sigmund Sivertsen —_177 Taxonomic history of the oomycete genus Thraustotheca. Will H. Blackwell and Martha J. Powell 183 [Contents continued overleaf] ISSN 0093-4666 MYXNAE 47:1-508 (1993) Published quarterly by MYCOTAXON, LTD., P. O. Box 264, Ithaca, NY 14851. For subscription details, availability in microfilm and microfiche, and availability of articles as tear sheets, see back cover. [Contents continued from front cover] Biostatistical characterization of the genus Oidium ................ Marco T. Ialongo Hypogeous mycorrhizal fungi of Spain. Isabel F. Alvarez, Javier Parladé, James M. Trappe and Michael A. Castellano Three xylariaceous fungi with scolecosporous conidia. Y..-M. Ju, Felipe San Martin Gonzalez and Jack D. Rogers Biscogniauxia and Camillea in Mexico Felipe San Martin Gonzalez and Jack D. Rogers New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp. nov., and its anamorph Kellermania yuccifoliorum, sp. nov. and Planistromella uniseptata, sp. nov., the teleomorph Of Kellermanid yuccipend. othe ace aie ota Annette W. Ramaley 4-O-Methyllividic acid, a new lichen depsidone. .. John A. Elix and Debra A. Venables Anaeromyces, and earlier name for Ruminomyces. Y. W. Ho, D. J. S. Barr, N. Abdullah, and S. Jalaludin A new species of Piromyces from the rumen of deer in Malaysia. Y. W. Ho, D. J. S. Barr, N. Abdullah, S. Jalaludin, and H. Kudo Type specimens of lichens and lichenicolous fungi in the Canadian Museum OF Nature (CANE) ea5 oie oo LE ere ek ae eee see Pak Yau Wong Neosartorya primulina, a new species of food-borne ascomycetes. Shun-ichi Udagawa, Noritsuma Toyazaki and Haruo Tsubouchi Studies on Boleiss SeCUOWLURIAE! All boas yy Memes Sean eu ei eo akg R. Treu Two new Crinipellinae (Tricholomataceae: Marasmiae) from South America. Roy E. Halling Septoria thymi sp. nov. from Bulgaria. . Ekaterina F. Sameva and Ganka G. Bakalova Three new species and a new variety of Pluteus from the United States. Partha Banerjee and Walter J. Sunberg World list of cetrariowschens. 2.9 aol hn ware ee Tiina Randlane and Andres Saag Two new species of Cristinia (Basidiomycotina, Aphyllophorales) and a survey OF ThE SOnus 0). eee Kurt Hjortstam and Helga Grosse-Brauckmann Comments on recent work on Ophiostoma and its synnematous anamorphs. H. P. Upadhyay Trematosphaeria pachycarpa and Herbampulla crassirostris gen. et spec. nov. (Ascomycetes): Ti taclar eae ine cette naan aren amet te C. Scheuer and A. Nograsek New or interesting lichenicolous fungi. 3. Karsteniomyces llimonae sp. nov. and Sclerococcum serusiauxii sp. nov. (Deuteromycotina). Montserrat Boqueras and Paul Diederich Amanita reidii - a new species from South Africa. Albert Eicker, J. V. Van Greuning and Derek A. Reid Phytophthora hibernalis and P. syringdé...........0 0c eee H. H. Ho and S. C. Jong A new Cortinarius from Southeastern (Fexas. 2s ce. eee Orson K. Miller Book Reviews. ssc 8 sis th noid Bias Sie Gee cee eee Cee te eer rene L. M. Kohn Notice: A new computer program for the identification of lichen SUDS(ATICES ..0 Wea seks Esther Mietzsch, H. Thorsten Lumbsch and John E. Elix Pthor TND EA oe asl ie eee & SS tae ini ec ane ome a dee ine ty eae rs er we INDEX to fungous and lichen taxa.” “ees tn oc a Ce cn a ie es, yy Errata, Publication Dates, MYCOTAXON Volumes 45 (corrected), 46............... Reviewers oe BD WPS. Oe. ER ee) lene! Tene he Leh eS [si ee fe) 10 ee eo le we, selma! ile te) rene A! (Gg he) wl Gt ce (@ Ghirennee-e (ee ie a 6) Se ie whe is. MY COTAXON AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS WEI INIGPe ALE i 19.93 ALBERT R. MANN LIBRARY JUN 2 9 1993 ITHACA, NY 14853 COMPLETE IN ONE VOLUME CONSISTING OF iv + 508 PAGES INCLUDING FIGURES EDITOR-IN-CHIEF JEAN BOISE CARGILL Harvard University Herbaria 22 Divinity Avenue, Cambridge, MA 02138, USA ASSOCIATE EDITORS LINDA M. KOHN Book Review Editor Botany Department, University of Toronto — Erindale Mississauga, Ontario LSL 1C6, Canada GREGOIRE L. HENNEBERT French Language Editor Laboratoire de Mycologie systématique et appliquée Université Catholique de Louvain, B-1348 Louvain-la-Neuve, Belgium ROBERT DIRIG Index Editor Bailey Hortorium, Mann Library Building Cornell University, Ithaca, NY 14853, USA EDITORIAL ADVISORY BOARD OVE E. ERIKSSON, Umea, Sweden (1990-93, Chm.) GREGOIRE L. HENNEBERT, Louvain-la-Neuve, Belgium (1990-96) JAMES W. KIMBROUGH, Gainesville, Florida (1992-97) RONALD H. PETERSEN, Knoxville, Tennessee (1990-94) JACK D. ROGERS, Pullman, Washington (1993-98) AMY Y. ROSSMAN, Beltsville, Maryland (1990-95) Published by MYCOTAXON, LTD., P.O. BOX 264 ITHACA, NY 14851-0264, USA Printed in the United States of America Table of Contents, Volume Forty-seven April-June 1993 New and interesting Polypores from Mexico .. . Leif Ryvarden and Gaston Guzman Glomales of Taiwan: III. A Compartative study of spore ontogeny in Sclerocystis (eslOMaACCAGs, CSIOMALCS ) aa Nira tenis andere ae Pliers bee, Chi-Guang Wu Systematic and biological studies in the Balansieae and related anamorphs. II. Ascospore and macroconidial germination as a taxonomic criterion. Ryan A. Phelps and G. Morgan-Jones Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian BOUIN SU Ab we: Peua Pr iene, M. Muntanola-Cvetkovic and A. G6mez-Bolea Lectotypification of Ophiobolus trichellus (Dothideales, Ascomycetes). Christian Scheuer Taxomyces andreanae, a proposed new taxon for a bulbilliferous hyphomycete associated with Pacific Yew (Taxus brevifolia). Gary Strobel, Andrea Stierle, Don Stierle and W. W. Hess Two new South American species of Corticiaceae (Basidiomycetes) with amyloid SDOLC Srunteeric Malet Alec haha deel ee oe Kurt Hjortstam and Leif Ryvarden Dematiaceous Hyphomycetes on Freycinetia (Pandanaceae). 4. Cryptophiale. E. H. C. McKenzie and A. J. Kuthubutheen Genetics of a Tuber aestivum population (Ascomycotina, Tuberales). G. Pacioni, G. Frizzi, M. Miranda and C. Visca New species in the lichen family Parmeliaceae (Ascomycotina) from Australia. John A. Elix Taxonomic studies in the genus Mycosphaerella. 2. Notes on some additional species DCCUUIAEFONL EST ASSICACCACI Gnas ahatner ail an ements Mee Ae A Michael Corlett Development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy and observations on the fine structure of the conidium. J. Muthumary, J. A. Jayachandra and M. Bhagavathy Preetha Hydnangium pila Pat., an older name for Martellia mediterranea Moreno, Galan SSN LONIECEIN Gee Pe veeae Ch tee econ Meee gree te it G. Moreno and R. Galan Computer coding of strain features of the genus Rhizopus. Shung-Chang Jong and Candace McManus Three new hyaloscyphaceous fungi from Norway and Greenland. Sigurd Olsen, John Haines and Sigmund Sivertsen Taxonomic history of the oomycete genus Thraustotheca. Will H. Blackwell and Martha J. Powell Biostatistical characterization of the genus Oidium.............. Marco T. Ialongo my pogeous mycorrhizal fungi of Spain..................... Isabel F. Alvarez, Javier Parladé, James M. Trappe and Michael A. Castellano Three xylariaceous fungi with scolecosporous conidia. Y..-M. Ju, Felipe San Martin Gonzalez and Jack D. Rogers Biscogniauxia and Camillea in Mexico Felipe San Martin Gonzalez and Jack D. Rogers New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp. nov., and its anamorph Kellermania yuccifoliorum, sp. nov. and Planistromella uniseptata, sp. nov., the teleomorph of Kellermania yuccigena................. Annette W. Ramaley itt 25 4] oy 67 701 101 iv 4-O-Methyllividic acid, a new lichen depsidone. John A. Elix and Debra A. Venables Anaeromyces, and earlier name for Ruminomyces. Y. W. Ho, D. J. S. Barr, N. Abdullah, and S. Jalaludin A new species of Piromyces from the rumen of deer in Malaysia. Y. W. Ho, D. J. S. Barr, N. Abdullah, S. Jalaludin, and H. Kudo Type specimens of lichens and lichenicolous fungi in the Canadian Museum ob Nature (CANT) Sante cre mee ae Rien Mt Goel arises ane Pak Yau Wong Neosartorya primulina, a new species of food-borne ascomycetes. Shun-ichi Udagawa, Noritsuma Toyazaki and Haruo Tsubouchi Studies’on Boletus \SectiOndUunidii anne eae aoe ke eis SoA i ed ete R. Treu Two new Crinipellinae (Tricholomataceae: Marasmiae) from South America. Roy E. Halling Septoria thymi sp. nov. from Bulgaria. Ekaterina F. Sameva and Ganka G. Bakalova Three new species and a new variety of Pluteus from the United States. Partha Banerjee and Walter J. Sunberg World hst of céetranoid lichenss iano es eee Tiina Randlane and Andres Saag Two new species of Cristinia (Basidiomycotina, Aphyllophorales) and a survey Of the SenuUS aus oer We enc Ann Kurt Hjortstam and Helga Grosse-Brauckmann Comments on recent work on Ophiostoma and its synnematous anamorphs. H. P. Upadhyay Trematosphaeria pachycarpa and Herbampulla crassirostris gen. et spec. nov. C(ASCOMYCELES) 7 AMR nO da erie een Tak C. Scheuer and A. Nograsek New or interesting lichenicolous fungi. 3. Karsteniomyces llimonae sp. nov. and Sclerococcum serusiauxii sp. nov. (Deuteromycotina). Montserrat Boqueras and Paul Diederich Amanita reidii - a new species from South Africa. Albert Eicker, J. V. Van Greuning and Derek A. Reid Phytophthora hibernalis and P. syringae............. H. H. Ho and S. C. Jong A new.Cortinarius from Southeastern Texas .....-.......0..0. Orson K. Miller Book Revie wseanee tea ee hee trie eee Stee On at ot ft aeeaitane L. M. Kohn Notice: A new computer program for the identification of lichen substances. Esther Mietzsch, H. Thorsten Lumbsch and John E. Elix Author, END Exar ids See dancin Sai oto eae nate is oe, ae geen fer TUG W ihre ean Sete ae INDE X*torTangous and miche pn taxkarge eager men, er hee eee er oon eee ee Errata, Publication Dates, MYCOTAXON Volumes 45 (corrected), 46............... REVICWETS "GRA ue OP ee, Bararnd Gee ae BL ies Cee diel! SMR Oia re Rd Oem ana Me MY COTAXON Volume XLVI, pp. 1-23 April-June 1993 NEW AND INTERESTING POLYPORES FROM MEXICO by Leif Ryvarden UNIVERSITY OF OSLO Box 1045, BLINDERN N-0316 OSLO 3 NORWAY and Gaston Guzman INSTITUTO DE ECOLOGIA APARTADO POSTAL 63 XALAPA, VERACRUZ 91000 MEXICO SUMMARY Forty three species of polypores from Mexico studied by the authors since 1989 are discussed, of which 19 are new records for the Mexican mycoflora and 23 are reported from new localities. Most of the studied specimens are from tropical vegetation, except a few from temperate regions or from coniferous forests. The studied species are reported from the following states of the country: Baja California, Distrito Federal, Jalisco, Morelos, Quintana Roo, Veracruz and Yucatan. RESUMEN Se discuten 43 especies de Poliporaceos de México estudiadas por los autores desde 1989, de las cuales 19 son nuevos registros para la micoflora mexicana y 23 se presentan de nuevas localidades. La mayoria de los especimenes proceden de vegetacién tropical, excepto unos pocos que son de zonas templadas 0 de bosques de coniferas. Las especies consideradas se adscriben a las entidades federativas: Baja California, Distrito Federal, Jalisco, Morelos, Quintana Roo, Veracruz y Yucatan. INTRODUCTION The polypores of Mexico have been studied during the last century, through the works of Kickx (1841) to Saccardo (1888), and in the present centuary through several contributions by Murrill (Murrill, 1905, 1907-1908, 1912, 1915), and modern revisions of Anell and Guzman (1988), Ojeda-L6pez et a/. (1986), Valenzuela and Nava (1991), Valenzuela and Chacdon-Jiménez (1991) and Vazquez and Guzman-Davalos (1991). Nevertheless the knowledge of the group is still scanty, as shown by the revision on some collections in XAL Herbarium. Among these the authors found 19 new records for the country, and 23 from new localities not only from the tropics where the mycological studies are few but also from temperate regions, where the mycology has a good development. The present paper is based on the study of more than 70 specimens of polypores of XAL Herbarium (mostly duplicates from ENCB), with duplicates in O. All the material was studied by the authors between 1989-1991. The nomenclature following in this paper, in according to Gilbertson and Ryvarden (1986-1987). CONSIDERED SPECIES A total of 43 species of polypores were studied, of which 19 are new records from Mexico, and 23 are reported from new localities. Tables 1 and 2 show the studied species. Phylloporia spathulata is considered in connection with a discussion of P. veraecrucis from a new locality. Albatrellus subrubescens (Murr.) Pouz. This is an unusual North American species, only known from Arizona and Florida in the U.S.A., and recently reported from Mexico, from the States of Michoacan (Diaz-Barriga et al., 1988) and Chiapas, Mexico, Nuevo Leon and Veracruz (Valenzuela & Nava, 1991). The studied material is characterized by the basidiocarps with yellowish, smooth to squamulose pileus, pore surface greenish yellow with irregular reddish-orange tints, context cream colored and unchanging, and by the absence of clamp connections and amyloid spores. 3 Material studied. STATE OF VERACRUZ, Region E of Cofre de Perote, S of Tembladeras, El Revolcadero, Pinus forests, Villarreal 1424. Amauroderma camerarium (Berk.) Furt. It is a widespread fungus in the neotropics (Furtado, 1981), and recognized by its pale yellowish context. Welden et a/. (1979) and Anell & Guzman (1987, 1988), reported this fungus from the State of Veracruz. | Material studied. STATE OF QUINTANA ROO, road to the sea from the road Puerto Morelos to Tulum, near the deviation to Vallarta, tropical forest, Lopez 1848. Amylosporus campbellii (Berk.) Ryv. This is a new record from Mexico. It is a pantropical species; in America known from southern United States, besides of Bermuda (Gilbertson & Ryvarden, 1986-1987), and the Caribbean zone. Murrill (1915) described this species as Tyromyces graminicola Murr. Usually the species grows in clusters of semistipitate to fanshaped basidiocarps on the ground in subtropical or tropical vegetation. The pileus is whitish to pale brownish, grayish brown or cinnamon brown, irregularly and finely tomentose. The poroid surface is almost concolorous with the pileus. Microscopically the small warted and strongly amyloid spores are characteristic and point to relationships with the Hericiaceae where these features are common. Material studied. STATE OF MORELOS, road Cuernavaca to Cuautla, Canon de Lobos, km 14, tropical forest, Guzman 6777. Anomoporia albolutescens (Rom.) Pouzar This is a temperate species, known from Europe and North America through the coniferous forests (Gilbertson & Ryvarden, 1986-1987). The Mexican collection is the first record to Mexico. The thin resupinate, loosely attached, rhizomorphic and pale yellowish, yellow or orange yellowish basidiocarps are typical in this fungus. Welden et al. (1979) reported A. myceliosa (Peck) Pouzar (as Poria myceliosa Peck) from Acayucan, Ver. a species with white basidiocarp. Material studied. STATE OF VERACRUZ, Region E of Cofre de Perote, S of Tembladeras, El Revolcadero, Pinus forest, Montoya 318. 4 Antrodia gossypina (Speg.) Ryv. A new record from Mexico. This species has a wide distribution in temperate zones, mostly in coniferous forests (Gilbertson & Ryvarden, 1986-1987). It is recognized by its resupinate thick, soft to cottony, white to whitish or whitish rose basidiocarp, with a rhizomorphic margin and by the ellipsoid spores, 4.5-6 x 2-3 um. Material studied. STATE OF VERACRUZ, Municipio de Totula, El Mirador, subtropical forest with Quercus, Ventura 16294. Antrodia malicola (Berk. & Curt.) Donk This species was known from Mexico (as Trametes malicola Berk. & Curt.) from a report by Sharp (1948), without specific locality. It is widespread in warmer or temperate zones, common in North America, but rare in Europe and Africa. The basidiocarps are resupinate, effused-reflexed or sessile, forming nodulose to imbricate pilei, whitish or pale woody color, with tomentose to agglutinated glabrous surface . Material studied. STATE OF VERACRUZ, Los Tuxtlas Region, near El Jical, tropical forest, Guzman 19702. Antrodia vaillantii (Fr.) Ryv. Lowe (1966) reported this species as Poria vaillantii (Fr.) Lowe from Mexico, but without specific locality. It is a widespread and conspicuous crusty fungus, very close to A. gossypina (see that). Many reports of this fungus are based on sterile specimens having a well developed rhizomorphic margin, common also in that species. A. vaillantii has spores (5-8 x 3-4 um) larger than A. gossypina. Material studied. STATE OF YUCATAN, between Xkantam and Progreso, plantation of henequen (Agave fourcroydes Lem.), Guzman 21365. Antrodiella angulatopora Ryv. This species was recently described from Venezuela (Ryvarden, 1987), and this is the second report of the fungus. The basidiocarp is sessile, broadly attached, imbricate and semicircular; the upper surface is whitish to ochraceous with narrow orangish yellow zones, and the pore surface is pale yellowish in dry conditions. Material studied. STATE OF VERACRUZ, road Xalapa to 5 Tlapacoyan, near Atzalan, La Calavera, subtropical forest with Quercus and Cyathea, Ventura 12958. Antrodiella versicutis (Berk. & Curtis) Gilbn. & Ryv. This is a neotropical fungus, first reported from Mexico. The allantoid spores separate this species from others of the genus. The studied specimens have sessile basidiocarps or short or long Stipitate, fanshaped pileus with the upper surface glabrous, ochraceous to whitish yellow; pore surface almost concolorous with the pileus, pores 6-8 per mm. Murrill (1907-1908) described this species as Coriolus ochrotinctellus Murr. \t is very interesting to observe the apparently abundance of A. versicutis in the tropical vegetation of the eastern part of Peninsula of Yucatan, and the absence of records from other tropical regions of Mexico, which suggest that this is a Caribbean and South American species, as those discussed by Guzman (1986). Material studied. STATE OF QUINTANA ROO, road Felipe Carrillo Puerto to Vigia Chico, 20 km from Felipe Carrillo Puerto, tropical forest, Guzman 20777. Road Puerto Morelos to Tulum, near the deviation to Vallarta, tropical forest, Guzman 23830; Lopez 1815; 1882. Road Coba to Nueva Xcan, near the deviation to Sajaca-Chen, tropical forest, Lopez 1925. N of Reserva de la Biésfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23853. Aurificaria luteo-umbrina (Rom.) Reid Murrill (1907-1908) described this species as Pyroporellus subblinteus Murr. from Mexico (from the coasts of the State of Sonora). A. luteo-umbrina is common in America, from the SE of the U.S.A. to Brazil, and in Australia, but unknown from Africa (Ryvarden & Johansen, 1980; Gilbertson & Ryvarden, 1986-1987). The basidiocarps are sessile on logs, rusty to yellowish brown and brittle, up to 80 mm wide; the context is bright yellowish brown, instantaneously staining red with KOH; the pore surface turns brownish red or cinnamon brown when touch in fresh conditions. The lack of setae and the globose to subglobose spores, that stain olivaceous brown in KOH define the species. Material studied. STATE OF QUINTANA ROO, N of Reserva de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23862. 6 Coriolopsis floccosa (Jungh.) Ryv. This species was reported by Ojeda-Lopez et al. (1986) from the State of Guanajuato. It is a pantropical fungus, apparently more common in the old tropics than in the neotropics. The basidiocarps are sessile, broadly attached and semicircular, with brown to grayish pileus, with black concentric zones in dry specimens. It may be confused with C. rigida (see that); however this latter species is more thinner and flexible, and with pale brown pore surface. Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Tulum, near the deviation to Vallarta, tropical forest, Guzman 23950. Coriolopsis rigida (Berk. & Mont.) Murr. Murrill (1907-1908, 1912), Welden & Lemke (1961), Castillo et al. (1969), Castillo & Guzman (1970), Guzman (1972) and Anell & Guzman (1987) reported this species from the State of Nuevo Leon, Colima and Veracruz. This is a very common fungus in the neotropics, but it is widespread even in the Eastern U.S.A. The basidiocarp is variable in habit, from effused or connate to imbricate clusters (See discussion of C. floccosa). Material studied. STATE OF YUCATAN, 5 km S of Xochel, road to Huhi, near Sacabah, tropical forest, Guzman 23746. Daedalea microsticta Cooke Figs. 4-5 This is a widespread neotropical fungus. It was known from the States of Colima and Veracruz (Murrill, 1912; Guzman, 1972, Anell & Guzman, 1987). Here it is recorded from the States of Nuevo Leon and Quintana Roo. The basidiocarp is applanate, more or less thick, sessile, light color; the pileus is ivory color in dry conditions, and reach more than 120 mm wide. Some studied specimens were found in ENCB as Daedalea elegans Spreng. : Fr., a neotropical fungus widespread in Mexico, that differs from D. microsticta for its white funnel shapped basidiocarp with a thin pileus. Material studied. STATE OF NUEVO LEON, Municipio Santiago, La Chancaca, Castillo 211; El Cercado, Sepulveda 19: Ramirez s.n., all in subtropical forests. STATE OF QUINTANA ROO, road Vigia Chico to Felipe Carrillo Puerto, Rancho Yaras, tropical forest, Guzman 23934. vf Datronia caperata (Berk.) Ryv. A pantropical species reported from Mexico as Polyporus caperatus Berk., Coriolopsis caperata (Berk.) Murr., and C. caperatiformis Murr., from the States of Campeche, Colima, Hidalgo, Quintana Roo, Tabasco and Veracruz by Murrill (1912, 1915), Guzman (1972, 1975), Welden et a/. (1979), and Anell & Guzman (1987). The sessile, thin and flexible basidiocarp is very variable in form, from applanate, dimidiate, conchate to flabelliform, up to 50 mm wide. In wet and shady places the pileus becomes almost smooth with a tomentum restricted only to certain zones, but in drier and sunny localities, it is covered with a characteristic dark tomentum in distinct zones and below the tomentum there are well defined black zones. The pore surface is white, but in old and dried specimens is pale brownish, grayish brown to deep brown; in fresh conditions turns brown when touched. Material studied. STATE OF MORELOS, highway Mexico to Cuernavaca, near the deviation to Cuautla, subtropical forest with Quercus, Guzman 18114. STATE OF QUINTANA ROO, N of Reserva de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23869. | Diplomitoporus lenis (Karst.) Gilobn. & Ryv. A new report from Mexico, this is a common circumglobal species in subtropical and temperate areas (Gilbertson & Ryvarden, 1986-1987), but rare in the tropics. Ryvarden and Johansen (1980) reported only one specimen from Africa [as Antrodia lenis (Karst) Ryv.]. The spores are small, allantoid to Iunate and the basidiocarp is resupinate, white to whitish, forming large crusts under the logs. Material studied. STATE OF QUINTANA ROO, N of Reserva de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23871. Flavodon flavus (KI.) Ryv. This species has hitherto not been reported from America. It is very common in Africa, Asia and Australia (Ryvarden & Johansen, 1980) and otherwise shows the relationships between tropical regions of Mexico and Africa, observed by Guzman et al. (1989). The intense yellow to orangish yellow basidiocarps are very conspicuous. The hymenophore is hydnoid to iripiciform and the pileus turns red with KOH. Material studied. STATE OF QUINTANA ROO, road Coba to Nueva Xcan, near the deviation to Sajaca-chen, Lopez, 1941; 1947. Fomitopsis melliae (Underw.) Gilbn. & Ryv. pals bates It is a new report from Mexico. This is a North American species (Gilbertson & Ryvarden, 1986-1987), and the collections from Mexico represent the southern limit so far known today. The basidiocarp is up to 80 mm wide, thin to thick, sessile with a pileus surface ivory to tan or cinereous color, frequently glabrous and azonate (the collection from Quintana Roo is faintly zonate and thin); the pore surface is whitish to ochraceous and the context is whitish to pale buff. Material studied. STATE OF QUINTANA ROO, N of Reserva de la Biésfera de Sialan Kalan, Chuyaxchen, Archeological zone of Mugil, tropical forest, Guzman 24147. STATE OF VERACRUZ, N of Xalapa, Rancho Lucas Martin, subtropical disturbed vegetation, Guzman 1915 (this specimen was reported by Guzman, 1963-A, as Daedalea ambigua Berk.). Ganoderma coffeatum (Berk.) Furt. Figs. 6-7 This species also placed in Amauroderma by Murrill or Humphreya by Steyaert (1972), was recently reported from Mexico by Cappello & Hernandez (1990), Vazquez & Guzman-Davalos (1991) and Valenzuela & Chac6on-Jiménez (1991) from the States of Jalisco, Tabasco and Tamaulipas. It is a neotropical and widespread fungus, very common in South America. Its light colored context, the brown ferrugineous velutinous pileus and stipe, the whitish pore surface and the coarsely ornamented spores with a prominent reticulated pattern of coherent ridges, are distinct good features of this species. Macroscopically this fungus resembles an Amauroderma. Here it is reported for first time from the States of Quintana Roo and Veracruz. Material Studied. STATE OF QUINTANA ROO, N of Reserva de la Biosfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23902. STATE OF VERACRUZ, Uxpanapa region, tropical rain forest, Guzman 30467. Grammothele fuligo (B. & Br.) Ryv. This is a tropical corticoid fungus, reported as Porogramme fuligo (B. & Br.) Pat. by Lowe (1963) from Mexico without specifing locality. The species occurs exclusively on dead leaves or stems of 9 monocotyledoneous, specially on all types of palms. It is easy to overloook due to its resupinate basidiocarp with dark gray bluish paints. Material studied. STATE OF QUINTANA ROO, road to Xcan, 15 km from Coba, tropical forest, Lopez 1913. Incrustoporia nivea (Jungh.) Ryv. This is a circumglobal temperate species. It was reported from Mexico by Sharp (1948) as Polyporus semipileatus Peck, without specifing locality. Murrill (1912) reported it from the State of Morelos as Tyromyces semipileatus (Peck) Murr. It is known in North America as Skeletocutis nivea (Jungh.) Keller (Gilbertson & Ryvarden, 1986-1987). It presents white to cream color annual basidiocarp, effused-reflexed to resupinate. Material studied. STATE OF BAJA CALIFORNIA, road Ensenada to Ojos Negros, NE of Agua Viva, Quercus forest, Guzman 24302-A. Inonotus hispidus (Bull. : Fr.) Karst. This is a common fungus in the United States (Gilbertson & Ryvarden, 1986-1987), characterized by its strongly hispid upper surface and the pigmented spores. The species was known from Mexico from the States of Baja California, Hidalgo, Mexico, Michoacan, Nuevo Leon and Veracruz (Bandala et a/., 1992). In the present paper it is reported from Jalisco. Material studied. STATE OF JALISCO, 11 km NE of Tamazula, Rio Contla, tropical disturbed vegetation, Guzman 8593. Inonotus munzii (Lloyd) Gilbn. A subtropical to xerophitic warm temperate American fungus, easy to recognize from the other species of the genus by the branched setigerous elements of the pileus and lack of setae. The basidiocarps are sessile, frequently ungulate and bright yellowish brown to reddish brown or cinnamon brown. The species was first reported from Mexico by Guzman (1963-A, -B) in subdesert lands from the north to the central regions of the country. Castillo & Guzman (1970) reported it from the State of Nuevo Leon. It is reported here for first time from Distrito Federal. Material studied. DISTRITO FEDERAL, Mexico City, Santo Tomas Colonia, Campus of the National School of Biological Sciences, on a living Schinus molle L., Fanti 74. 10 Junghuhnia nitida (Pers. : Fr.) Ryv. This species was reported from the State of Oaxaca by Welden et al. (1979). It is a cosmopolitan fungus, recorded from Northern Norway to Australia and New Zealand. It grows on any type of dead wood of angiosperms. Curiously in Mexico it is only known from tropical regions. The pinkish-cinnamon color of the pore surface and the so common thick-walled incrusted cystidia are characteristic features of this species (Gilbertson & Ryvarden, 1986- 1987). Material studied. STATE OF QUINTANA ROO, near the road Puerto Morelos to Felipe Carrillo Puerto, Archeological zone of Chunyaxche, tropical forest, Guzman 20646. STATE OF VERACRUZ, Los Tuxtlas Region, 3 km S of Montepio, tropical rain forest, Guzman 7257. Microporellus obovatus (Jungh.) Ryv. This is a pantropical species reported from Mexico by Cappello & Hernandez (1990), Vazquez & Guzman-Davalos (1991) and Valenzuela & Chacon-Jiménez (1991) from the States of Jalisco, Tabasco and Tamaulipas. The white to yellowish orange or straw color, tomentose to velvety striate, round to reniform or spathulate pileus, sometimes radially striate, laterally stipitate, with a white to concolorous pore surface and with very small pores (6-8 per mm) are features that distinguish this peculiar species (Gilbertson & Ryvarden, 1986-1987). Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Vallarta, San Matilde, tropical forest, Guzman 20539. State of Veracruz, Municipio de Totutla, El Mirador, subtropical forest, Ventura 17324. Los Tuxtlas Region, 7 km S of Montepio, Biological Station of the University of Mexico, tropical rain forest, Guzman 7160. 2 km SW of Xalapa, near Rio Coapexpan, subtropical forest with Quercus and Liquidambar, Bandala 227. Oligoporus fragilis (Fr.) Gilobn. & Ryv. A new report for the State of Morelos. This species was only known from the States of Guanajuato and Michoacan (Cifuentes et al., 1990; Ojeda-Lopez et a/., 1986) [in both cases as Tyromyces fragilis (Fr.) Donk]. It is a boreal fungus following pine trees in high altitudes. It is easy to recognize by the white to buff basidiocarps that turn reddish brown when bruised or dried. 11 Material studied. STATE OF MORELOS, Lagunas de Zempoala National Park, near the principal lagoon, Pinus forest, Guzman 3174. Perenniporia ohiensis (Berk.) Ryv. Welden & Lemke (1961) and Castillo et a/., (1969) reported this fungus as Fomes ohiensis (Berk.) Murr. from the State of Nuevo Leon. The large (up to 13 um long) thick-walled, dextrinoid spores are the principal features of this American species, as well as its small effused-reflexed or ungulate basidiocarps, with pileus whitish or ivory to brown or blackish, glabrous and azonate or fairy zonate, and almost concolorous with the pore surface and context (Gilbertson & Ryvarden, 1986-1987). Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Tulum, near the deviation to Vallarta, Botanical Garden of CIQRO, tropical vegetation, Guzman 23788. N of Reserva de la Biosfera de Sialan Kalan, Archeological zone of Mogil, tropical forest, Guzman 23872. STATE OF YUCATAN, 5 km S of Xocchel road to Hohi, near Sacabah, tropical forest, Guzman 23753. 5 km S of Conkal, road to Telchac, tropical forest, Guzman 23595. Perenniporia tephropora (Mont.) Ryv. A new record from Mexico. It is a pantropical species, known in the State of Louisiana (U.S.A.), the Caribbean region and Africa (Gilbertson & Ryvarden, 1986-1987; Ryvarden & Johansen, 1980), and considered as Loweporus by Ryvarden or Fomitopsis cubensis by Murrill. It is characterized by its basidiocarps, which are resupinate, large and thick, crustose and perennial, grayish to dark ochraceous or milk coffee color to ferrugineous brown. The basidiocarps frequently grow on vertical surfaces of the trunks. Material studied. STATE OF YUCATAN, road Merida to Valladolid, 3 km from Tixpeual, tropical forest, Guzman 23636. Phellinus chryseus (Lév.) Ryv. A new record from Mexico. It is a rare neotropical species, Originally described from Colombia (Ryvarden, 1981; Ryvarden & Johansen, 1980). The fairly large setae and small hyaline dropshaped basidiospores, define the species, besides its typical brown ferrugineous context and pore surface, as well as the rimose and zonate grayish black pileus. The Mexican collection represents the northern limit so far known to this fungus. The studied specimen 12 was growing in a disturbed tropical vegetation. Material studied, STATE OF JALISCO, Tamazula region, near Agua Caliente, tropical deciduous forest, Guzman 12486. Phellinus contiguus (Fr.) Pat. It is a widespread species in the tropics and warmer rare of Europe (Gilbertson & Ryvarden, 1986-1987). The large hyaline and ellipsoid spores and tramal setae characterize the species. The basidiocarp is perennial and resupinate, forming a thick, rusty brown crust below the logs. It is reported for first time from Mexico from tropical forests of the Peninsula of Yucatan. Material studied. STATE OF QUINTANA ROO, N of Reserva de la Bidsfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23882. Phellinus ferrugineo-velutinus (Henn.) Ryv. This is a rare resupinate neotropical species, new recorded from Mexico, and characterized by small hyaline spores and scattered setae (Gilbertson & Ryvarden, 1986-1987). Probably overlooked as other resupinate Phellinus species, which are often neglected because of the considerable difficculties presented in their identification. Ph. ferrugineo-velutinus forms large and thick, ferrugineous brown crusts under the logs, with a conspicous mustard yellow color margin. Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Tulum, near the deviation to Vallarta, road to the sea, tropical forest, Lopez 1849. Phellinus grenadensis (Murr.) Ryv. New record from Mexico. It is a wide-spread species in the Caribbean zone and Central America. It belongs to the. complex group of pileate species without setae and rather uniformely ellipsoid spores (Gilbertson & Ryvarden, 1986-1987). The studied specimen has a small basidiocarp, no more than 20 mm wide, subungulate, rusty brown and with a crustose pileus. Material studied. STATE OF QUINTANA ROO, Municipio Lazaro Cardenas, S of Chinquilla, tropical disturbed vegetation in a grassland, Guzman 20657. Phellinus rhytiphloeus (Mont.) Ryv. This is a rare tropical species, previously known only from 13 Brazil (Ryvarden & Johansen, 1980). It has solitary, dimidiate, thick, applanate and semicircular basidiocarps, with the typical brown cinnamon color. When cut it presents a distinct black line below the pileus. It is reported for first time from Mexico. Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Tulum, near Puerto Morelos, tropical forest, Lopez 1319. Phellinus rimosus (Berk.) Pilat Fig. 1 This is a complex species with a widespread distribution in the tropical and subtropical regions of the world. It grows in Africa (Ryvarden & Johansen, 1980), but its presense in the U.S.A. is doubtful (Gilbertson & Ryvarden, 1987). It was reported from Mexico as Fomes, Cooke, from several temperate and subtropical localities (Bandala et a/., 1992). It is reported here for first time from Yucatan. The perennial, ungulate basidiocarp, with brownish to blackish and tomentose to rimose pileus, yellowish-brown pore surface and lack of setae are characteristic in this species. Material studied. STATE OF YUCATAN, Municipio Progreso, between Chelen and Chuburna, tropical vegetation, Guzman 21373-B. Phylloporia chrysita (Berk.) Ryv. A pantropical species, recorded from Mexico from the State of Tamaulipas (Valenzuela & Chacon-Jiménez, 1991) and from the Uxpana region in the State of Veracruz (Welden et a/., (1979). In the presented paper it is reported from a new locality from the State of Veracruz. The sessile, thick, rusty-brown basidiocarps, with the pileus covered by a thick and velvety spongy compressed tomentum, as well as the thin context separated by a dark line from the tomentum, are good features that define the fungus and separate it from Ph. frutica (Berk. & Curt.) Ryv., also known from Mexico as parasitic on several tropical trees (Guzman, 1977). Material studied. STATE OF VERACRUZ, Municipio Minatitlan, Rio Coachapa, El Remolino, tropical forest, Pérez Ortiz 894. Phylloporia spathulata (Hook.) Ryv. It is a pantropical species, known as Polyporus by Cooke, Coltricia by Murrill and Polystictus s. auct., rather common although often overlooked due to its small size and dark brown color basidiocarps, growing on the ground often among leaves and debris inside the tropical forests. It seems to be a parasitic fungus, 14 at least specimens from Colombia were attached to living roots, and others from Mexico present a long pseudorrhiza. This fungus was reported from Mexico from the States of Quintana Roo and Yucatan by Guzman (1983) and from Veracruz by Guzman and Guzman-Davalos (1984). The species is considered here, in spite of that the studied material is from the same above regions of the country, because it is closely related to Ph. veraecrucis (see below), which was considered as synonym, and actually as an independent species; both species have golden- yellow to cinnamon brown stipitated basidiocarps. Ph. spathulata has pileus 10-60 mm broad and spores 3-4 x 2-3 um, while Ph. veraecrucis presents pileus up to 100 mm wide and spores 4-4.5 X 3-3.5 um, as shown in the study of the type at K. Material studied. STATE OF QUINTANA ROO, road Escarcega to Chetumal, 10 km N from Tomas Garrido, Guzman 20910; 23884. Road Puerto Morelos to Tulum, near the deviation to Vallarta, Guzman 21003; 21065; 23958; Lopez 1826; 1831. STATE OF YUCATAN, road Merida to Valladolid, between Ticopo and Tahnek, Guzman 23680. 5 km S of Xocchel, near Sacabah, Guzman 23694; 23710 (all the collections from tropical forests). Phylloporia veraecrucis (Berk. ex Sacc.) Ryv. This is an interesting fungus described as Polyporus veraecrucis Berk. from the State of Veracruz in the last century (Berkeley, 1867), based in a collection on roots of a tropical tree in 1854. It was validated and redescribed by Saccardo (1888). Murrill (1905) studied the type at K and considered the species as Phaeolopsis veraecrucis (Berk.) Murr. Ryvarden & Johansen (1980) considered Ph. veraecrucis as synonym of Ph. spathulata (see that), but recently, Ryvarden (1991) recognized this species and presented a modern description based on a study of the type. In the present paper this fungus is reported for the first time from the State of Quintana Roo. Material studied. STATE OF QUINTANA ROO, Cozumel region, Coba, tropical rain forest Guzman 20729. Piptoporus soloniensis (Dub. : Fr.) Pil. A new record from Mexico. This is a rare species distributed in the SW of the United States (Gilbertson & Ryvarden, 1986-1987). The Mexican collection seems to be the southern limit in the continent. The basidiocarp is similar to Laetiporus sulphureus 15 (Bull. : Fr.) Murr. (Polyporus pseudosulphureus Long is a synonym of P. soloniensis), but the discussed species has not the bright orange and yellow sulphur coloration. The clamp connections in the generative hyphae are a distinct feature. Material studied. STATE OF VERACRUZ, road Xalapa to Tlapacoyan, Municipio Atzalan, La Calavera, subtropical forest with Quercus, Ventura 17315 (Anell & Guzman, 1988, reported this specimen as Laetiporus sulphureus). Polyporus varius Fr. A circumglobal fungus specially common in temperate zones, but scattered in the tropics (Ryvarden & Johansen, 1980, did not report this species from Africa). The stipitate light coloured basidiocarps with a black stipe make it rather distinct in the genus except for P. elegans Bull. : Fr., that differs in the lack of the striations of the pileus. The young specimens present light coloured basal stipe as most of the studied specimens. This species has been recorded from the States of Chiapas, Michoacan, Mexico Tamaulipas and Veracruz (Bandala et a/., 1992). Here it is reported for first time from Morelos, Quintana Roo and Yucatan. Material studied. STATE OF MORELOS, IV Mushroom Exposition of Morelos, Guzman 21911. STATE OF QUINTANA ROO, N of Reserva de la Bidésfera de Sialan Kalan, Archeological zone of Mugil, tropical forest, Guzman 23873. Road Puerto Morelos to Tulum, near the deviation to Vallarta, tropical forest, Guzman 21015; 21077. STATE OF YUCATAN, 5 km S of Xocchel, road to Huhi, near Sacabah, tropical forest Guzman 23687. Rigidoporus vinctus (Berk.) Ryv. var. cinerea (Bres.) Setliff. Anew record from Mexico. The variety vinctus was reported from different localities in the State of Veracruz (Lowe, 1963, Guzman, 1972, Welden et a/., 1979, Anell & Guzman, 1987) as Poria vincta (Berk.) Curt. (except in the last reference, where it was cited as Rigidoporus). The var. cinerea differs from the typic by the grey to dark brown to almost black pore surface (pale ochraceous buff to light pinkish ochraceous in the latter), and by the perennial widely effused or effused-reflexed basidiocarp, with pores 6-12 per mm. Material studied. STATE OF QUINTANA ROO, 20 km from Felipe Carrillo Puerto, road to Vigia Chico, tropical forest, Guzman 20773-B. 16 Trichaptum biforme (Fr.) Ryv. Fig. 3 A very common fungus in Mexico in temperate and tropical regions, frequently reported as Polyporus pargamenus Fr. (Guzman, 1977). It is closely related with Trichaptum abietinus (Dicks. : Fr.) Ryv. from which differs mainly in the more purplish bluing color of the pore surface and larger basidiocarps. T. biforme is reported here for first time from the State of Quintana Roo. Material studied. STATE OF QUINTANA ROO, road Puerto Morelos to Tulum, near the deviation to Vallarta, road to the sea, tropical forest, Lopez 1830. STATE OF VERACRUZ, Uxpanapa region, tropical rain forest, Guzman 30466. Tyromyces fumidiceps Atk. This species was known only from Eastern U.S.A. and Canada (Gilbertson & Ryvarden, 1987). The light greenish color of the pore surface in herbarium specimens is characteristic. The Mexican material has laterally stipitate basidiocarp, and whitish to brownish yellow or brown chocolate color, azonate and tomentose pileus. Its habitat is always close to stream and rivers, where the basidiocarps are found on drift wood and similar debris. Material studied. STATE OF MORELOS, old road Mexico City to Cuernavaca, 3 km S of Tres Marias, Colonia Atlixtac, near a river, Pinus- Quercus forest with subtropical elements, Guzman 8474. Tyromyces galactinus (Berk.) Lowe A new record from Mexico. This common North American species (Gilbertson & Ryvarden, 1986-1987) is recognized by its soft, white to whitish gray or pale ochraceous, strigose to tomentose basidiocarps, with duplex context and fragant odour. The studied specimen presents imbricate basidiocarps growing on a living trunk of Platanus lindeniana Mart. & Gal. Material studied. STATE OF VERACRUZ, old road Xalapa to Coatepec, deviation to Rancho Guadalupe, way to Instituto de Ecologia, subtropical (mesophitic) disturbed forest, Bandala-Munoz Py Tyromyces humeana (Murr.) Lowe The species was only known from the type locality in Florida (U.S.A) (Gilbertson & Ryvarden, 1986-1987). The Mexican collection agrees well with Murrill’s fungus, except by the anise odour in fresh basidiocarps which was not indicated in the studied specimens. The 17 basidiocarp is white but in dry condition turns brownish even cinnamon brown in the center of the pileus. Material studied. STATE OF MORELOS, road El Parque to Tepoztlan, near Tepozteco mountain, subtropical forest, Guzman 5451. Table 1. Polypores new from Mexico Amylosporus campbellii (Berk.) Ryv. Anomoporia albolutescens (Rom.) Pouzar Antrodia gossypina (Speg.) Ryv. Antrodiella angulatopora Ryv. A. versicutis (B. & C.) Gilbn. & Ryv. Diplomitoporus lenis (Karst.) Gilbn. & Ryv. Flavodon flavus (KI|.) Ryv. Fomitopsis meliae (Underw.) Gilbn. & Ryv. Perenniporia tephropora (Mont.) Ryv. Phellinus chryseus (Lév.) Ryv. Ph. contiguus (Fr.) Pat. Ph. ferrugineo-velutinus (Henn.) Ryv. Ph. grenadensis (Murr.) Ryv. Ph. rhytiphloeus (Mont.) Ryv. Piptoporus soloniensis (Dub. : Fr.) Pil. Rigidoporus vinctus var. cinerea (Bres.) Selif. Tyromyces fumidiceps Atk. T. galactinus (Berk.) Lowe T. humeana (Murr.) Lowe 18 19 Figs. 1-3.- 1: Phellinus gilvus (Kruse, s.n., from Guerrero). 2: Fomitopsis melliae (Guzman 1915, from Veracruz). 3: Trichaptum biforme (Guzman 30466 from Veracruz). Figs. 4-7.-4-5: Daedalea microsticta (Sepulveda 19, from Nuevo Leon). 6-7: Ganoderma coffeatum (Guzman 30467, from Veracruz). 20 Table 2. Polypores from new localities in Mexico atrellus subrubescens (Murr.) Pouz. Amauroderma camerarium (Berk.) Furt. Antrodia malicola (B. & C.) Donk A. vaillantii (Fr.) Ryv. Aurificaria luteo-umbrina (Rom.) Reid Coriolopsis floccosa (Jungh.) Ryv. C. rigida (Berk. & Mont.) Murr. Daedalea microsticta Cooke Datronia caperata (Berk.) Ryv. Ganoderma coffeatum (Berk.) Furt. Grammothele fuligo (B. & Br.) Ryv. Incrustoporia nivea (Jungh.) Ryv. Inonotus hispidus (Bull. : Fr.) Karst. !. munzii (Lloyd) Gilbn. Junghuhnia nitida (Pers. : Fr.) Ryv. Microporellus obovatus (Jungh.) Ryv. Oligoporus fragilis (Fr.) Gilbn. & Ryv. Perenniporia ohiensis (Berk.) Ryv. Phellinus rimosus (Berk.) Pilat. Phylloporia chrysita (Berk.) Ryv. Ph. veraecrucis (Berk. ex Sacc.) Ryv. Polyporus varius Fr. Trichaptum biforme (Fr.) Ryv. ACKNOWLEDGEMENTS The authors acknowledge the authorities of their institutions for the facilites to their researches. The authors are grateful to Victor M. Bandala and Fidel Tapia of the Mycological Collection of XAL for his help in the herbarium and to Leticia Montoya from Instituto de Ecologia for revising this paper. Dr. Ake Strid, from Museum of Natural History, Stockholm, Sweden, kindly revised critically the text. LITERATURE CITED Anell, J.C. and G. Guzman, 1987. Especies de Poliporaceos citadas del Estado de Veracruz. Rev. Mex. Mic. 3: 137-148. 21 Anell, J.C. and G. Guzman, 1988. Nuevos registros de los hongos del grupo de los Poliporaceos del Estado de Veracruz. Fev. Mex. Mic. 4 : 25-42. Bandala, V.M., G. Guzman and L. Montoya, 1992. Los hongos del grupo de los Poliporaceos conocidos en México. Universidad Autonoma de Nuevo Leon, Linares (en prensa). Berkeley, M.J., 1867. On some new fungi from Mexico. Linn. Soc. Jour. (Bot.) 9: 423-425. Cappello, S. and H. Hernandez, 1990. Lista preliminar de los hongos (macromicetos) y mixomycetes de Tabasco, México. Universidad y Ciencia 7 (13) : 15-21. Castillo, J. and G. Guzman, 1970. Estudios sobre los Poliporaceos de Nuevo Leon, Il. Bol. Soc. Bot. Mex. 31 : 1-47. . Castillo, J., G. Guzman and G. Sepulveda, 1969. Estudios sobre los Poliporaceos de Nuevo Leon. |. Ciencia, Mex. 27 : 9-18. Cifuentes, J., M. Villegas, L. Perez-Ramirez, M. Bulnes, V. Corona, M. del R. Gonzalez, |. Jiménez, A. Pompa and G. Vargas, 1990. Observaciones sobre la distribuciOn, habitat e importancia de los hongos de Los Azufres, Michoacan. Fev. Mex. Mic. 6: 133-149. Diaz-Barriga, H., F. Guevara-Fefer and R. Valenzuela, 1988. ContribuciOn al conocimiento de los macromicetos del Estado de Michoacan. Acta Botanica Mex. 2: 21-44. Furtado, J.S., 1981. Taxonomy of Amauroderma (Basidiomycetes, Polyporaceae). Mem. N.Y. Bot. Gard. 34 : 1-109. Gilbertson, R.L. and L. Ryvarden, 1986-1987. North American Polypores. Fungiflora, Oslo (vols. 1 & 2). Guzman, G., 1963-A. Frecuencia y distribucion de algunos Basidiomycetes lignicolas importantes en Mexico. An. Esc. Nac. Cienc. Biols. 12: 23-41. Guzman, G., 1963-B. Macromicetos de las zonas aridas de Mexico, |. An. Esc. Nac. Cienc. Biols. 12: 43-60. Guzman, G., 1972. Macromicetos mexicanos en el Herbario The National Fungus Collections de E.U.A. Bol. Soc. Bot. Mex. 322731-55. Guzman, G., 1975. Hongos mexicanos (macromicetos) en los herbarios del extranjero Ill. Bol. Soc. Mex. Mic. 9: 85-102. Guzman, G., 1977. /Identificacién de los hongos. Ed. Limusa, Mexico City. 22 Guzman, G., 1983. Los hongos de la Peninsula de Yucatan, Il. Nuevas exploraciones y adiciones micolégicas. Biotica 8: 71-100. Guzman, G., 1986. Distribuci6n de los hongos en la region del Caribe y zonas vecinas. Caldasia 15 : 103-120. Guzman, G., V.M. Bandala, L. Montoya and Y. Saldarriaga, 1989. Nuevas adiciones sobre las relaciones micofloristicas entre Africa y el neotropico. El genero Rugosospora Heinem. (Fungi, Agaricales). Brenesia 32: 107-112. Guzman, G. and L. Guzman-Davalos, 1984. Nuevos registros de hongos en el Estado de Veracruz. Bol. Soc. Mex. Mic. 19: 221-244, Kickx, J., 1841. Notice sur quelques champignons du Mexique. Bull. Acad. Sc. Brux. 8: 72-81. Lowe, J.L., 1963. A synopsis of Poria and similar fungi from the tropical regions of the world. Mycologia 55 : 453-486. Lowe, J.L., 1966. Polyporaceae of North America. The genus Poria. State Univ. College of Forestry, Syracuse University, Tech. Publ. 90, Syracuse. Murrill, W.A., 1905. The Polyporaceae of North America. XIl. A synopsis of the white and bright-colored pileate species. Bull. Torrey Bot. Club. 32: 469-493. Murrill, W.A., 1907-1908. Polyporaceae. /n: North American Flora 9. N.Y. Bot. Garden, New York. Murrill, W.A., 1912. The Polyporaceae of Mexico. Bull. New York Bot. Gard. 8 (28) : 137-153. Murrill, W.A., 1915. Tropical Polypores. Lancaster, New York. Ojeda-Lopez, S., M. Sandoval and R. Valenzuela, 1986. Los Poliporaceos de México. |. Descripcién de algunas especies del noreste de Guanajuato. Rev. Mex. Mic. 2: 367-436. Ryvarden, L., 1981. Type studies in the Polyporaceae 13. Speccies described by J.H. Léveille. Mycotaxon 13: 175-186. Ryvarden, L., 1987. New and interesting polypores from tropical | America. Mycotaxon 28: 525-541. Ryvarden, L., 1991. Genera of Polypores. Nomenclature and Taxonomy. Synopsis Fungorum 5, Fungiflora, Oslo. Ryvarden, L. and |. Johansen, 1980. A preliminary polypore flora of East Africa. Fungiflora, Oslo. Saccardo, P.A., 1888. Sylloge Fungorum. Vol. 6 (Reprint 1944, Edwards Brothers, Ann Arbor). 25 Sharp, A.J., 1948. Some fungi common to the highlands of Mexico, Guatemala and Eastern United States. Mycologia 52 : 856-876. Steyaert, R.L., 1972. Species of the Ganoderma and related genera mainly of the Bogar and Leiden Herbaria. Persoonia (és her ike): Valenzuela, R. and R. Nava, 1991. E/ género Albatrellus (Polyporaceae) en México. Memorias IV Congreso Nacional de Micologia, Tlaxcala. Valenzuela, R. and S. Chacén-Jiménez, 1991. Los Poliporaceos de Mexico. Ill. Algunas especies de la Reserva de la Biosfera El Cielo, Tamaulipas. Rev. Mex. Mic. 7 : 39-70. Vazquez, L.S. and L. Guzman-Davalos, 1991. Nuevos registros de Poliporaceos estipitados de Jalisco. Rev. Mex. Mic. 7: 113-120. Welden, A.L., L. Davalos and G. Guzman, 1979. Segunda lista de los hongos, liquenes y mixomicetos de las regiones de Uxpanapa, Coatzacoalcos, Los Tuxtlas, Papaloapan y Xalapa (México). Bol. Soc. Mex. Mic. 13: 151-161. Welden, A.L. and P.A. Lemke, 1961. Notas sobre algunos hongos mexicanos. Bol. Soc. Bot. Mex. 26: 1-24. meee as ® at J iP) Ba PEt ell a Bes - aan +4 1 an rete ed a Senet Ke im i es ” eG Ra Rune YAO SAA “ore bir CNN ny aie i) ae ew const ese paar The as hte! os he Fas or ier ee One ; Lg a: KN ae pi Hy ital Ue a shih at. eave se ee i fh 4 ie ahs Wk 4 eye APY ee CAI: reve ee anata? ameter: MiG. aves? Yai ae ora te ‘prcrhan oe ee af ey rary Ay) Seats RSE a wh) oat sapere om i erie: in Ws, ‘goat not) a gt i ra , ‘4 We iy 4 wtih Ha ? Wer enh sibae Asta eae y ta ay re rh ice OG . et bite lait 4, ai faker Pon ae ei ae hey gl het yn wae i MR TAL Re Ng Hs ah aalh ee ty phate ea ie, » eT nee PL iat er Eine | MPDEH Laas ‘eta? PROG ati: 4 Cal, ort es Ae hal ae ave A ah ( eh Rise oe | 7 a ince Anny caviatale ih \ cakes Pee Bat, arian ey a 3 | ; | HN sida Ae i Widen i vy Nan a ae Renee: | Ba 0 ’ i . ; | Pian a qin "e i i ‘ | 7 NM art me | ba ; . - inn ee a | VP TRAM ho Tia ASL Bg HE iy eat ‘wits ca: Bora? eo : eens nih: Cay, Phe ee Te Ne ea MO Paes | aes ny ae a ; a ; a aye Teds a er (es he i es 4 Pa fat Ms Mat Le ie re i ne i a Fe aim asa AGP Meee eee) ae! is bg : iy a ; t “i | . : | VANE, SRN S EXt oe Geb ead: ep Ri ae Phd iat a han i a Bt wate . ae ee ; Me a dices pt ee. BED. sei tia 7) fight iat ane Race pen ae ei | es lah i ast a sy fanaa Pie a ‘ mi thot ian Ny he ; Sh Rex tid Sete ee IR te a ear res y ied my , ne any sii ee ae ' sy ry | é MY COTAXON Volume XLVI, pp. 25-39 April-June 1993 GLOMALES OF TAIWAN: Ill. A COMPARATIVE STUDY OF SPORE ONTOGENY IN SCLEROCYSTIS (GLOMACEAE, GLOMALES)! CHI-GUANG WU Soil Microbiology Lab, Agricultural Chemistry Department, Taiwan Agricultural Research Institute, Wu-feng, Taichung, Taiwan, R.O.C. KEY WORDS: Sclerocystis, sporophore, sporocarp formation, spore germination, spore ontogeny. ABSTRACT During the course of investigation on the Glomales of Taiwan, numerous sporocarps of Sc/erocystis were isolated and identified as SS. coremioides, S. clavispora, S. sinuosa, S. rubiformis, S. taiwanensis, and S. liquidambaris. After detailed comparison among these species by optical and electron microscopy, the similarity of spore ontogeny and spore arrangement in sporocarps was determined. Chlamydospores of S. coremioides with long, slender subtending hyphae also produced sporophores the same as those of other species of Sclerocystis and Glomus. The chlamydospores of S. coremioides are not always occluded by septa. The suggestion to define Sc/erocystis as a monotypic genus is rejected. A modified generic concept and a hypothesis to explain sporocarpic evolution of Sclerocystis are proposed. 1. Contribution No. 1637 from Taiwan Agricultural Research Institute. 26 INTRODUCTION Glomus and Sclerocystis were considered as closely related groups and Sclerocystis appeared to be evolutionarily advanced in sporocarp morphology from G/omus (Gerdemann and Trappe, 1974). Recently, Almeida and Schenck (1990) proposed to transfer most of the species of Sclerocystis to Glomus and retained only S. coremioides Berk. and Br. in the genus, because there were at least four uniquely different features in S. coremioides, i.e. (i) each spore forms individually on a subtending hypha without branching to form a sporophore near the spore base, (11) spores have a well-defined septum, (iii) spores are arranged in a hemispherical layer, and (iv) sporocarps form distally or laterally from the older sporocarps and appear fused in a column or branch. Almeida and Schenck (1990) concluded that these characters were not found in other species of Sclerocystis and Glomus. They applied Madelin's (1970) mode of sympodial conidial formation to distinguish spore ontogeny of Glomus from Sclerocystis, and postulated that S. rubiformis Gerdemann and Trappe (1974), S. clavispora Trappe (1977), S. sinuosa Gerdemann and Bakshi (1980), S. faiwanensis Wu and Chen (1987), and S. liquidambaris Wu and Chen (1987) should be transferred to Glomus. They have the same type of spore ontogeny as found in G. ambisporum Smith and Schenck, G. heterosporum Smith and Schenck (Smith and Schenck, 1985), and other species of Glomus. Glomus species have the ability to generate a lateral sporophore in the subtending hypha near the spore base. During the survey of Glomales of Taiwan, abundant sporocarps of Sclerocystis were collected from a variety of sites. These materials provided all of the presently described species of Sclerocystis and were very beneficial to the taxonomic study. In this paper, new evidence of spore ontogeny and sporocarp formation, a modified generic concept of Sclerocystis, and a hypothesis of sporocarpic evolution are presented. MATERIAL EXAMINATIONS Sporocarps were isolated by a wet-sieving and decanting (Gerdemann and Nicolson, 1963) and were observed by optical and scanning electron microscopy. In order to observe the central plexus of sporocarps, fruiting bodies were ultrasonicated to break down peridial tissues and chlamydospores. af The following specimens were examined: Sclerocystis clavispora-T AIWAN: Chi-tou, from rhizosphere of Chimonobambusa quadrangularis (Fenzi) Markino, Wu-850821; Chi- tou, from the rhizosphere of Erechthites valerianifolia (Wolf) DC., Wu- 8508201; Chi-tou, from the rhizosphere of Gingo biloba L., Wu- 8508202; Chi-tou, from the rhizosphere of Colocasia formosanum Hayata, Wu-8607151; Kenting, from the rhizosphere of Colocasia formosanum, Wu-8605171; Pintong, from the rhizosphere of asparagus (Asparagus officinalis L.), Wu-860813. Sclerocystis coremioides-T AIWAN: Tainan, from rhizosphere of Ageratum conyzoides L., Wu-850725; Tainan, from the rhizosphere of orange (Citrus sinensis Osb.), Wu-8608141; Taipei, from the rhizosphere of Bignonia sp., Wu-850810; Chi-tou, from the rhizosphere of Colocasia formosanum, Wu-8607152; Pintong, from the rhizosphere of asparagus (A. officinalis), Wu-860813; USA: Oregon, F55365S. Sclerocystis liquidambaris-T AIWAN: Taipei, Holotype, from rhizosphere of Liguidambar formosana Hance, Wu-860203; Isotype, Wu-860627; Kaoshiun, Liou-Kuei, from the rhizosphere’ of Cunninghamia lanceolata (Lamb.) Hook, Wu-870117. Sclerocystis rubiformis-T AIWAN: Chi-tou, from the rhizosphere of Erechthites valerianifolia, Wu-8508203; Chi-tou, from the rhizosphere of Gingo biloba, Wu-8508204; Chi-tou, from the rhizosphere of Colocasia formosanum, Wu-8607153; Kenting, from the rhizosphere of Colocasia formosanum, Wu-8605 172. “Sclerocystis sinuosa-TAIWAN: Tainan, Anpin; from the rhizosphere of Miscanthus sp. in Casuarina equisetiffolia L . woods, July 25, 1985, Wu-850725; Tainan, Anpin, from the rhizosphere of ground covers in C. equisetifolia woods, September 18, 1985, Wu- 850918; Pintong, by Kau-ping River, from the rhizosphere of legume (Glysine sp.), August 14, 1986, Wu-8608142; Pintong, by Kau-ping River; from the rhizosphere of asparagus (A. officinalis), August 13, 1986, Wu-860813; Pintong, Tongang, from the rhizosphere of unidentified herb grass, August 13, 1986, Wu-ME2; Tainan, Charlu, from the rhizosphere of citrus (Citrus sp.), August 14, 1986, Wu-8608143; Nantou, Chi-tou, from the rhizosphere of Sinocalamus sp., August 21, 1985, Wu-8508144; Taichung, Wu-feng, from the rhizosphre of maize 28 (Zea mays L.), November 14, 1991, Wu-111401; Orchid Island, Taitung, from the rhizosphere of C. equisetifolia, July 7, 1986, WU- 860707; U.S.A., INVAM collection, Gainesville, Florida (SSNS 122). Sclerocystis taiwanensis-T AIWAN:Chi-tou, Holotype, from the rhizosphere of Crassocephalum rabens (Juss. ex Jacq.) S. Moore, Wu- 8508205; Chi-tou, from the rhizosphere of Colocasia formosanum, Wu- 8607154; Taipei, from the rhizosphere of unknown host, Wu-850810; Tainan, from the rhizosphere of orange (C. sinensis), Wu-850724. RESULTS SPOROPHORE OF SCLEROCYSTIS: Sporophores branching from the base of chlamydospore are very commonly found in the every species of Sclerocystis. The sporophores are of various shapes such as membraneous vesicles (Figs. 1, 3, 4 ), dichotomous branches (Figs. 2, 6), or irregular forms. Sporophores are not uncommon in S. coremioides. They may be globular structures or tubular hyphal branches (Figs. 4, 7). The formation of sporophores is by branching from the chlamydospores with or without septa. Sporophores of S. /iqguidambaris are further differentiated into clubshaped structures which are parts of the peridium (Eigse oy SEPTA OF CHLAMYDOSPORES: Septa are very often formed at the bases of chlamydospores in Sclerocystis (Figs. 3, 4). The frequency of septa depends on the species and represents one of character in the species description. The septum is very commonly, but not always, found below the base of chlamydospores in S. coremioides (Figs. 4, 7). A Figures. 1-6. Micrographs of Sclerocystis spp. Fig. 1. Chlamydospores of S. sinuosa with a sporophore (arrow head). Fig. 2. Chlamydospores of S. liquidambaris with a sporophore (arrow head). Note paraphysis-like structures (P) enclosing the chlamydospore. Fig. 3. Chlamydospores of S. taiwanensis with sporophores (arrow head). Note the septa formed at the bases of spores. Fig. 4. A sporophore of S. coremioides formed below the spore base (arrow head). Note the presence of a membraneous septum (arrow) at the spore base. Fig. 5. Mature (upper) and vesicular young spores (YS) of S. clavispora. Note the mature spore enclosed by a exospore wall (arrows). Fig. 6. Hyphal branches formed from the spore base of S. clavispora. Scale= 30 tm. 29 30 similar type of septum occurs in S. sinuosa and S. taiwanensis (Fig. 3). Septa are also present in the subtending hyphae of S. rubiformis; however, they are much thinner than those of S. coremioides. ARRANGEMENT OF CHLAMYDOSPORES IN SPOROCARPS: The chlamydospores of S. coremioides are arranged in a hemispherical layer (Figs. 10, 16). The intersporal spaces are traversed by hyphae which grow radially from the central plexus and terminally connect to the peridium. Similar intersporal hyphae occur in S. sinuosa and _ “S. liquidambaris (Figs. 2, 9). In young sporocarps of S. rubiformis, chlamydospores are also arranged in a hemispherical layer (Fig. 12). However, with the subsequent formation of additional chlamydospores, these immature sporocarps are transformed into globular sporocarps with radially arranged chlamydospores. CHLAMYDOSPORE AND SPOROCARP FORMATION: All of the chlamydospores in sporocarps of Sclerocystis are produced from a central plexus. The central plexus ranges from a single flat, stellate cell as found in S. rubiformis and S. taiwanensis (Figs. 12, 15), to a mass of interwoven hyphae as found in S. clavispora, S. liquidambaris, S. sinuosa, and S. coremioides (Figs. 8, 10, 13, 16, 19, 20). The central stellate cell of the plexus in S. rubiformis is slightly different from that of S. taiwanensis. The former is a thick-walled structure connecting with a monohyphal stalk; however, the latter is a swollen cell initiated by fusion with more than one monohyphal stalk (Figs. 12-15). Among the interwoven plexal hyphae, flat stellate cells as found in S. rubiformis are also frequently observed and connected with chlamydospores_ or Figures 7-11. Micrographs of Sclerocystis spp. Fig. 7. Chlamydospore of S. coremioides not occluded by a septum. Note a branch (arrow head) formed on a subtending hypha. Fig. 8. Partial sporocarp of S. coremioides showing a slender subtending hypha (arrow heads). Fig. 9. Partial sporocarp of S. liguidambaris showing chlamydospores (S) embedded in club shaped structures (P) protruding from the central plexus. Fig. 10. Cross section of sporocarp of S. coremioides showing chlamydospores arranged radially in a hemispherical layer. Note the intersporal hyphae connecting between the central plexus and peridium. Fig. 11. Two naked sporocarps of S. clavispora (SC) and S. taiwanensis. S. taiwanensis producing remarkable smaller sporocarp. Scale bars in figures 7, 8, 9 =30 um. 31 | © © in a x @ N 32 intersporal hyphae (Fig. 19). Sporocarps of S. coremioides are frequently produced in columns or branched structures through lateral or distal proliferation. Nevertheless, the sporocarps of S. rubiformis are frequently interconnected into a flat piece (©2 cm diam.) by a monohyphal stalk. Among the collections of S. taiwanensis, a few young sporocarps were isolated and all of the chlamydospores were vesicular in appearance (Fig. 14). The apical wall of chlamydospores is formed last as sporocarps mature (Figs. 3, 17, 18). A similar type of spore formation is also found in S. clavispora (Fig. 5). HYPHAL STALK OF SPOROCARP: One of the main characters in S. coremioides is its hemispherical sporocarp with a hyphal stipe (Fig. 16). The hyphal stipe is a rope of interwoven hyphae. However, other species of Sclerocystis are only associated with one or more monohyphal stalks (Figs. 12, 14, 20). These monohyphal stalks are usually thick-walled. Young sporocarps of §. rubiformis with a monohyphal stalk are very similar to those of S. coremioides, excluding the absence of peridia and intersporal hyphae (Fig. 12). GERMINATION OF SPORES: Germinating sporocarps of 5S. coremioides and S. sinuosa have often been observed. Spores germinate from intact sporocarps, producing extramatrical vesicle-like structures. Figures 12-16. Micrographs of Sclerocystis spp. Fig. 12. A young sporocarp of S. rubiformis showing spores arranged in a broom shape. Note the plexus connecting with a monohyphal trunk (asterisk). Scale= 30 um. Fig. 13. A fragment of sporocarp of S. sinuosa showing sinuous peridial hyphae and simple interwoven plexal hyphae. Fig. 14. A primordial sporocarp of S. faiwanensis connecting with three monohyphal stalks (arrow heads). Note spores in a vesicular stage (V). Scale= 30 um. Figs. 15. A swollen central plexus of S. taiwanensis actively projecting out to form spores. Note the presence of a broad hyphal trunk (HT) and vesicles (arrows). Scale= 30 um. Fig. 16. Cross section of sporocarps of S. coremioides showing a hyphal stipe (HS), central plexus (PL) and spores produced in a hemispherical layer. Central plexal hyphae are comparatively broader than subtending hyphae of spores. Scale= 100 um. 33 DISCUSSION The Species of Sclerocystis included in the present study form a natural grouping on the basis of method of sporophore formation, spore germination, chlamydospore and sporocarp. production, ‘and _ the appearance of the septa and hyphal stalks of the sporocarps (Table 1). The affinity of S. coremioides is closer to the other species of Sclerocystis than to Glomus. Although three species of Glomus, i.e., G. ambisporum, G. heterosporum (Smith and Schenck, 1985), and G. dimorphicum Boyetchko and Tewari (1986) produce sporocarps similar to those of S. rubiformis the ability to produce dimorphic chlamydospores clearly distinguishes them from Sclerocystis. Dimorphic species, G. ambisporum, G. heterosporum and G. dimorphicum, appear to be transitional taxa linking G/omus and Sclerocystis, because they exhibit generic features of both. To date, these three species appeared to be only distributed in North America. The chlamydospores in sporocarps of Glomus were mostly arranged randomly and could be enclosed by a peridium or naked. Evolutionary trends of sporocarps in Sc/lerocystis are visible through a comparison of their structures (Fig. 21). Sporocarps of S. rubiformis are considered as primitive due to their lack of a peridium and producing sporocarps from a simple, broad, thick-walled plexal cell. Sporocarp aggregates were not uncommonly isolated and were interconnected by a single "fecund" hypha which was the same as that found in G. ambisporum (Smith and Schenck, 1985). In the naked sporocarps of §. ftaiwanensis , the central plexus is more complicated than in S. rubiformis and is formed by more than one monohyphal stalk (Fig. 15). Sclerocystis clavispora produces sporocarps similar to S. taiwanensis, but the central plexus is replaced by interwoven hyphae. Nevertheless, broad, thick-walled stellate cells similar to those of Figures 17-20. Micrographs of Sclerocystis spp. Fig. 17. Crushed young sporocarp of S. faiwanensis showing thin-walled spores (SP). Scale= 30 um. Fig. 18. Mature sporocarp of S. taiwanensis showing the gradually thickened apex of spore walls. Note a monohyphal stalk (arrow head) connecting with plexus. Scale= 30 um. Fig. 19. A broad, thick-walled cell (arrow head) in the central plexus of a sporocarp of S. liquidambaris. Scale= 30 um. Fig. 20. Aperidial sporocarp of S. sinuosa showing four monohyphal stalks (arrow heads). Scale= 100 um. 36 S. rubiformis are often mixed in the plexus. Consequently, S. clavispora is thought to be advanced S. faiwanensis, and both are senior to S. rubiformis. Sclerocystis liquidambaris appears to be derived from the S. clavispora/S. taiwanensis type, with the hyphal branches arising from the base of the spores becoming thick walled. The peridium of S. liquidambaris is thought primitive if compared with that of S. sinuosa and S. coremioides, since it is only tightly packed by thick-walled, clubshaped cells. In the sporocarps of S. sinuosa and S. coremioides, however, a peridium of interwoven hyphae is well developed. The peridial hyphae originate from the central plexus (Figs. 10, 16). Sclerocystis sinuosa has a number of monohyphal stalks radiating out of sporocarps. Thus, it resembles S. coremioides, except that the monohyphal stalks of this species are all combined into a coarse stipe. A single monohyphal stalk has the ability to produce a broomlike fruiting body, and further develops into a globose sporocarp as found in S. rubiformis (Fig. 12). However, the fusing of many monohyphal stalks into a stipe would result in a lose of the ability to produce spores in a globular arrangement. The result would be a hemispherical sporocarp. Results of this study support the retention of the generic concept of Sclerocystis as proposed by Gerdemann and Trappe (1974). However, the definition of Sclerocystis is modified as following. SCLEROCYSTIS Berk. & Br. emend. Wu Sporocarps globose or subglobose or hemispherical, enclosed by a peridium or naked, with a multihyphal stipe or monohyphal stalks; chlamydospores arranged side by side in a single layer, radiating out from a central plexus of hyphae; central plexus composed of a broad, stellate thick-walled cell or interwoven hyphae; development of chlamydospores within sporocarps synchronous or asynchronous; sidebranches frequently produced from the base of chlamydospores, becoming spores or intersporal hyphae. Fig. 21. Diagram of sporocarpic evolution of Sclerocystis spp. A. S. rubiformis, B. S. clavispora and S. taiwanensis, C. S. liquidambaris, D. S. sinuosa, E. S. coremioides. 21 OF, 38 TABLE 1. COMPARISON OF SPORE ONTOGENY AND SPOROCARP FORMATION IN SCLEROCYSTIS CHARACTERISTICS | S. COREMIOIDES | SCLEROCYSITS spp. SPOROCARP in chains or branches _| in flat pieces FORMATION (S. rubiformis) SPOROCARP vesicle structures vesicle structures GERMINATION (S. sinuosa) SPOROCARP STALK monohyphal stalk (S. rubiformis) more monohyphal stalks (S. sinuosa, S. taiwanensis..etc SPOROPHORE FROM | present present BASE OF CHLAMYDOSPORES ACKNOWLEDGEMENT This research was financially supported by National Science Council of Taiwan, Republic of China (NSC 82-0409-B-055-013). Appreciation is extended to Drs. R. E. Koske, James W. Kimbrough, and G. L. Benny for their critical reviews of this manuscript and for providing specimens used in this research. LITERATURE CITED Almeida, R. T. and N. C. Schenck. 1990. A revision of the genus Sclerocystis (Glomaceae, Glomales). Mycologia 82:703-714. Sh Gerdemann, J. W. and J. M. Trappe. 1974. The Endogonaceae in the Pacific Northwest. Mycologia Memoir No. 5, 76p. Iqbal, S. H. and B. Perveen. 1980. Some species of Sclerocystis (Endogonaceae) from Pakistan. Trans. Mycol. Soc. Japan 21:57- 63. Madelin, M. F. 1979. An appraisal of the taxonomic significance of some different modes of producing blastic conidia. Pp. 63-80. Jn: The Whole Fungus. Vol. 1. Ed., B. Kendrick. National Museum of Canada, Ottawa. Smith, G. W. and N. C. Schenck. 1985. Two new dimorphic species in the Edogonaceae: Glomus ambisporum and Glomus heterosporum. Mycologia 77:566-574. Trappe, J. M. 1977. Three new Endogonaceae: Glomus constrictus, Sclerocystis clavispora, and Acaulospora _ scrobiculata. Mycotaxon 6:359-366. Wu, C.-G. and Z. C. Chen. 1986. The Endogonaceae of Taiwan. I. A preliminary investigation on Endogonaceae of bamboo vegetation at Chi-tou areas, Central Taiwan. Taiwania 31:65-88. Wu, C.-G. and Z. C. Chen. 1987. The Endogonaceae of Taiwan. I]. Two new species of Sclerocystis from Taiwan. Trans. Mycol. Soc. Rep. China 2:73-83. ie ' be rh Le na Wha a ae my ini Hy qi A gt ia ivi ae saci fa wih ie pina bint. on ( ¥ 4 uiwld \ iP ha are a Me j " ‘4 a i ME DEN eV ib irr. 3 1. sie be ascii nuaneiny ih ee eae mn iF ‘ oe 2 “ Ba " wortla eae ” nat . ‘| Ss tea ny Was ; ‘1, Rule ay eh ye i ' er h +i a ry py yg! ae iby ide “ule poh Ae heal ‘ ae ir vie we i ve -f f fy i ” ry 1 ! i hah a af aNaie ain tS Legale) the a ae Md af ae ve ied Aa vei: aie ‘hy | | eae re la sh al ire iG ‘i 7 mal RAL es wy inant ee! : Pir none Ap aa aces FRE A RG a, ane: ceva, | mee gt re wee se ee elem nr. An Mey Tg FY, Fgh «ls ag as a a 7 sie wr ; yy hd ey vy : 7" 4 ha ' 7 enh 1h gee By | ve ‘ y. i : RM ee | f I Ne ek ee t Tas i WOM Riga thet aw The! os ; Mi r Bf r f ; i oh t Oy , £ q x nnpheehsek i ty st ie Da een ka ea 9 ne _ Too ; oe ew : prs 1) ley ny ve : 4 ‘ rr vee . i: 1 4 if 7 1 . bach ales Teo a at ae) eee ae: A : \ eit ie Te De heater Je Tea os a 2 ma . : me a r eee a a | ni rs Peer. ul Pe ha eee RT Wa as ie aia “yt . Aber ' pen ley i oe he We a, na fi \ our AO) " a J afi : fave v , rh Bg ay Bi 6 We, el ie" 4 , 4 aie di a in ¢ re et i . - ' ka F 1 we UPL i, D a yt a v os .. ; ’ 4 i : 7 - } ’ Fi oe a! is | ee hy. bel i ; te iT ata a.) ie { ay ; f a s i Y i UJ Ny i aU ef at ba 1. ‘ ‘ at eA ( ny ey i i) , ni , MY 4 ar an ; f ah ~ Ve ee me eta ek h q per [ Di u AA ay f P A ike hay Aa ’ wat co | ' at + ao We : bay ote ae ee ey a, ‘i - oe ry uk ie Ties das Sa A wee: OU ahs oe (oe a ih MI y i i P is P ‘ = ' i MAD eae Su oi te sy fl : Le Pee a Lead ly oye -, Me eaeair nm, im i 4 a aang re gene! A een aba ie oe, iid a i Na ‘iy ri ey uh pa Git ews J ; nit Aber 7 “ = Pa a vs ig ye 7 na MY COTAXON Volume XLVI, pp. 41-57 April-June 1993 SYSTEMATIC AND BIOLOGICAL STUDIESIN THE BALANSIEAE AND RELATED ANAMORPHS. III. ASCOSPORE AND MACROCONIDIAL GERMINATION AS A TAXONOMIC CRITERION RYAN A. PHELPS and G. MORGAN-JONES Department of Plant Pathology, College of Agriculture and Alabama Agricultural Experiment Station, Auburn University, Auburn, Alabama 36849 ABSTRACT Ascospore and Ephelis macroconidial anamorph germination in vitro on agar media is described in Balansia aristidae Diehl and B. epichloe (Weese) Diehl, and ascospore germination in Myriogenospora atramentosa (Berk. & Curt.) Diehl. A review of the literature on the subject of spore germination in the Balansieae is presented. Consideration is given to the usefulness of differing germination patterns in the taxonomy of the tribe, including species of Atkinsonella Diehl, Balansia Speg. and Epichloe (Fr.) Tul. It appears that, in the taxa so far studied in detail, differences occur that are reflective of both generic and specific distinctions. INTRODUCTION As alluded to in the prologue to this series of papers (Morgan-Jones et al., 1992), a number of problems exist in the taxonomy of the Balansieae. These have arisen partly because of a limited number of criteria, including host relationships, and overall morphology of conidial and perithecial stroma, used in delimiting taxa. In many instances, species distinctions are not readily apparent solely on the basis of host specificity and/or stroma morphology. In the case of Balansia aristidae and B. strangulans (Mont.) Diehl, for example, their stromata are essentially similar except for the fact that in the former perithecia are more prominent. Likewise, their Ephelis anamorphs are virtually indistinguishable. During the course of the present studies, attempts are being made to evaluate additional criteria of potential taxonomic usefulness. Among them are details of host-pathogen interactions, in vitro colony growth characteristics and peculiarities of ascospore and Ephelis macroconidial germination. It is also anticipated that analysis of molecular sequence data will aid in resolving questions of relatedness among taxain this complex. Alabama Agricultural Experiment Station Journal Series Number 18-923366 42 Diehl (1950) reported studying the germination of the Ephelis macroconidia of six species of Balansia in tap and rain water and on the surface of sterile agar media. The taxa observed in this regard, except for Balansta obtecta Diehl, and the composition of the media, were not specified. Germination of the conidia of Ephelis mexicana Fr., the anamorph of B. obtecta, was illustrated by a line drawing. No clear distinction was found in conidial appearance or germination among the species. Both a single primary apical germ tube and also lateral ones were reported to be produced. Diehl (loc. cit) described the acicular Ephelis conidia as being characterized by indentations that were thought to be due to the presence of septa, although these could not be discerned in most species. A tendency for conidia to break apart at one or two slight constrictions was also considered to indicate occurrence of septa. Septa were reported to occur in the macroconidia of Balansia linearis (Rehm) Diehl and ascospores were said to develop septa before or at germination. When the latter occurs, each cell may form one or more germ tubes perpendicular to the axis of the ascospore. Some swelling of the segments of the ascospores of B. linearis at germination was illustrated. Ullasa (1969), in a study of Balansia claviceps Speg. in vitro, investigated conidial and ascospore germination in hanging drop cultures. Conidia were found to swell considerably in water and water agar prior to germination. After forty-eight hours, one to five polar and lateral germ tubes were formed. At this stage conidia became variously septate and individual cells became somewhat bulbous and produced more than one germ tube. After seventy-two hours germ tubes branched or remained simple and formed whorls of secondary conidia at their tips similar in morphology to the parent conidia. Such conidia either germinated or produced tertiary conidia by a budding process. On potato dextrose agar slow-growing yeast-like colonies consisting mainly of masses of conidia were produced. Ascospores germinated similarly by germ tubes, at the tips of which whorls of conidia were formed. Luttrell and Bacon (1977) reported that in Myriogenospora atramentosa, which they classified in the Balansieae, one-celled part-ascospores consistently germinated by forming an aborted foot at one end and a coiled germ tube at the other. The germ tube failed to develop into a hypha on common laboratory media. It was also stated that in species of Balansia each ascospore cell may germinate by a coiled germ tube produced laterally just below each septum. The Ephelis anamorph was described as "fusoid conidia produced on phialides at the surface of the stroma" but no account of conidial germination was given. Subsequently Rykard et al. (1982) showed the Ephelis state of this species to have holoblastic, sympodial conidiogenous cells and described conidial germination. This was found to be characterized by the development of bipolar or lateral germ tubes. In a liquid medium (referred to as M102), conidia germinated by forming bulbous, flattened, spoon-shaped ends which produced elongated germ tubes. On cornmeal malt extract (CMM) and water agar (WA), branched or unbranched, determinate germ tubes functioned as conidiogenous cells, producing a sequence of secondary conidia. No mention was made of any conidial septation. Conidium production from ascospores, without an intervening hyphal phase, is known to occur in a number of ascomycetes. Several didymosporous, 43 phragmosporous and dictyosporous species belonging to the Nectria Fr. complex, for example, produce microconidia, often referred to as ascoconidia, from ascospores while still remaining within asci. In Nectria aquifolii (Fr.) Berk. and N. coryli Fuckel numerous microconidia are formed from minute, open- ended, tube-like extensions at both ends of their one-septate ascospores, and their ontogeny is considered to be phialidic (Booth, 1959). Likewise, microconidia are produced from the multiseptate ascospores of Scoleconectria cucurbitula (Tode:Fr.) C. Booth through minute openings in their walls. (Booth, loc. cit.). Rossman (1983) described the ascospores of this fungus as ‘budding’ to form ascoconidia. A similar process occurs in species of Thyronectria Sacc., including T. balsamea (Cooke & Peck) Seeler, T. berolinensis (Sacc.) Seaver and T. Jamyi (Desm.) Seeler. In all of these taxa ascospores never produce germ tubes but rather disintegrate following microconidial production. Ascospore germination in Epichloe typhina (Pers.:Fr.) Tul. was investigated by Bacon and Hinton (1988) on CMM and water agar. A process, referred to as iterative germination, was reported to occur in which ascospores produced lateral, aculeolate, phialidic conidiogenous extensions perpendicular to their long axis. One such entity was produced from each cell of the ascospore and sometimes delimited from it by a transverse septum, sometimes not. Each conidiogenous element produced several conidia terminally. Under varied conditions, germ tubes leading to hyphal development werenever produced. Germination of discharged ascospores of Atkinsonella hy poxylon (Peck) Diehl and A. texensis (Diehl) Leuchtmann & Clay was studied by Leuchtmann and Clay (1989) on cornmeal-dextrose agar. This was found to be of an identical pattern in both species. Within the ascus ascospores are usually seven-septate but split at their median septum, where they are constricted, into two four- celled part-ascospores prior to germination. Germ tubes were found to be initially produced, one per cell, at both ends of the part-ascospores and laterally, immediately below the septa delimiting the end cells. No germ tube was produced in the vicinity of the median septum of each part-spore. At a later stage, the part-ascospores often split further, at their median septum, and additional germ tubes were formed from free ends or from the base of existing tubes. No study of Ephelis anamorph conidial germination in these species was reported. We have had opportunity to study Balansia aristidae, B. epichloe, and Myriogenospora atramentosa in vivo on grasses in Alabama and the former two have been brought into pure culture. It is clear from previous reports that ascospore and/or conidial germination patterns differ between genera and, at least in some instances, between species of the Balansieae. In order to further evaluate the possible usefulness of modes of germination in the taxonomy of the tribe, ascospore and conidial germination characteristics of the two Balansia species, and ascospore germination in M. atramentosa, have been investigated and are reported upon herein. MATERIALS AND METHODS Plants of Aristida purpurascens Poir., Eragrostis capillaris (L.) Nees, and Paspalum notatum Fliiggé, bearing stromata of B. aristidae, Balansia £4 epichloe, and Myriogenospora atramentosa, respectively, were collected from sites in Lee, Montgomery, and Tallapoosa counties during early to mid summer 1991. Voucher specimens from these collections have been deposited in the Auburn University Mycological Herbarium (AUA) and at BPI. Infected plants were transplanted into 15 cm diameter plastic pots containing local sandy loam and grown in a Conviron growth chamber. An environment of 90% relative humidity and alternating 12-hour light/dark periods (provided by incandescent and fluorescent lights) at temperatures of 32° C and 24° C respectively, was maintained. Using a sterile dissecting needle, macroconidial masses were scraped from superficial ephelidial pustules on culm-borne stromata of B. aristidae and from leaf-borne stromata of B. epichloe. No conidia were harvested from Myriogenospora atramentosa. Ascospores were harvested from all three taxa by cutting off the tops of several mature, stroma-borne perithecia horizontally with a razor blade, followed by removal of their exposed contents with a sterile needle. Both conidia and ascospores harvested by these methods were immediately suspended in sterile distilled water. Spore suspensions were then diluted to appropriate concentrations, and plated on each of three media. The media used during this study were 2% WA, CMM, and EC- 14 (3 g KH>PO,, 2 g K,HPO,-3H20, 2 g NH4S0O,q, 5 g MGSO,-7H30, 40 g mannitol, 10 g yeast extract, 20 g agar, + water to equal 1 L), a medium developed by the late Edward M. Clark and Richard A. Shelby at Auburn University for grass endophyte culture (unpublished). Using a glass rod, approximately .3 ml of spore suspension was spread onto the surface of the medium in each petri plate. Ascospores were also harvested by taping mature perithecial stromata inside the lids of petri plates and awaiting the forcible discharge of ascus contents onto the agar surface below. After 1, 2, or 4 hours the stromata were removed and the plates sealed. Germinating spores were observed in situ with a Zeiss microscope using low power objectives. Limitations of such observations included compromised resolution and contrast resulting from the relatively thick agar background, and inability to study the spores at high magnification due to lack of clearance for an oil immersion lens. Attempts to transfer germinating spores from petri plates to microscope slides generally resulted in damage to the spores. To enhance imaging, and enable viewing at high magnification, spores were allowed to germinate on medium- coated 22 mm #1 slide coverglasses. Coverglasses were dipped in hot agar medium and placed inside petri plates on the surface of the same medium to prevent desiccation. After adding the spore suspension to the upper exposed surface of the coverglasses and allowing the excess liquid to evaporate, the petri plates were sealed and incubated at 25° C. Inoculated coverglasses were PLATE 1. Balansia aristidae. A-F, Ephelis conidia; A, from nature; B, initial swelling on agar medium; C, apical swelling (early stage); D, apical swelling (later stage); E & F, apical swelling (final stages), and initiation of primary germ tubes: G-L, part-ascospores; G & H, from mature perithecium (median septa indicated by arrows); I, median disarticulation and terminal swelling (early stage); J-L, median disarticulation and terminal swelling (later stages), lateral germ tube initiation (indicated by arrowheads) and disarticulation of swollen ends. 45 46 removed periodically during germination, inverted, and mounted on hanging-drop slides with the surface bearing spores downward. Prior to viewing, excess agar was gently wiped from the upper surface of the mounted coverglass. When standard slides were used instead of hanging drop slides, germinating spores were frequently damaged. RESULTS Balansia aristidae. Conidia. Conidia of this species are long, cylindrical to somewhat fusiform, obtuse at the apex, subtruncate at the base, 10 to 15 ym long and less than 1 pm wide (Plate 1, A; Figure 1, A). Germination by germ tube occurred after 4 days on EC-14 and CMM media and was preceded by morphological changes. Following 48 hours on these media, conidia swelled appreciably at their apex to form an ellipsoid to oblong portion about 3 ym in length and 2 ym wide, while the opposite, basal end remained more or less truncate (Plate 1, A & B; Figure 1, B). After 5 days the swollen portion became much more elongate (up to 7 pm), assuming a long-oblong or more or less cylindrical aspect or, in some cases, became narrowly obpyriform (Plate 1, D; Figure, 1, B). Initial germination was mostly monopolar but variable. A primary germ tube often originated from the conidium immediately below the base of the terminal swelling or a short distance below it (Plate 1, E & F). Alternatively, more rarely, a germ tube originated apically. Occasionally this was followed shortly thereafter by disarticulation of the terminal swelling from the remainder of the conidium. After 10 days, some swollen terminals became up to 12 pm long where apical germination did not occur. Following initial germ tube production a secondary germ tube sometimes originated proximal to the first. In some instances, following a period of 12 days, a germ tube was also produced from the opposite, basal end of the conidium. Many conidia underwent the described morphological changes but failed to germinate after 10 days. Although most conidia failed to germinate on WA, germination followed a similar course except that terminal swelling was delayed 24 to 48 hours and never developed beyond 3.5 ym in length. After 12 days a small percentage developed a swollen spherical structure up to 6 pm in diameter (Figure 1, C), located distal to the already swollen oblong, terminal portion, frequently giving rise to one or two germ tubes. Such germ tubes generally developed into extensive hyphal systems. Though rare, hyphae sometimes also emerged from the basal end. PLATE 2. A, Balansia aristidae part-ascospores 72 hours after direct discharge onto agar medium; B-G, part-spores after 4 to 7 days on agar media showing various stages of terminal swelling, median and terminal septation and disarticulation (indicated by arrowheads), and sympodial germ tubes; H, chlamydospore-like cells arising from a part-spore on agar medium; I-L, Myriogenospora atramentosa part-ascospores showing stages of germination and germ-tube branching. 47 48 C (Bike Be a et} Sym FIGURE 1. Balansia aristidae, Ephelis conidia. A, from nature; B & C, stages and patterns of germination. Spm | FIGURE 2. Balansia aristidae part-ascospores. A, from nature; B & C, various stages of germination, septation and disarticulation. 50 Ascospores. Ascospores of Balansia aristidae disarticulate within the asci, forming one-septate part-spores which are then discharged. Part-spores are narrow and cylindrical with truncate ends, 21 to 34 jm in length and less than 1 pm wide (Plate 1, G & H; Figure 2, A). After 48 hours on EC-14 medium, part-spores swelled at both ends forming bulbous, ellipsoid terminals 3.5 to 4.5 pm in length and disarticulated at the median septum (Plate 1,1; Figure 2, B). Following another 48 hours, the swollen ends became nearly 9 pm long. Lateral germ tubes developed directly or a short distance below the terminal swellings (Plate 1, H; Plate 2, B& C). Concurrently or immediately before germ tube initiation, a septum was laid down just above its point of origin. Disarticulation of the swollen terminal from the remainder of the half part-spore frequently occurred at this septum (Plate 2, D). After 7 or more days, some swollen ends reached 14 zm in length and a second germ tube was occasionally produced from the base of the half part-spore (Plate 2, G; Figure 2, C). A few half part-spores swelled appreciably, becoming multiseptate and forming thick-walled, melanized chlamydospore-like structures (Plate 2, H). Part-spores behaved similarly on WA. Septae and terminal swellings, however, were not as well-defined, and spores became generally more irregular in shape. Lateral and terminal germ tubes occurred frequently after 48 hours, and quickly developed into mycelia. Thick-walled, chlamydospore- like cells also began to occur at 48 hours. After 4 days these structures were extremely common. Following 7 days, part-spores had either failed to swell and began deteriorating, formed mycelia, or formed chlamydospore-like structures. Balansia epichloe. Conidia. Conidia of this species are acicular to fusiform, acute at the apex, subtruncate at the base, 16 to 21 um long and less than 1 pm wide (Plate 3, A; Figure 3, A). Within 24 hours on EC-14 medium conidia swelled slightly and germinated by apical germ tubes, one per conidium. Moderate swelling occurred evenly over the basal two-thirds of the spore, the apical portion remaining more attenuate (Plate 3, B). Just prior to apical germination, the extreme tip swelled slightly (Plate 3, C). After 24 hours germ tubes averaged 7 pm, emerging from a Slight enlargement at the apex of the tapered conidium (Figure 3, B). A similar enlargement developed at the basal end also. After 48 hours apical germ tubes approached 12 pm in length. Germination remained monopolar for nearly PLATE 3. Balansia epichloe. A-H, Ephelis conidia; A, from nature; B, initial swelling; C & D, further swelling and early stage of germination; E, well- developed apical germ tubes; F & G, disarticulation of initial germ tube(indicated by arrowhead) and initiation of sympodial germ tube; H, bipolar germination; I-L, part-ascospores; I, from mature perithecium; J & K, median disarticulation (indicated by arrowhead) and terminal germination; L, sympodial development of secondary germ tubes (indicated by arrows) and disarticulation of initial terminal germ tubes (indicated by arrowhead). Hy oy B ) FIGURE 3. Balansia epichloe, Ephelis conidia. A, from nature; B & C, stages and patterns of germination. A eee > a Fagen TE Ome a FIGURE 4. Balansia epichloe part-ascospores. A, from nature; B, various stages of germination. 54 5 days, at which time a second germ tube occasionally developed from the basal enlargement (Plate 3, H). More commonly, sympodial initiation of a second germ tube occurred at the apical end of the spore (Plate 3, F & G). Subsequent disarticulation of the initial germ tube typically ensued. Germination rarely occurred on WA, and only after 72 hours. The characteristic swelling prior to germination was more pronounced than on EC- 14 medium. Following 5 days on WA germ tubes up to 5.5m had developed. Though infrequent, some conidia swelled appreciably, approaching 40 pm in length and 2.5 ym in basal width, the apical half remaining quite narrow. Even after 10 days germination failed to progress further. Ascospores. Ascospores of Balansia epichloe disarticulate within the asci, forming one-septate part-spores which are then discharged. Part-spores are broadly filiform with truncate ends, 32 to 46 ym in length (Plate 3, I; Figure 4, A). Although half part-spores seldom separated completely, their partial disarticulation at median septae was a common phenomenon. Germination on EC-14 medium occurred within 24 hours. A germ tube developed from the outer exposed end of each half part-spore, and usually grew sharply to one side or the other (Plate 3, J &K). Since half part-spores typically remained connected, this germination pattern resembled bipolar germination of the part-spore. After 72 hours germ tubes had reached at least 10 to 14 um in length on WA, and at least 17 pm in length on EC-14. No further development occurred on WA. On EC-14 medium, sympodial initiation of secondary germ tubes, and disarticulation of initial germ tubes followed (Plate 3, L; Figure 4, B). Myriogenospora atramentosa. Ascospores. The germination pattern characteristic of ascospores of this species is particularly distinctive. Long, narrow, filiform, aseptate part-spore with acutely tapered ends are discharged from each ascus. Rarely, these part-spores remain joined end-to-end in pairs. Within 48 hours on WA and EC-14 media, part-spores exhibited monopolar germination by the development of a germ tube from a slight apical enlargement. A similar swelling on the other end remained dormant throughout germination, which continued for several days and then ceased. At first germ tubes were distinctly curled, becoming undulate or loosely circinate to somewhat coiled (Plate 2, I-L; Figure 5, B). On EC-14 medium, branching was frequent, each branch continuing the same growth habit. FIGURE 5. A, Myriogenospora atramentosa part-ascospores from nature; B, part-spores germinating; C, germinating ascospores of Atkinsonella hy poxylon (after Leuchtmann and Clay, 1989); D, segment of Epichloe typhina ascospore showing iterative germination (after Bacon and Hinton, 1988). 56 DISCUSSION Ascospore and conidial germination peculiarities clearly vary appreciably between the member genera of the Balansieae and, to a lesser extent, between species. It is interesting to note that ascospore germination in the two species of Balansia investigated during the course of this study is more closely similar than between these and species of Atkinsonella. Rykard et al. (1984) questioned the desirability of maintaining Atkinsonella and Balansiopsis Hohnel as separate genera from Balansia. Further information on ascospore germination in additional species of Balansia will, perhaps, help resolve that question. As pointed out by Leuchtmann and Clay (1989), disarticulation of seven-septate ascospores into two three-septate part-spores, and the number and location of germ tubes formed subsequently, appears to be unique for the genus Atkinsonella. This is certainly different from species of Balansia where the released part-spores are one-septate and those of Myriogenospora which are aseptate. That the two known species of this genus, A. hypoxylon and A. texensis, have identical ascospore germination (Leuchtmann and Clay, loc. cit.) indicates that they are closely related. The latter, on morphological grounds alone, was originally classified as A. hypoxylon var. texensis by Diehl (1950). It was recently recognized as a separate species mainly on the basis of host association, isozyme variation, colony differences in vitro and host cross- inoculation incompatibilities (Leuchtmann and Clay, loc. cit.). The distinctive ascospore discharge and germination characteristics of Epichloe typhina and Myriogenospora atramentosa are further indicative of the usefulness of this feature in determining relatedness. The single-celled part- spores of the latter and the very different germination morphology are reflective of the genetic distance between Myriogenospora, a monotypic genus, and others classified in the tribe Balansieae. The monopolar germination characteristic of each part-spore also indicates this. Myriogenospora was not included by Diehl (1950) in the tribe but added by Luttrell and Bacon (1977) despite its considerably different epiphytic habit and ascus morphology. Luttrell and Bacon (loc. cit.) considered the type of germination in M. atramentosa to be significant, noting that in species of Balansia each cell in the filamentous ascospore may germinate by a coiled germ tube produced from the side of the cell just below the septum. That is, of course, a simplification and generalization for, as indicated in the results noted above, there are variable patterns of ascospore disarticulation, secondary septation, and secondary breaking apart of part-spores, associated with the sequence of germ tube initiation. Germ tubes in Balansia do not usually appear to be undulate, circinate or coiled in the manner of those formed from ascospores of Myriogenospora. The swelling of the Ephelis conidia of M. atramentosa in liquid culture reported and illustrated by Rykard et al. (1982) is, interestingly, reminiscent of the morphological changes, described herein, undergone by conidia of Balansia aristidae prior to germination. The latter differs, however, in that the swelling occurs at the apex of the conidium only and not at each end, as was noted to occur in M. atramentosa by Rykard et al. (loc. cit.). The sequence of secondary conidia sympodially produced from the tip of a determinate germ tube in M. atramentosa form in essentially the same manner as the primary conidia. 57, Such sympodial growth is also seen occurring during germination of conidia and ascospores of both B. aristidae and B. epichloe. The sequences of events occurring during macroconidial and ascospore germination in Balansia aristidae and B. epichloe are distinctive for each species and, therefore, of use in their characterization. The considerable degree of difference in ascospore growth pattern, morphology and disarticulation is of particular interest. The secondary septation and the eventual breaking up of the original one-septate part-spore into four separate cells in B. aristidae is, to our knowledge, unique in the genus. There is, in this case, a sequence of primary, secondary and tertiary disarticulation of the original ascospores delimited within the asci. Whether such differences exist between other species of the genus remains to be determined. The account by Ullasa (1969) of germination in Balansia claviceps in hanging drop cultures is not comparable to the present study because of the differing conditions and less detailed observations. It would be interesting, for example, to investigate if there are differences in regard to germination between B. aristidae and B. strangulans, two species that are considered close (Diehl, 1950), and likewise between B. epichloe and B. henningsiana (Moell.) Diehl, two species that are superficially similar, and from time to time have been considered to be the same fungus, but are now recognized as separate and distinct entities. There is reason to believe that, even if all individual species lack clear-cut differences in respect to germination peculiarities, this criterion is nevertheless of some use in indicating relatedness and/or separateness. ACKNOWLEDGEMENT This research was supported by a National Science Foundation (BSR- 8922157) grant to J. F. White, Jr. and G. Morgan-Jones. We thank Dr. J. Leland Crane for reviewing this paper. LITERATURE CITED BACON, C.W. and D.M. HINTON. 1988. Ascosporic iterative germination in Epichloe typhina. Trans. Br. mycol. Soc. 90: 563-569. BOOTH, C. 1959. Studies of Pyrenomycetes. IV. Nectria (part 1). Mycol. Pap. 73: 1-115. DIEHL, W.W. 1950. Balansia and Balansiae in America. Agric. Monograph No. 4, USDA, Washington, D.C. 82 pp. LEUCHTMANN, A. and K. CLAY. 1989. Morphological, cultural and mating studies in Atkinsonella, including A. texensis. Mycologia 81: 692-701. LUT TRELL, E.S. and C.W. BACON. 1977. Classification of Myriogenospora in the Clavicipitaceae. Can. J. Bot. 55: 2090-2097. MORGAN-JONES, G., R.A. PHELPS and J.F. WHITE, JR. 1992. Systematic and eee studies in the Balansieae. I. Prologue. Mycotaxon 43: 401-415. ROSSMAN, A.Y. 1983. The phragmosporous species of Nectria and related genera. Mycol. Pap. 150: 1-164. RYKARD, D.M., E.S. LUT TRELL and C.W. BACON. 1982. Development of the conidial state of Myriogenospora atramentosa. Mycologia 74: 648-654. eee on 1969. Balansia claviceps in artificial culture. Mycologia 61: 72-579. = 6) Vy Cory i ee Lewy y a ‘ . f Wah ; ark Ware oad ‘ { k \ LAER, i} \ ty : . heres h BA set > PY, lJ , | 1) iF, a a : ‘y iy ek frit “aii ite \ f ; i fie pa ay | Ney M ts Peel ales he ESE TA heme Sent ha ings ? i at , ' 4 i ‘ f ony fap it rN Hi ia as nay : A in see ‘oe itl re ey it de my, fh si \ ' oye Atyey meg ye a NY inde Gon : J r ( i ae Te uae nae RG one h IB RR A Tey ee 4 Tere iT wh mR: ye H! Sy vax | Te: vray yey A res bal ry [aA ena i® i ake oA are A A Mi Lika WAM eee ree es BP a ar : i , i} % j Ley ; i ie Wie ‘ K : ‘eh Vea 7 1 ak i ‘D if a ly 7 ack ae Un r { : er Vc netrie. ot 4 lu i) ; ; i ie ‘ F , Mes ibe Beis tt et Te i ‘ d ny ' ¥ tae eat 0 Adee * og Fra ay iF « “sie a: pat isang ah i | ds aint wea fren } ’ oh ; ment int = et r] nay Py i , f ~- ' ‘ ‘ r Wh ’ j ; i ‘) { 4 ii } ' v1 i 1 * i ‘4 a F 7 ale i « ed my i aie Line , a 4) ~ if . 1h ate i b 4 s i) bi } r j \ ae 4 > t 1) Me ary ae NS ars na ee Ce may te } nM / mS , \ nt ile ‘1 dial’) oyehat a ay tip ab AP MBOR Eytan. | ALT AFT CAVE ioe NS Gaiam eh ates Ma j ; niet ) i fale | A an y ¥ iN vue Ws ais ual ew ai Hay ah "| i 1 aA T PT aaa ee i, A ae K ASI int i A } a) ; att a } , F sa ) *5 ry 7 iy Ly iy Vee ‘ Le #1 ng we 1 gay A ease ere i il alah j +} 4 ru i vi soli RT hn ba L7, stat ah ee MY COTAXON Volume XLVI, pp. 59-65 April-June 1993 Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian Peninsula M. Muntafiola— Cvetkovic* and A. Gémez-— Bolea* *Department of Plant Biology, Unit of Botany, Faculty of Biology, University of Barcelona, Diagonal 645, 08028 Barcelona, Spain. Abstract. Chaetospermum chaetosporum (Pat.) Smith & Ramsb. was encountered on graminicolous leaves and sheaths with small distorted areas, in the Tibidabo mountain, Catalonia. The conidiomatal structures of the fungus showed an_ evident resemblance to gelatinous sporodochia. This is the first record of the above mentioned taxa in the Iberian Peninsula. Key words: Coelomycetes, Chaetospermum, C. chaetosporum. In July 1990 several graminicolous decaying plants were collected in the Tibidabo mountain, near the city of Barcelona (Catalonia, Spain), showing coloured, slightly thickened, distorted areas on the leaves and sheaths. A closer examination of the lesions revealed the presence of orange —yellow structures, gelatinous to the touch, resembling groups of sporodochia. The microscopic preparations made from such crushed structures showed abundant subcylindric conidia with bipolar appendages characteristic of the genus Chaetospermum Sacc. The sporodochium —like bodies and the Chaetospermum conidia were seen in subsequent collections of similar lesions on leaves of decaying Gramineae growing near the same place. Some conidia of other fungi (mainly Alternaria and Cladosporium species), together with those of Chaetospermum, were observed eventually in the slides, as well as some few conidia of Drechslera (probably D. victoriae (Meehan & Murphy) Subramanian & Jai). Since no published reports have been found concerning the presence of Chaetospermum species in the Iberian Peninsula, the description of the fungus encountered in the Tibidabo is herein presented. 60 Material and Methods Collections studied. The material studied consisted of decaying grass leaves and sheaths showing small distorted areas with minute sporodochium — like conidiomata. The material was collected in the Tibidabo mountain, Serra de Collserola (S00 m altitude), near Barcelona, Catalonia, Spain, the 1st, 7th and 8th July 1990, a month with no rains, that followed a dry period. Microscopic preparations and examination. The conidiomata were removed from the plant substrates with sterilized needles and mounted onto glass microscope slides in lactophenol slightly stained with safranin. Preparations fro, thirteen different collections were sealed with a nailpolish double coat and labeled EE. 90, 30-42. Microscopic examinations were made using a Leitz Diaplan microscope and 1,000 magnifications by oil—immersion, bright field, phase-contrast and the Nomarski interference optic systems. Micrographs were taken with a Nikon F-—610 camera and Ilford Pan F, 50 ISO film. Slides labeled EE. 90. 30 and 33 were deposited in the Mycological Herbarium BCC (University of Barcelona). Results The morphologic features of the fungus encontered in the Tibidabo mountain were the following: Conidiomata consisting of orange—yellow, sessile, gelatinous, mostly aggregate and roughly spherical structures more similar to sporodochia than to typical acervula. Conidiophores constituted by tufts of hyaline hyphae, mostly 30-40 x 15-2 wm (Fig. 1, arrowed), with terminal or lateral conidiogenous cells, swelling at their tips to give rise the conidia. Conidia elongate — cylindrical, straight to slightly bent smooth—walled, hyaline, (15—) 30-35 (-45) x (5-) 7.5 (-10) pm, aseptate, with granular contents (Figs. 1 and 2), with the apices irregularly obtuse or sometimes almost truncate and bearing three to six terminal or subterminal appendages. Appendages filiform, flexuous, not branched, not septate, 1 wm _ width, generally as long as the conidium body, or slightly shorter; with no protoplasmatic continuity with the conidium body when the internal contents retracts. Figs. 1 and 2. Chaetospermum chaetosporum (Pat.) Smith & Ramsb. (Mycol. BCC Herbarium NO EE. 90. 30). Conidiophores (arrowed) and conidia showing the granular protoplasm conspicuously retracted (separated) from the cell—wall, and the apical and subapical insertion of the appendages. Scale bars = 10 um. 61 62 Discussion The several generic names that have been given to the fungi now gathered in the genus Chaetospermum reflect the views of their authors concerning the conidiomatal structure of these fungi. The conidiomata, initially roughly spherical and later rupturing irregularly, show such,a variation in morphology that the species have been placed in the Tuberculariales, Melanconiales and Sphaeropsidales. Patouillard (1888) considered a sporodochium the conidiomata of the fungus that he described as Tubercularia chaetospora, for which Saccardo (1892) erected the genus Chaetospermum, with the following short diagnosis: Sporodochia gelatinosa. Sporophora ramosa. Conidia ovoidea, hyalina, utrinque 7-8 setosa. Later on the conidiomata of Chaetospermum were again referred to as sporodochia (Smith & Rambsbottom, 1913; Agnihothrudu, 1962). However, the Chaetospermum conidioma was considered essentially a pycnidium by other authors (Héhnel, 1924; de Fonseka, 1960; Nag Raj & Kendrick, 1972), while after Sutton (1980) it is acervular. The conidiomatal structures of the fungus collected in the Tibidabo evidently grade into sporodochial conidiomata. Sutton (1980) put into synonymy with Chaetospermum the genera Ciliospora Zimmerman, Mastigonema Speg., Entomopatella Petrak, and Chaetospermella Naumov, as he had discussed early (Sutton, 1977). Of the five Chaetospermum species keyed by Agnihothrudu (1962), Sutton (1980) tentatively recognized only three, mainly based on conidium size. This author maintained C. gelatinosum Petch in the genus, although recognized that the conidia are much thicker—walled than in the other accepted taxa and the appendages are very thin, perhaps not "cellular" in origin. In Tab. 1 the main morphometric characters of the fungus collected in the Tibidabo have been compared with those of C. chaetosporum, the type species of the genus, and other congeneric taxa. It can be concluded that the Chaetospermum structures found on the plants collected in the Tibidabo fit well with the morphology and dimensions of C. chaetosporum. It could be only remarked here that septate spores like those reported and illustrated by de Fonseka (1960) from 3-4 month-old conidiomata were not seen in our collections, although the retraction of the conidium body protoplasm observed in some slides could indicate that the fructifications were quite old. The retraction was not considered a manipulation artefact, since the phenomenon was observed in water preparations as well as in the lactophenol — mounted slides. Much attention was paid in this study to the appendages of the conidia since many species of the Coelomycetes show similar structures. Their features have been considered by Sutton & Sellar (1966) and Sutton (1968, 1973) of great importance. Sutton’s criteria have been sustained by Pirozinski & Shoemaker (1971), Nag Raj (1981, 1988) and others. According to these criteria the conidial appendages of the fungus found in the Tibidabo must be considered of "cellular" origin. 63 086T suoTz wil og wt [Z-91XPP-9€ ‘u0}4NS 0} dn ‘21-9 “‘peTTemM-yoTyy wr ¢xoE yojed WnsouTzeTes “OD Z96T suo, wi 22 wil (9-) 7-€ wi Z-[xXZT 0} dn ‘yTUusy 0} dn ‘6-<¢ x(92-) 22-9T sTereyey eyewr}[N ‘ylusy aeTTTeuled *D = 086T wi [Xgt-P1 *u0}4NS 0} dn °6-S wr (9-) ~-€X9Z-9T Ui %=1 x71 02.00 Issey], Wneuled ‘9 ee ee eae ee a ee ie a ee eee Z68T ‘2098S ‘opres90eS 8-1 win QT-8X8p-8F sSeptotsze[nozeqny “OD = 88st *yeq ePIOdsSoOyseyo “qed wil QT-gxep-8E erre[noteqny, = 086T suo, wi cy 0} dn ‘qsuey 3 YWIWS ("}ed) *u0}4NS OT-€ win GTt-8XxSP-9Z wil Zx0¢€-0T uintodsoyeeyo “DO 096T unl TXg9-GE unl GI-€TXESG-8Z ‘qsuey 8 YyTUIS ("}ed) ‘eyosuoy op *TT-€ PeTTeM UTUY winztodsojyeeys *O €T6l “qswuey 4y8UeT ‘qsuey 8 YytUS ("3ed) R UWS WNTptuod se ‘g-/ wid QT-9X€Pp-8E wniodsojseeyo °9 wr (OT-) L-S suo, wi cp 07 dn X(GP-ST) SE-SZ oqeprqrL wor *9-€ peTiemM-utyy win 7-S"TX0b-0€ uinultedsozeeryo (ezIs pue Jewnu) sooustejoy sodsepueddy etptuog so1oydorptuog exe, exe} peye[el Iey}O JO ssoy} YIM pereduios ‘OqepIqtL, 24} UT pe,eTjoo wnazodsojyeeyo wnuedsoyeeyD jo sueutoeds ey} Jo sein}yess OTrJoWOYdIOW “T “qeL 64 C. chaetospermum appears not to be a common fungus in the Iberian Peninsula. Records of this species or other related taxa are not found in the literature. Since July 1990 we were not able to collect again this species. C. chaetosporum has been reported on stems, twigs and leaves of a variety of plants belonging to very different genera, but no studies have been found emphasizing this organism as a pathogenic one. However, the lesions observed on the leaves and sheaths from which the fungus was recovered in the present study were similar to those attributed by Sprague (1950) to Dilophospora alopecuri (Fr.) Fr. Controlled inoculation tests of healthy plants with C. chaetosporum should dilucidate the questions related to _ its pathogenicity, specially taking into account that conidia of other fungi were found in the samples examined. Acknowlegments M. M.-C. is grateful to the staffs of the "Centre de Documentacié i Experimentacié de Ciéncies", Barcelona, and of the "Departament de Biologia Vegetal, Unitat de Botanica, Facultat de Biologia", Barcelona, for the use of their facilities. The authors are indebted to Profs. X. Llimona and J. Guarro for critically reading the typescript. References AGNIHOTHRUDU, V. 1962. Notes on fungi from North-East India X. Chaetospermum camelliae sp. nov. on tea (Camellia sinensis (L.) O. Kuntze). Mycopathologia et Mycologia Applicata 16, 113 — 116. FONSEKA, R.N. de 1960. The morphology of Chaetospermum chaetosporum. Transactions of the British mycological Society 43, 631-636. HOHNEL, F. von 1924. Uber die Gattung Chaetospermum Sacc. Mitt. bot. Lab. techn. Hochsch. Wien, 3, 86 — 88. NAG RAJ, T.R. 1981. Coelomycete systematics. In The biology of conidial fungi. Vol. I. Edited by G.T. Cole & B. Kendrick. Academic Press: New York. NAG RAJ, T.R. 1988. Genera coelomycetarum. XXIV. Chithramia anamorph gen. nov. Canadian Journal of Botany 66, 903 — 906. 65 NAG RAJ, T.R. & KENDRICK, W.B. 1972. Genera Coelomycetarum III. Pestalozziella. Canadian Journal of Botany 50, 607-617. PATOUILLARD, M.N. 1888. Note sur une Tuberculariee graminicole. Bulletin de la Société Mycologique de France 4, 39-40. PIROZYNSKI, K.A. & SHOEMAKER, R.A. 1971. Some Coelomycetes with appendaged conidia. Canadian Journal of Botany 49, 529-541. SACCARDO, P.A. 1892. Sylloge Fungorum 10 (p. 706). Padua. SMITH, A.L. & RAMSBOTTOM, J. 1913. New or rare fungi. Transactions of the British mycological Society 4, 318-330 (p. 328). SPRAGUE, R. 1950. Diseases of cereals and grasses in North America (Fungi, except smuts and rusts). Ronald Press: New York. SUTTON, B.C. 1968. Kellermania and its generic segregates. Canadian Journal of Botany 46, 181-196. SUTTON, B.C. 1973. Coelomycetes. In the Fungi. An Advanced Treatise. Vol. IV A. Edited by G.C. Ainsworth, F.K. Sparrow & A.S. Sussman. Academic Press: New York. SUTTON, B.C. 1977. Coelomycetes VI. Nomenclature of generic names proposed for Coelomycetes. Mycological Papers 141, 1-253. SUTTON, B.C. 1980. The Coelomycetes. Fungi Imperfecti with Pycnidia Acervuli and Stromata. Commonwealth Mycological Institute: England. SUTTON, B.C. & SELLAR, P.W. 1966. Toxosporiopsis n. gen., an unusual member of the Melanconiales. Canadian Journal of Botany 44, 1505 - 1513; peed ii bebe i sea hue me hs a) tien cate ORY im RTE, Mek. on Th mi y an ines whi rusia sige 7 LORIE ate smack Oe es i ar abide tans ea en ae RA is EL he "ah eh re in i Katie eet kaa | cat alee Tenant: as Tonal aa . ae oF wg he te 6 rain seers A vol i, Aye ant wy ependinte A Ace! Ries 6 2 eit mea, Oe seek “‘Dathigh ees, i) A ryigw iy ro ans al UK: diy i. bee ney A nn vt ‘ a 4° OM wi) # Lf wt, ie) ili | iunssirg ene pai menenHraey kink. Sak eye anne ie A! A “prrade. ES ) aA ae “ Ree, fit ava? aes ieee id itt. Me ~ t 9 P i shite OW. is, ine ™ thee ns coi ath oer a. d SOAR : CS San ny co ont: iene, ho ee fate & meahgt gsi dgtca} i nee rainty at cha mt of tis ae Onl ht el, « Capea eT - 4 , » ae a Bi ee: othe eM Te we bgt a CMa che te hoo ee Pee ne OUR , a J 41 yaks ‘avg abe Title a Fine by Re bee otal Ay ba al mere ter ont ene sa ipa 2 err ie Re a ae ate Ek a a | ; . % ‘dou por Mar ts a) - m4), i oN / i > } » ; on A, Mas ee ae he ee iy ots ; (apo See ie Saal a ah tind ei eS A pe | itd reo pe Arh ee ere eS tie Py he nlite sa CArsee walao ) nor a 1 Stes rma Si , ee hee is hay ‘adlinis ay "Ks auld ai Ce eer: i se en B q He pate be ake sh apd tis tytn nnn Wa ieben : saa eae tee ia A Layee ; (haben es a ; ¢ Me Pee a a yi a pie a thy Mit ie - a thes 7 $40 f bhe F seat, iy 7 ae phat pitty Wisin : a sant sven righ init oe ne ae: * a MK i hte | rae Pa a) nf a “ ip Nae Alay stabi Wg Pai: oe area _ Ome ; Heys Aes \ Soaitaahe s(qioe. egih yer ; ih nt? uh re ie opens, life - Pe < ou ’ ce ‘ . | A ian J een o~ r if Ls Nits 6 ee Bets ie nate i, mM fe ie had ihe hp ®t Wi4. 4 eae aa wy) — v) a be Hai Pile a a) ad AEE ye | am er prio ee er eee ie ahi 2 ale tae ay el ae 7 hae ; iat? : nee Yu ; “J ok MY COTAXON Volume XLVI, pp. 67-69 April-June 1993 LECTOTYPIFICATION OF OPHIOBOLUS TRICHELLUS (DOTHIDEALES, ASCOMYCETES) Christian Scheuer Institut fiir Botanik der Karl—Franzens—Universitat Graz, Holteigasse 6, A—8010 Graz, Austria A lectotype specimen is designated for Ophiobolus trichellus SACC., BOMM. & ROUSS. The species, recently placed in Tubeufia by SCHEUER (1991), is briefly described and illustzated. Key words: Tubeufia trichella, taxonomy. SCHEUER (1991: 814) has recently published a new combination of Ophiobolus trichellus SACC., BOMM. & ROUSS., Tubeufia trichella. Although the British material examined agrees with the original description in every respect, the new combination was somewhat doubtful, because no type material of Ophiobolus trichellus had been available at that time. A quite satisfactory syntype specimen from Knocke (Belgium) could be located in PAD. It agrees with the protologue and with the British material studied and illustrated by SCHEUER (1991). No specimen from Oostende (Belgium), the second locality given by the authors in the protologue (in BOMMER & ROUSSEAU 1890: 259), could so far be found in any herbarium. The syntype specimen in BR (Sur les feuilles d'dmmophila arenaria, [Belgium,] Knocke, Aoit 1890, leg. M. ROUSSEAU) consists only of a small leaf fragment bearing no ascomata of Ophiobolus trichellus. Nevertheless, because of the two localities given in the protologue, a lectotype must be designated. Ophiobolus trichellus P. A. SACCARDO, BOMMER & ROUSSEAU in BOMMER & ROUSSEAU, Bull. soc. roy. bot. Belgique 29: 259 (1890), later cited erroneously as 'O. trichellus BOMM., ROUSS. & SACC.' in Syll. Fung. 9: 934 (1891). 68 = Ophiochaeta trichella (SACC., BOMM. & ROUSS.) SACC., Syll. Fung. 11: 352 (1895). = Tubeufia trichella (SACC., BOMM. & ROUSS.) SCHEUER, Mycol. Res. 95(7): 815 (1991). Ascomata (Pseudoperithecia) + scattered on dead leaves of Ammophila arenaria (L.) LINK, usually epiphyllous, nesting among the leaf hairs, brown, + globose, ca. 100 um in diam., with a fringe of dark brown, simple setae around the ostiole. No branched setae have been found in the lectotype material. Setae ca. 20-40 um in length, tapering from the base, but always rather torulose and irregular in outline, thick-walled, usually non- septate. The ostiole is surrounded by small, knob-like peridial cells. Peridium in surface view composed of irregular, angular cells up to ca. 8 wm in diam., consisting of 3 - 5 layers of slightly flattened cells. Pseudoparaphyses present in mature loculi, ca 1.5-2 um thick. Asci bitunicate, + cylindrical, 45-66 x 10-15 “~m (SACCARDO, BOMMER & ROUSSEAU 1890), 8-spored. Ascospores hyaline, flexuous, cylindric-filiform, ca. 48- 65 x 2.5 wm, 14—-16-(?18-)septate, guttulate. The ascospores are usually coiled in the ascus, before the ascus expands at maturity. LECTOTYPUS designated here: A la face inférieure des feuilles du Psamma arenaria [=Ammophila arenaria], [Belgium,] Knocke; ex Herb. E. BOMMER & M. ROUSSEAU (Herbarium Mycologicum P. A. SACCARDO, 2729, PAD). Lectotype of Ophiobolus trichellus: (A) Ascoma. (B) Ascospores (lactophenol permanent slide). — Bars = 20 um. 69 This material differs only very slightly from the British collections studied and illustrated by SCHEUER (1991: 814, 815). No branched setae have been found in the lectotype, but in some (not all!) of the British collections. In addition, apparently some of the ascospores of the two ascomata mounted from the lectotype have more septa (up to 18?) than those of British material (up to 15). Thanks to Dr. Sergio CHIESA (Padova) for his help in tracing the type specimen in PAD, and to PD. Dr. Orlando PETRINI (ETH Ziirich) for acting as pre—submission reviewer. References BOMMER, E. & M. ROUSSEAU 1890. Contributions a la flore mycologique de Belgique. — Bulletin de la societé royale de botanique de Belgique 29: 205-302. SCHEUER, Ch. 1991. Taphrophila (Dothideales: Tubeufiaceae) and two new species of Tubeufia with dark setae. - Mycological Research 95(7): 811-816. a) oe OO Ee b rte LF 7 pry Liat J ra Nai oe 6 A RAL aA War Yoh Vi “de Re 1 rd . at . ty ¢ i] : HM i y Ee ‘oe @ i Mae ies 4 5 en Y ; A Ps Curh. eH erg i 7 i ‘ if ¥ Ma \ wi ic ina ores: aN a fs ea x V ro) Ps - ea hegeis' ys rene 4 abe es Lie fing ) ib ts Pe fia 3 a) mi 4 ‘ ay oy : A ca va a *¥ y, “— ene it sa nal 7 a" iP 0 f " fae abil ae f ‘ a Fat at ; ai r ny ib ay : “gy ay Sire Pea , t ae t 1 Aa PE eae RS kate ti ay aes a hea + it 4 =f y ; - Ay p. Sari , - 4 ah at ahs Tw Athi! Ne Agi Fas Lean cs y * era 4 D rf bo wep Te. Wi . pay oy isin Aes 4 we ‘ pate, . PIG bps Md ta aT aah, a eal ‘ ' 4 , . : i rif ws j i" "ly oY, Vir» ole wee be i { eo 4 : Lae i: i a) Mh al ; +> Mey Pet cae fa 2) ae » af { 5 Vo Be We j 4 n ‘ \ “+ 3 ’ i r Faced Ye pane ey par nea A bay Fas 2.) igh: i pasts Lat a MYCOTAXON Volume XLVI, pp. 71-80 April-June 1993 TAXOMYCES ANDREANAE, A PROPOSED NEW TAXON FOR A BULBILLIFEROUS HYPHOMYCETE ASSOCIATED WITH PACIFIC YEW (TAXUS BREVIFOLIA) GARY STROBEL and ANDREA STIERLE Department of Plant Pathology Montana State University Bozeman, Montana 59717 DON STIERLE Department of Chemistry Montana College of Mining Technology Butte, Montana 59701 and W. M. HESS Department of Botany & Range Science Brigham Young University Provo, Utah 84601 ABSTRACT Taxomyces andreanae is described as a novel endophytic fungus associated with the inner bark of Taxus brevifolia Nutt. (Pacific yew). This fungus has small septate hyphae which average 1.2 wm and large septate hyphae which average 3.75 wm in diameter. Hyphal cells of this fungus are multinucleate. It characteristically forms clumps of loosely constructed cells (bulbil-like). These clumps are of various shapes and sizes typically ranging from ca. 5x5 to 16x30 wm in diameter and length. The nucleated cells in these clumps average about 1.5x2.5 wm and appear to be loosely packed in the bulbil and are incapable of germination. This fungus grows rapidly on many common laboratory media, covering the plates with its mycelium in 3-4 days. itaelacks cclamp connections, and dolipore septations. Its telemorph is unknown. 72 INTRODUCTION The Pacific yew (Taxus brevifolia Nutt.) is the primary source of taxol, a promising anticancer drug isolated from the inner bark of this tree (1,2). In a search for microbes that may influence the production and/or fate of taxol (and related taxanes)in this tree, we have made a concerted effort to isolate and identify parasitic and endophytic microbes associated with it. In the present report we describe a novel endophytic hyphomycete, Taxomyces andreanae, that was isolated exclusively from the inner bark on small limbs of a specific yew tree in Northern Montana. MATERIALS AND METHODS The fungus was isolated from the upper limbs of a shrub- like Pacific yew tree growing as undercover in a mature, undisturbed cedar forest in Flathead County, Montana. Small limbs (0.5 - 1.0 cm) were surface treated with 70% ethanol. The outer bark was peeled back and pieces of the white inner bark (phloem/cambium) were aseptically removed and placed on H,O agar. Hyphal tips of fungi growing from the pieces of the plant were placed on mycological agar and fungal growth was enhanced. Subsequent attempts to isolate the fungus from other sites in Montana, Idaho, Oregon and Washington were unsuccessful. The growth pattern of the fungus was studied on other additional plant species. Leaf, stem, and bark samples of various plant species growing near the tree from which this fungus was obtained were collected near the Hungry Horse dam site on the Flathead National Forest. Small pieces (0.5 - 2 cm) of leaves, stems and bark of these species were placed over 4 layers of cheesecloth, thoroughly dampened, and autoclaved. Agar blocks (1.0x1.0 cm) supporting fungal growth were then placed on the sterilized plant material and fungal growth observed and measured after 1 week. The growth of the fungus was measured daily after placement of 0.5x0.5 cm agar blocks on standard freshly prepared agar media plates (Difco) eg. potato dextrose broth agar, nutrient agar, oatmeal agar, cornmeal agar, lima bean agar, water agar, and malt agar. Agar blocks having mycelia and bulbil-like structures were fixed and dehydrated as for transmission and scanning electron microscopy (SEM) (3,4,5,6). For SEM, the material was then critical point dried, gold coated and sputter coated, and observed with a JEOL 840A scanning electron microscope. Fungal structures were measured on SEM micrographs after critical point drying of tissues. This 1S, drying procedure causes some shrinkage of biological structures (ca. 10%) which means that they are probably Slightly larger, and the clumps of cells more tightly packed when in the living state. Nuclei were stained with the 4,6 =diamidino-2- phenylindole reagent (5ug/ml of 20% ethanol) and observed under UV at 365 nn. RESULTS AND DISCUSSION Taxonomic Treatment and Description Taxomyces andreanae Strobel, Stierle and Hess gen. et sp. NOVen 4 ELOS s.1 =6)i. Fungus endophyticus e cortice interiora Taxo brevifolo Nutt.; hyphae dimporphae -- parvae 1.25 wm et magnae ca 3.75 um latae et longae; bulbilus cellularum ca 1.25x2.5 um et laxe contiguus et apparenter non germinans; mycelium celiter crescens, hyphis fibulis nullis et doliporis septis nullis; telemorphus ignotus. Mycelium superficial, composed of a network of highly branched, septate, multinucleate, usually hyaline, smooth walled hyphae. Smaller hyphal cells average 1.25 pum in diameter. Larger cells average 3.75 wm in diameter (Figs. 1- 2). Cells are budded from fructigenous hyphae forming clumps which vary enormously in shape from spherical to ovoid to longiform and in size from 5x5 wm in diameter (for spherical bulbils) to 16 - 30 wm in length (for elongate bulbils). Bulbil cells remaining colorless. The cells seem to be loosely packed in the bulbil and are ovoid ca. 1.5x2.5 wm and are never observed to germinate. The "clumps" of cells in Fig. 2 and 3 are not located on sterigmata, but seem to arise by a "budding process" from hyphal pores (Fig. 3). The number of cells in each clump Variesawidely (Pig. £2 &) 3)". We have referred to these "clumps" as bulbils after the broad definition of deBary (7), thatsers, "Small pluricellular bodies incapable of germination". In 7. andreanae, the cells of the bulbils, unlike most bulbilliferous fungi, appear to be loosely packed, but nevertheless connected with fibrous material (Figs. 3,6). These clumps of cells might also be considered as conidial masses but since germination has never been observed (in sterile H,O and nutrient broth), the clumps of cells seem to better fit the broad description of a bulbil. Furthermore, transmission electron microscopic examination of these bulbil cells reveals that they are engorged with cytoplasmic structures including lipid bodies and each has a nucleus (Figs. 5,6)(8,9). They also possess a bilayered cell wall 74 (Figs. 5,6). Nevertheless, these bulbils differ from the bulbils of other standard bulbilliferous fungi by lacking pigmentation, certain sclerotial-like qualities (outer rind- like cells and inner swollen cells) and in the manner of their formation (10). T. andreanae may be related to Cladorrhinum spp. by virtue of certain conidial and hyphal characteristics (11). However, our SEM and culture observations made on the type species - Cladorrhinum foecundissimum revealed the absence of bulbils, the presence of spherical conidia, and the presence of small, numerous phialid-like structures on the hyphae, presumably from which the spores are budded (11). Furthermore, C. foecundissimum and related species are heavily pigmented (green- black colonies) and are commonly isolated from soil and dung sources. In addition, T. andreanae was inhibitory to C. foecundissimum. These findings are all in sharp contrast to the descriptions given in this report of T. andreanae. Type species: Taxomyces andreanae Strobel, Stierle, & Hess. Holotypus: Based on material taken from the bark of Taxus brevifolia Nutt., collected by Andrea and Don Stierle, August, 1991, in Flathead County, Montana, USA. Agar slants containing the type culture are deposited with the MSU mycological collection, D. E. Mathre, Department of Plant Pathology, Montana State University - col. no. 738. Duplicate cultures are deposited at the CBS, Baarn, Netherlands; International Depository Authority, CBS 279.92. Dried specimens are on deposit at the Farlow Herbarium, Harvard University (FH). Etymology The source tree of the fungus is Taxus, hence the genus designation - Taxomyces. The species name, Taxomyces andreanae, is in honor of Andrea Stierle. Andrea is a colleague, research associate and dedicated scientist. She and her husband, Don Stierle, were the people who originally isolated this organism. Cultural Characterization When an agar plug of inoculum was placed in the center of most freshly prepared agar plates enriched with various nutrients it grew so rapidly that it reached the edge of the plate in 3 days (corn meal agar, lima bean agar, nutrient agar, malt agar, oatmeal agar). Bulbils did not form on any of these media up to 6 days after inoculation. However bulbils were noticed on the inoculum piece on the corn meal agar after 6 days. Some bulbils were noticed at the edges of ihe) the malt agar plate 7 days after inoculation. Numerous fluffy aerial mycelia were especially observed on malt agar and after 6-7 days the mycelium on the malt agar developed a deep reddish-brown coloration and a thick mycelial mat. When the fungus was placed on the autoclaved leaves, fragments of small limbs, and bark taken from various tree species located in the geographical area of Taxus brevifolia, the best mycelial growth and bulbil formation occurred on Pacific yew (Table 1, Fig. 4), followed by River birch (Betula nigra) (Table 1). In contrast, there was no growth on, or bulbil formation on, Larix occidentalis, or Tsuga heterophylla, (Table 1). Other species differentially supported weak fungal growth and light bulbil formation, eg. Pinus monticola, Picea engelmanni (Table 1). These observations suggest the likelihood that some host preference of Taxomyces andreanae exists in nature and that it would be unlikely to be found in and on many species other than Taxus or Betula. This organism appears to be a saprophyte or endophyte with the latter preferred since it was found in association with living tissue. After one year it was still possible to recover the organism from the inner bark of the source tree. There is no evident gross pathology of this host tree. Furthermore, attempts, using agar blocks infested with T. andreanae placed under the bark of yew also failed to cause any disease manifestation. We also noted that the thicker hyphae ca. 3.75 um in dia, were the ones that typically extended the mycelial mat from one object (leaf or limb fragment or agar block) to another. These might be considered "exploratory hyphae". Careful study of the cultural, mycelial and bulbil characteristics in comparison to other bulbilliferous fungi nicely demonstrated the uniqueness of Taxomyces andreanae (10). ACKNOWLEDGEMENTS We acknowledge the financial assistance of the National Science Foundation and the Montana Agricultural Experiment Station. Appreciation is also expressed to the Hungry Horse Ranger District of the Flathead National Forest and the Superintendent of Glacier National Park. We also acknowledge discussions with Dr. Amy Rossman, ARS, USDA; Dr. Don Mathre and Gene Ford of Montana State Univesity; and Dr. Ed Butler, University of California, Davis. The description was rewritten in taxonomic format and translated by S. L. Welsh of Brigham Young University. The authors appreciate the help of Ms. Pam Berger in typing and laying out this manuscript. Fig. 1. A scanning electron micrograph of hyphae and fructigenous hyphae of T: andreanae. Bar=10 um. Fig. 2. A scanning electron micrograph of a series of various shaped bulbils of T. andreanae. Bar=10 um. Fig. 3. A scanning electron micrograph of a single bulbil of T. andreanae illustrating the organization of the cells of the bulbil. Bar=10 pm. Fig. 4. Growth of T. andreanae on the inner bark of Pacific yew (left) and growth and bulbil formation on the leaves and small limb fragments of T. andreanae (right). 78 Transmission electron micrograph of bulbil cells of T. andreanae illustrating a dense cytoplasm in each cell. Bar=1 pm. Transmission electron micrograph of TI. andreanae bulbil cells at a higher magnification. Note that the cell wall has two distinct layers, and the fiberous material between the cells. Bar=1 um. 79 Table 1. Growth and bulbil formation of Taxomyces andreanae on various tree species normally growing in the vicinity of Taxus brevifolia. All observations were recorded 1 week after inoculation with a 1.0x1.0 agar block supporting fungal growth. Plant Species Av growth from Formation edge of agar of block (cm) bulbils* Twigs/leaves Bark Twigs/leaves Bark Taxus brevifolia Nutt. Pacific Yew 2.0 0.25-0.5 heavy heavy Betula nigra L. River Birch 1.0 0.75 heavy moderate Pinus monticola Doug]. Western white pine 0.1 0.2 light light Tsuga heterophylla Rafn Sarg. 0.0 O71 none none Western hemlock Pseudotsuga taxifolia (Poinjebritt: Ou 0 none none Douglas fir Thuja plicata Donn. Western Red Cedar 0125 075 light to heavy moderate Picea engelmanni Parry exEngelm. 0.1 0 moderate none Engelmann spruce Larix occidentalis Nutt. 0 0 none none Western larch *Bulbil formation is given in terms of heavy (completely covering the area where mycelium is growing) to moderate, to light (little or sparce bulbil formation). 80 REFERENCES 1) WANI, M. C., H. L. TAYLOR, M. E. WALL, P. COGGON, and A. T. McPHAIL. LO Fass Plant antitumor agents, VI. The isolation and structure of taxol, a novel antileukemic and antitumor agent from Taxus brevifolia. J. Am. Chem. OS tego eoteled « 2) ROWINSKY, E. K., L. A. CAZANAVE, and F. C. DONEHOWER. 1990. Taxol: a novel investigational antimicrotuble agent. J. Natl. Cancersitnst. $2355124/—-1259- 3) UPADHYAY, R., D. KENFIELD, G. A. STROBEL, and W. M. HESS. 1991. Ascochyta cypericola causing leaf blight of purple nutsedge (Cyperus rotundus). Can. J. Bot. 69:797-802. 4) HESS, W. M. 1966. Fixation and staining of fungus hyphae and host plant root tissues for electron microscopy. Stain Technol. 41:27-35. 5) REYNOLDS, E. S. 1963. The use of lead citrate at high pH as an electron opaque stain in electron microscopy. J. Cell Biol’) 172208=212.. 6) SPURR, A. R. 1969. A low viscosity epoxy resin embedding medium for electron microscopy. J. Ultrastruct. Res. 20°31-43% 7) DeBARY, A. 1887. Comparative morphology and biology of the Fungi; mycetozoa and bacteria. (English translation). Clarendon Press, Oxford. 8) WEBER, D. J. and W. M. HESS. 1975). Diverse Spores of Fungi IN Spores Vol VI. Am. Soc. of Microbiol. p. 97- 111. 9) HESS, (Wo oM: 1981. Fungal Organelles and Other Cell Structures IN The Fungal Spore Morphogenetic Controls, G. Turian and H. R. Hohl (eds) Academic Press, N.Y. pp 21-41; 10) WERESUB, L. K., and P. M. LeCLAIR. 1971. On Papulaspora and bulbilliferous basidiomycetes Burgoa and Minimedusa. Cans. Bot 4932 208-221 3:. 11) COMPENDIUM OF SOIL FUNGI. 1980. Cladorrhinum (Sacc and Marchal), [{K. H. Domsch, W. Gams and T-H Anderson eds. ] Academic Press, N.Y. MY COTAXON Volume XLVI, pp. 81-85 April-June 1993 TWO NEW SOUTH AMERICAN SPECIES OF CORTICIACEAE (BASIDIOMYCETES) WITH AMYLOID SPORES KURT HJORTSTAM Malaregatan 12 S-441 35 Alingsas, Sweden and LEIF RYVARDEN University of Oslo, Department of Biology Division of Botany Box 1045 Blindern, N-0316 Oslo, Norway Abstract Two new South American species of Corticiaceae (Basidiomycetes), are described: Amyloathelia aspera Hjortst. & Ryv., with asperulate spores from Chile and Aleurodiscus exasperatus Hjortst. & Ryv. from Brazil. Amyloathelia aspera Hjortst.& Ryv. sp.nov. Fig. 1. Basidioma resupinatum, effusum, laxe adnatum, tenue, pelliculare. Hymenophorum leve, album vel cremeum, postea lutescens; margine indeterminato. Systema hyphale monomiticum; hyphae basales hyalinae, tenuitunicatae vel plerumque crassitunicatae, plus minus rectae, leves, 5-6 pm latae; hyphae subhymeniales tenuitunicatae, circiter 4 pm latae; hyphae omnes fibulatae. Cystidia nulla. Basidia subclavata vel plerumque tubularia, tenuitunicata, constricta et sinuosa, 30-35(-40) x 5-7 pm, 4 sterigmatibus. Sporae subglobosae, crassitunicatae, asperae, (6-)7-8(-9.5) x (5.5-)6.-6.5(-7) um, amyloideae, cyanophilae. Holotypus: Chile, Osorno, P.N. Puyehue, Antillanca, 1500 m., on dead branch in Nothofagus and Drimys winteri forest, 15 April 1988, Rajchenberg No. 4185 (BAFC). Isotypi: GB and O. BASIDIOME resupinate, effuse, loosely adnate, rather thin and pellicular. HYMENOPHORE smooth, at first white to cream-coloured, then pale yellowish, margin indeterminate or usually paler than the fertile part. HYPHAL SYSTEM monomitic; basal hyphae thin-walled to more commonly with a slight wall thickening or even thick-walled next to the substratum, hyaline, rather straight and long-celled, smooth, often branching at right angles, approximately 5-6 um wide, subhymenial hyphae comparatively short-celled, thin-walled, about 4 pm wide, all hyphae with clamp-connections. 82 Fig. 1. Amyloathelia aspera A) part of the hymenium, B) hyphae from the subiculum, C) spores, D) SEM of the spores, bar = 1 um. From the holotype. 83 CYSTIDIA absent. BASIDIA subclavate or when fully developed almost tubular, thin-walled throughout, constricted and sinuous, 30-35(-40) x 5-7 pm with four rather long (6-8 pm) sterigmata and a basal clamp-connection. SPORES subglobose, thick-walled, often with oil drops in the protoplasm, appearing completely smooth in Cotton blue, but slightly rugose in KOH as well as in Melzer’s reagent, (6-)7-8(-9.5) x (5.5-)6-6.5(-7) pm, distinctly amyloid and cyanophilous. The species is easily recognized by its thick-walled, asperulate spores. Its closest relative seems to be Amyloathelia amylacea (Bourd.& Galz.) Hjortst. which, however, has smooth, slightly longer spores and with a stronger amyloidity. The basidia of the latter are in general also shorter. It should be noted that Corticium vallum G.H. Cunn. was stated to be the same as A. amylacea by Staplers (1985). This must be a misinterpretation of A. amylacea since Stalpers described Cunningham’s species with gloeocystidia. A. amylacea (Corticium amylaceum) has no cystidia, only occasional hyphal ends between the basidia. Original material of Corticium vallum has not been studied in this case. Aleurodiscus exasperatus Hjortst. & Ryv. nov.sp. Big Basidioma resupinatum, arcte adnatum, tenue, quasi 0.1-0.2 mm crassum, plus minus coriaceum. Hymenophorum leve vel leviter rugosum, primum cremeum tum griseum vel dilute brunneolum. Systema hyphale monomiticum; hyphae distinctae, tenuitunicatae, modice ramosae, (2.5-)3-3.5 pm latae, sine fibulis. Gloeocystidia generaliter 50-80 x 8-10(-12) pm, tenuitunicata, sulphopositivan contentia. Dendrohyphidiae numerosae, inclusae vel plerumque projectae, ramosissimae, leviter incrustatae. Basidia clavata, grandia, generaliter 70-100 x 12-15 pm, 4 sterigmatibus. Sporae ellipsoideae vel aliquantum fusiformes, tenuitunicatae vel leviter crassiusculae, (15-)20-23 x 8-10 pm, exasperatae, amyloideae. Holotypus: Brazil, Sao Paulo, Santos, Cananeia, Ilha do Cardoso, 2-5 Feb. 1987, Hjortstam 16760. (K). Other specimen examined: Argentine: Tucuman, Ruta 38, km 19, no date indicated, leg. H. Gomez. Herb. BAFC 24547. BASIDIOME resupinate, closely attached to the substratum and hardly detachable, thin to moderately thick, approximately 0.1-0.2 mm thick, more or less coriaceous, confluent. HYMENOPHORE smooth or slightly pulverulent, cream-coloured or with a greyish tint, with age pale brown. Margin indeterminate. HYPHAL SYSTEM monomitic; hyphae thin-walled, moderately branched, (2.5-)3-3.5 um wide, forming a rather dense subiculum, all hyphae without clamp-connections. GLOEOCYSTIDIA generally numerous, with granular content, 50-80 x 8-10(-12) p", blackish in sulphovanillin, thin-walled, with an obtuse tip, basally often with several protuberances. DENDROHYPHIDIA numerous, projecting slightly above the basidia, richly branched, encrusted. BASIDIA clavate, large, 70-100 pm long or sometimes longer, with a width of 12-15 um just below the four stout sterigmata, without a basal clamp-connection. Fig. 2. Aleurodiscus exasperatus A) part of the hymenium, B) dendrohyphidia, C) acanthophyses, D) spores. From the holotype. 85 SPORES ellipsoid to somewhat fusoid, thin-walled or with slight wall thickening, (15-)20-23 x 9-12 pm, appearing smooth in KOH but slightly rugose in Melzer’s reagent, amyloid. Aleurodiscus exasperatus is similar to A. aurantius (Fr.) Schroet., but differs primarily by its spore shape and gloeocystidia. In the latter species the spores are ellipsoid, never fusoid and distinctly echinulate to verrucose. Further more, the gloeocystidia are moniliform at the apex. Macroscopically, the new species is reminiscent of species of Vuilleminia Maire, but easily separated from species of this genus by the amyloid spores and sulfo-positive gloeocystidia. Acknowledgements. We acknowledge assistance from Dr. R. Watling, Royal Botanic Gardens, Edinburgh, Scotland who has read the manuscript and suggested improvements. References. Stalpers, J.A. 1985. Type studies of the species of Corticium described by G.H. Cunningham. N.Z.Journ.Bot. 23:301-310. ore | ree ft + ¥; 7 ihe Nig AO Te ) MA BMA MYCOTAXON Volume XLVII, pp. 87-92 April-June 1993 DEMATIACEOUS HYPHOMYCETES ON FREYCINETIA (PANDANACEAE). 4. CRYPTOPHIALE E.H.C. McKENZIE Herbarium PDD, Landcare Research Private Bag 92170, Auckland, New Zealand and A.J. KUTHUBUTHEEN Department of Botany, University of Malaya, 59100 Kuala Lumpur, Malaysia Abstract Two new species of Cryptophiale are described and figured, C. caudata from Malaysia and C. novae-caledoniae from New Caledonia. These fungi were found during a study of dematiaceous hyphomycetes on dead leaves of Fr- eycinetia. Introduction During an examination of over 70 samples of dead Freycinetia leaves from nine countries (McKenzie 1991), only two collections of Cryptophiale were found. Both of these, one from New Caledonia and the other from Malaysia, are distinct new species and are described and figured in this paper. Cryptophiale caudata McKenzie et Kuthubutheen sp. nov. Fig. 1 Coloniae conspicuae, effusae. Conidiophora e cellulis atro-fuscis, macrone- matosa, mononematosa, singula, eramosa, erecta, apicem versus parum curvata, laevia, septata, brunnea, apice acuta, usque ad 230 »m longa, ad basim 7-10 ym lata, ad apicem 5-7.5 ym. Zona fertilis subapicalis, scutel- liformis, 45-60 »m longa, 15-24 .m crassa; cellulae conidiogenosae scutello cellarum sterilium pallide fusco celatae; cellulae steriles 2.5-5 .m latae, extendentes 6-9 um ultra conidiophororum parietem. Conidia hyalina, ad- herentia, clavata vel fusiformia, subulata, laevia, guttulata, eseptata, (20-) 23-26 (-28) x 4-4.25 zm, apicem versus 10-25 ym longa, basi obconice trun- cata. Figure 1 Cryptophiale caudata, conidiophores and conidia (PDD 60584 - type). Specimen mounted in hydrous lactophenol. 89 In foliis mortuis pandanaceae Freycinetia sp. Holotypus PDD 60584. Colonies conspicuous, effuse. Conidiophores arising from dark brown, smo- oth, swollen cells, macronematous, mononematous, single, unbranched, erect, slightly curved towards the apex, smooth, septate, brown, abruptly tapered to an acute apex, up to 230 »m long x 7-10 «mm thick near the base, tapering to 5-7.5 um near the apex. Fertile region subapical, 45-60 »m long, 15-24 »m wide. Conidiogenous cells obscured by a shield of sterile, pale brown cells; each shield cell 2.5-5 um wide, extending 6-9 um beyond the conidiophore wall. Conidia produced in slime and adhering to the fertile part of the conidiophore in a lateral droplet; hyaline, clavate or fusiform, subulate, smooth, guttulate, not appearing septate, (20-) 23-26 (-28) x 4- 4.25 «.m, tapering to form a single, 10-25 ym long flexuous appendage at the distal end, base long, obconically truncate. On Freycinetia sp. Specimen examined: Malaysia, Selangor, Genting Highlands, Gunung Bunga Buah, 18.VIII.1992, E.H.C. McKenzie (PDD 60584 - holotype). The specific epithet refers to the tail-like appendage on the conidia. Cryptophiale novae-caledoniae McKenzie et Kuthubutheen sp. nov. Bigio2 Coloniae conspicuae, effusae. Conidiophora e cellulis atro-fuscis, macrone- matosa, mononematosa, singula, eramosa, erecta, recta vel flexuosa, lae- via, septata, obscure rubro-brunnea, apicem versus parce pallidiora, acuta, usque ad 450 ym longa, ad basim 10-11 ,m lata, ad apicem 4-6.5 ».m. Zona fertilis apicalis, scutelliformis, 35-80 «.m longa, 13-14 ym crassa; cellulae conidiogenosae scutello cellarum sterilium pallide fusco celatae; cellulae steriles 2.5-3.5 «m latae, extendentes 3-4 »m ultra conidiophororum pari- etem. Conidia hyalina, adherentia, fusiformia, laevia, medio septata, (9.5-) 14-16 (-18) x 2-2.25 um, apice rotundata vel acuta, basi rotundata vel ob- conice truncata. In foliis mortuis pandanaceae Freycinetia sp. Holotypus PDD 60585. Colonies conspicuous, effuse. Conidiophores arising from dark brown, smo- oth, swollen cells, macronematous, mononematous, single, unbranched, erect, straight or flexuous, smooth, septate, dark reddish-brown, slightly paler towards an acute apex, up to 450 »m long x 10-11 ym thick near the base, tapering to 4-6.5 «m in the fertile region. Fertile region apical, 35-80 »m long, 13-14 wm wide. Conidiogenous cells obscured by a shield of sterile, pale brown cells; each shield cell 2.5-3.5 um wide, extending 3-4 ».m beyond the conidiophore wall. Conidia produced in slime and adhering to 90 BrceeadaeAre ay oT yuna canis HES FOB BEDII0+60C2: 0-255 eS ne pe Re == Soe Se Se <= fees es conidiophores and conidia (PDD d in hydrous lactophenol. 1ae, -caledon Figure 2 Cryptophiale novae 60585 - type). Specimen mounte 91 the fertile part of the conidiophore in a lateral droplet; hyaline, fusiform, smooth, medianly 1-septate, (9.5-) 14-16 (-18) x 2-2.25 ,m, apex rounded or acute, base rounded or obconically truncate. On Freycinetia sp. Specimen examined: New Caledonia, Mt Panié, 1500m, 15.XII.1990, J.S. Dugdale (PDD 60585 - holotype). The specific epithet refers to the country where the species was found. Sutton et al. (1989) provided a key to nine species of Cryptophiale then known; a tenth species, C. secunda Kuthubutheen & Sutton having been previously transferred to Cryptophialoidea Kuthubutheen & Nawawi (19- 87). McKenzie (1993) has recently described a new species, C. insularis, from New Zealand. C. caudata and C. novae-caledoniae can be distin- guished from other species in the genus by the structure of the conidiophore apex, size of conidia, and position of the fertile region on the conidiophore. C. caudata is similar in several respects to C. aristata Kuthubutheen & Sutton (1985) and to C. enormis Sutton, Nawawi & Kuthubutheen (1989). The apex of the simple, setiform conidiophore in all three species extends not more than 10 »m beyond the fertile region. The conidiophore of C. enormis, however, is wider at the fertile region than above or below it. The conidia in all three species are caudate. The conidia of C. aristata are 23-27 x 2-3 «.m and 1-septate and the apical cell is drawn into a filiform appendage 7-12 »m long. Although the conidial length of C. caudata and C. aristata is similar, in C. caudata the conidia are broader, 0-septate, and the apical flexuous appendage is 10-25 »m long. The conidia of C. enormis, however, are twice as large as those of C. caudata, 4-7 septate, and have shorter (5-8 .m) appendages. C. novae-caledoniae is reminiscent of C. kakombensis Pirozynski (1968), a species widely distributed in the tropics and subtropics. The conidia of C. kakombensis, however, are falcate, longer but narrower (22-28 x 1-2 ,m), and acute at both ends compared to the conidia of C. novae-caledoniae. The fertile region of C. kakombensis is subapical and not apical as in C. novae- caledoniae. The apical location of the fertile region in C. novae-caledoniae is unique and not found in any of the known species in the genus. Acknowledgment We gratefully acknowledge the assistance of Dr G. Kuschel, Auckland, in preparation of Latin diagnoses. We thank Professor Gareth Morgan-Jones, Department of Plant Pathology, College of Agriculture, Auburn University, Alabama, USA for critical review of this paper. 92 References Kuthubutheen, A.J.; Nawawi, A. 1987: Cryptophialoidea gen. nov. on decaying leaves from Malaysia. Transactions of the British Mycological Society 89: 581-583. Kuthubutheen, A.J.; Sutton, B.C. 1985: Cryptophiale from Malaysia. Tran- sactions of the British Mycological Society 84: 303-306. McKenzie, E.H.C. 1991: Dematiaceous hyphomycetes on Freycinetia (Pan- danaceae). 1. Stachybotrys. Mycotaxon 41: 179-188. McKenzie, E.H.C. 1993: New hyphomycete species from litter in the Chat- ham Islands, New Zealand. Mycotaxon 46: 291-297. Pirozynski, K.A. 1968: Cryptophiale, a new genus of hyphomycetes. Cana- dian Journal of Botany 46: 1123-1127. Sutton, B.C.; Nawawi, A.; Kuthubutheen, A.J. 1989: Additions to Belem- nospora and Cryptophiale from Malaysia. Mycological Research 92: 354- 358. MYCOTAXON Volume XLVII, pp. 93-100 April-June 1993 GENETICS OF A TUBER AESTIVUM POPULATION (Ascomycotina, Tuberales) Gi, PACIONE 7 3Ge(ERLZAL 7 M. MIRANDA and C. VISCA Dipartimento di Scienze Ambientali and i Dip Biologia& Fisiologia Cellulare, Universita, 67100 L'Aquila, Italy Key-words: Tuberales, Tuber aestivum, Tuber uncinatum, truffle, allozymes, reproduction, sibling species. Abstract: A truffle population with of Tuber aestivum and Tuber uncinatum morphotypes was subjected to allozyme analysis via SGE and genotypically scored at fourteen loci. Ten of the loci were monomorphic and four polymorphic. Thus, seventeen new genotypic classes, with very low genetic differentiation, were found. However, heterozygosity was never found and all alleles seem to be fixed in individuals. These results indicate that Tuber aestivum and T. uncinatum are probably two extreme morphological forms of a complex of self-fertilizing sibling species. Introduction Some taxonomic complexes of the genus Tuber Mich. ex Fr. are very difficult to distinguish on a morphological basis. For Tuber aestivum Vitt., a European taxon, also found in Algeria and Turkey, there seemsjito be a’ long list,of probable synonyms. Beyond the nomenclatural problems, the difficulty in distinguishing species has produced confusion in regulating and marketing these species worldwide. CHEVALIER et al. (1978) in an attempt at’ separating Tuber uncinatum Chat. as a distinct species, tried without much success to juxtapose its morphological characters with those of T. aestivum. GROSS (1987) proposed to divide these two taxa on the basis of the height of the sporal reticulum. Moreover, in two previous reports, no heterozygosity was found for several loci in a number of Tuber species (PACIONI & POMPONI, 1989; 1991). An investigation at the population 94 level, to test the hypothesis that a selfing process of reproduction may be responsible for the homozygosity, therefore appears of some interest. This paper reports the results of using allozyme analysis to resolve the status of Tuber aestivum and T. uncinatum. Materials and Methods A sample of 66 specimens, morphologically related to Tuber aestivum and Tuber uncinatum, was collected during 1990 and 1991 ‘inthe, } bei Poz2ze"," aysmail) trutrie preserve; near che village of Santi, in’ LD Aguilav(The Abruzzi »yltalyysithe specimens, determined by the senior author, have been preserved, one portion cut in little sticks and then deep- frozen and a small portion dried and deposited in the mycological herbarium of L'Aquila (AQUI) as a voucher. Horizontal starch gel (SGE) was performed essentially according to PACIONI & POMPONI (1989) and FRIZZI & TAMMARO (1991). Some slight modifications were made in the staining solutions. The stains were basically prepared according to HARRIS (1966), exceptufor the’ SKDH. stain) (FINESCHI, 1983). Electrophoresis was conducted at a constant 8V/cm for approximately 3.5 hours per run. Details on the electrophoretic techniques used are given in Wavie Wis A preliminary survey showed fourteen enzymatic loci produced patterns clearly interpretable from a genetic point of view: Pgm-2 (PGM, phosphoglucomutase), G6pd (G6PD, glucose-6-phosphate dehydrogenase), 6-Pgd (6PGD, phosphogluconate dehydrogenase), Hk (HK, hexokinase), Mpi-1l (MPI, mannose phosphate isomerase), Ak (AK, adenylate kinase), Pgi (PGI, glucose phosphate isomerase), Mdh-2 (MDH, malate dehydrogenase), Me-1 and Me-3 (ME, malic enzyme), Idh (IDH, isocitrate dehydrogenase), Sod-1 (SOD, superoxide dismutase), Skdh (SKDH, shikimic dehydrogenase). With a view to comparing the resulting elecrophoretic data to all known data for Tuber, the previous reference for the mobility of the enzymes of Tuber melanosporum Vitt. (=100) was mantained. Electrophoretic data were analysed using the NTSYS-pc program (ROHLF,1987) and genetic divergence was estimated with the index of standard distance (D) proposed by NEI CL9I2)> Results Nine of the fourteen enzymatic loci examined ( G6pd, 6Pgd, Adk, Got, Pgi, Me-3, Skdh, Mdh-1, Sod) are monomorphic 95 showing the same phenotype as one band. The Pgm-2 locus is also monomorphic, however it is distinguished by three bands. The Pgm-2 pattern has been reported for all other taxa of Tuber already examined (PACIONI & POMPONI, 1989; 1991). The cause of the Pgm-2 pattern is still unexplained. The relative mobility of the above enzyme electrophoretic patterns are: 6Pgd 103, Adk 100, Got 105, Pgi 100, Me-3 110, Skdh 100, Mdh-1 100, Sod 93, Pgm-2:85. Only LOurgocie(Hk:, Mpi-1, Me-1, IdH) appeared polymorphic. At locus Hk three alleles (108; (110; 7112). were found, two. (95),..100:) at. Mpi-1, two (105, 110), at \Me-l*and’three (90, 95, 100) at.iIdh. Heterozygosity was not found and all alleles seem to be fixed individuals. The electrophoretic patterns revealed seventeen genotypic classes (ROYSE & MAY, 1982). Only the four polymorphic loci are reported (Table 2). Their genetic distances are very LOW ithe SiO), Oi 4 oe OmMEOr 6s SOK) IN LOD Un Ce Vales moti an intraspecific variability, confirming the overlapping of the morphocharacters. Figure 1 shows one of the possible clustering trees. The enzyme phenotype, belonging to the genotypic class labeled TA2, seemed to be the most common. Several phenotypes were only observed once. The number of phenotypes found is almost equal to that of the truffle habitats sampled. For loci in common, the genotypic classes found here are completely different from those previously observed in samples from other Italian localities (PACIONI & POMPONI, L991). Discussion The homozygosity of truffles has been reported previously in the studies on the Tuber aestivum-T. mesentericum group (PACIONI & POMPONI, 1991) as well as on several other taxa of Tuber (PACIONI & POMPONI, 1989). However vegetative cells of the truffle sporophores contain a pair of nuclei (FASOLO- BONFANTE & BRUNEL,1972) and therefore a certain percentage of heterozygosity should be present, unless the two nuclei arise from self-fertilization or mitosis. Our results strongly support the hypothesis that in dikaryotic hyphae, the two nuclei result from selfing; accordingly non-heterozygosity is observed. This is at variance with what is known about truffle reproduction, hitherto supposed to be heterothallic (DANGEARD 1895/96; GRENTE et al., 1972). Sexual reproduction by selfing would restrict, in this group of) Lungijevariabi laity toh tnter=and intrachromosomal recombination and mutation. As stated in the Results, only 96 four of fourteen loci examined appeared polymorphic and this may favour the hypothesis that a strong selection, due to the, peculiaryjecology; and ‘lute cycles ofs:trufives /ihas> been operating on the ten monomorphic loci, whereas it has been less stringent on the polymorphic loci. It should be noted, however, that, rnytrutites, very little inftormationeis available about the structures, functions and molecular properties’ of thetenzymes, anvéestigated here. jin tact, ethe polymorphism observed might be ascribed to alleles of the same locus, to alleles of entirely different loci differentially expressed in the various samples, or to conformational variants (DAWSON, 1972) of the same locus expressed in different sporophores. In the light of the present results, we think that the sporophores of Tuber aestivum or T. uncinatum represent only two extreme forms, at present unresolvable by morphological and biometrical methods. The existence of a great number of sibling species suggests that .oun knowledge! otmttber jreproduetionsismstallyvery scanty and requires more investigation. Research carried out with financial support from M.U.R.S.T., Scientific Research 40% "Ecology and biology of fungi and lichens.’ We are! andebbedi to Prof. 68.) Grilbis for his assistance with the English manuscript and Prof. R.D. Fogel, University of Michigan, for his suggestions. References BREWER,G. & SING,C.F., 1970. An introduction to isozyme techniques. London & New York: Academic Press. CHEVALIER; -G., DESMAS; C., FROCHOT, H. & RIOUSSET;, L:, 1978. L'éspece Tuber aestivum Vitt. : I. Definition. Mushroom SciencelO: 957-975. DANGEARD, P.A., 1895/96. La Truffle. Recherches sur _ son développement, sa structure, sa reproduction sexuelle. Botaniste 4/5:63-87. FASOLO-BONFANTE,P. & BRUNEL,A., 1972. Caryological features in a mycorrhizal fungus: Tuber melanosporum. Allionia 18: 5-11. FPINESCHI) (52% P983kiVariabilitay untraspecificaritiny /einus nigra, Arn... 4 iRisultatiervd)? Ganalisi ) sugvalcuni ssisteni isoenzimatici. L'Italia Forestale e Montana 38: 200-213. FRIZZI, G. & TAMMARO, F.; 1991. Electrophoretic study “and genetic affinity sin ‘the Campanula elatines and Gx fragilis (Campanulaceae) rock-plants group from Italy and W. Jugoslavia. Plant Syst. Evol. 174: 67-73. o7. GRENTE,J., CHEVALIER, Geese POLLACSER en 9:7 or La germination de l'ascospore de Tuber melanosporum et de la synthése sporale des mycorrhizes. C.R. Acad. Sc. Paris 21 Ot ea 3. GROSS, G., 1987. Zur den europaischen Sippen der Gattung Tuber. in,DERBSCH,H.,& SCHMITT, J.A., (Eds.):Atlas der Pilze des Saarlandes,2: Nacweise, Okologie, Vorkommen und Beschreibungen, pp.79-99. Saarbrucken, Germany: Ed. Delattinia. HARRIS, H., 1966. Enzyme polymorphism in man. Proc. Roy. soc. London B69 *% 3298-31107 NEI, M., 1972. Genetic distance between populations. Americ. NaG. 61062 (233-292. PACLONI7: 9G. &, POMPONI, G:, "1989. ‘Chemotaxonomy of ‘some Italian species of Tuber. Micol. Veget. Medit. 4: 63-72. PACIONI, G. & POMPONI, G., 1991. Genotypic patterns of some Italian populations of the Tuber aestivum-T. mesentericum complex. Mycotexon 42:3, 171-179. ROHLF, estllinan 1987. NTSYS-pc Numerical Taxonomy and Multivariate Analysis System. Setauker, N.Y., U.S.A.: Exeter Publishing. ROTHE, G., 1973. Shikimisaure-Dehydrogenase in keimenden Erbsen. Biochem. Biophys. Pflanzen 164: 475-486. ROYSE, 7 Di.- & MAY, 9B.) 61982. Use of, tseozyme: ‘variation to identify genotypic classes of Agaricus brunnescens. Mycologia 74: 93-102. SiOUANDER gk. Ws 2 OM LIn. jo MH. @YANG Sy, GONHSON, Wok. o& GENTRY, J.B., 1971. Biochemical polymorphism in the genus PeromnyScusi tl. a Variation pin. ene old field mouse (Peromyscus polionotus). Stud. Genet. 6: 49-90. SHAW, C.R. & PRASAD, R., 1970. Starch gel electrophoresis of enzymes. A compilation of recipes. Biochem. Genet. 4: 297-320. (OL6T ‘AWSWUd ¥% MVHS) O3Vet0G/euUesSteA/STIL-F S(IL6T ‘4°72 39 YAANVTS) II snout quoo aexeaqtTo/stan -e€ !(OL6T ‘ONIS ¥ UAMAUA) SQeeTeUW/STIAL -Z !(OL61 ‘AWSWUd B MWHS) e2etTOq/wNTYATT -T :pesn swaeqsks rez jng (€L6L) AHLOU 0°8 € Ge. abl Haus (OL6L) DNIS ¥ UAMAUE poke ‘: 6ePoceiS IDd (9961) SIUYUVH Q232 v Qe T EKG IdW (OL6L) DNIS ¥ UNMAUE Coal: Z [eng Siac Wd (OL6L) DNIS ¥ MAMA Cze/, Z Si Fon ard YW (OL6L) GWSWud ¥% MVHS 0°sS v Ter ssa /oucG YH (ITL6L)°Te@ 8% UAAGNWIES 0°8 € thea bose pnts LOD (9961) SIUUVH Z*8 : Tels ee aos (OL61T) DSNIS ¥ UAMHUG G%H T Cie a qd99 (OL61L) GWSWud ¥ MVHS 0°s8 i Vuela eee q9d9 (OL61T) DNIS ¥ YAMANE Z°8 T ha Ae MES HAdI (9961) SIUYUYWH £58 T OF bes AW (OL61T) DNIS ¥ UAMAUE Cael: Z Ges ee tea HAW seouertejel Hd Zezszgngq et Ghd Ul ges) cel sowAzuso “JATA WNATQZSEe Taqny, Ut sowdzo[Te pue sewkzost Jo wuoTtTAOeqjep AOT suoTRIpuCo ADS “*T eTqeL GOT/SOT G6/S6 Ostet 20 salen 00220 0p ee el Wd, GiOieS 0 (00/007. 011701001 00a oO LW di GOT/SOT G6/S6 OLT/OTT 06/06 (1L)STWa Oder G6/S6 OLE L0 LE 06/06 (2) oTWa OseaC te G6/G6 S007 80 00a 00 beat) &LWL OileLOeeL S6/GS6 Ole /-0A-b 0 Oe/-0.0 alae): CE OAS /207E:1 G6/G6 80T/80T G67 G6. SCG LL OREAOTT G6/S6 OLTAOLE G6/S6 (L)OTWL Ober 12. 001/001. 01140 TE 06/06 (7) 6WL OOLI- 001/00 Teac li fcit 06/06 (€) SWL OFL/OL1- -O00T/00Le S0E/S0T 06/06 (LT) LW Oqee/s0 le. 001001 =ClEAcle O0l,00l = OU OTE/OLE 0017001 011 /0TE— 001/007 (9) S¥a SOA25 0 La 001L/001= =01 TAO LE G6/S6 (Ll) Pwo Ors 08 0 Ot 00 tea deca | S6/S6 (Z)€WL One Ose 0b 70 0nee Oi 0 G6/S6 (€Z2Z)tWoL OMT 0 019200 te 90.29 0 G6/S6 (>) TWo [-OW T-TawW 4H YpI (sqeyoeriq uT umoYsS Ssuswtoeds Jo Toequinu 9Y44)‘AATA WNATQZSae Taqny fo uotqzetndod T[einjeu e UT peaAazZesqo (WL) sessetTo otdAjoush oy Fo T90T otydzowATog °zZ eTqeL 100 “IATA wNATZSee Teqny, JO sesseTo oTJeueb usejUSAES 2uq JOF seoueqstp OTJeueb s,TeN ey FO weiboipuegq - T°hta S00 OL'O SltO 020 G20 MY COTAXON Volume XLVI, pp. 101-129 April-June 1993 NEW SPECIES IN THE LICHEN FAMILY PARMELIACEAE (ASCOMYCOTINA) FROM AUSTRALIA JOHN A. ELIX Department of Chemistry, The Faculties, Australian National University, GPO Box 4, Canberra, ACT, 2601, Australia ABSTRACT : The species, Bulbothrix microscopica Elix, Canoparmelia herveyensis Elix, Canoparmelia subarida Elix, Canoparmelia whinrayi Elix, Flavoparmelia kantvilasii Elix, Hypotrachyna booralensis Elix, Hypotrachyna proserpinensis Elix, Hypotrachyna subpustulifera Elix, Neofuscelia kondininensis Elix, Neofuscelia imitatricoides Elix, Neofuscelia parasitica Elix, Neofuscelia scabrosina Elix, Neofuscelia subbarbatica Elix, Paraparmelia yamblaensis Elix, Parmelia crowii Elix, Parmelina euplectina Elix, Parmelina johnstoniae Elix, Parmelinopsis protocetrarica Elix, Parmotrema submerrillii Elix, Xanthoparmelia austroconstrictans Elix, Xanthoparmelia canobolasensis Elix, Xanthoparmelia hyposalazinica Elix, Xanthoparmelia masonii Elix and Xanthoparmelia trirosea Elix are described as new. A new name is given to Xanthoparmelia xanthofarinosa Elix (formerly xX. xanthosorediata). The following new combinations are made: Bulbothrix queenslandica (Elix & Stevens) Elix, Bulbothrix subtabacina (Elix) Elix, Canoparmelia owariensis (Asah.) Elix, Canoparmelia pustulescens (Kurok.) Elix, Canomaculina melanochaeta (Kurok.) Elix, Imshaugia evernica (Elix & Johnst.) Elix, Parmotrema pseudovirens (Gyelnik) Elix and Rimeliella haitiensis (Hale) Elix. Introduction As a result of further study of the lichen family Parmeliaceae in Australia (Elix & Johnston 1986, 1987a, 1987b, 1987c, 1988a, 1988b, 1988c; Elix, Johnston & Armstrong 1986; Nash & Elix 1987), I am describing 24 new species. Throughout the present work chemical constituents were identified by thin layer chromatography ( Culberson 1972; Culberson, Culberson & Johnson 1981; Culberson & Johnson 1982; Elix, Johnston & 102 Parker 1987, 1988), high performance liquid chromatography (Lumbsch & Elix 1985) and comparison with authentic samples. Bulbothrix microscopica Elix sp. nov. Fig. 1 Species cum thallo ut in Bulbothrix subtabacina sed ab hac specie thallo diminuto, lobis angustioribus, emaculatis et ciliis inflatis differt. Type: Australia. Queensland. On mangroves along the foreshore of Tin Can Inlet; Tin Can Bay, 25°54'S, 153°0V'E, Tom iy. Av Elix 22862) 3.vili 1989; holo: CBG. Thallus corticolous, adnate to tightly adnate, mineral grey, 1-2 cm diam. Lobes sublinear, 0.1-0.5 mm wide, subdichotomously branched, margins densely bulbate-ciliate, bulbate cilia shiny, markedly inflated. Upper surface plane, emaculate, moderately to densely isidiate, the isidia . simple, less than 0.15 mm high. Medulla white. Lower surface black shiny, moderately to densely rhizinate to the margins, rhizines black, simple or tufted at apices. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow, medulla K+ yellow-red, C-, P+ orange; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (minor). The minute thallus, very narrow lobes, simple isidia, black lower surface and medullary salazinic acid distinguish this species. P. subtabacina Elix a related species (vide infra), has broader (0.5-1.0 mm wide), sublinear- elongate lobes with a distinctly maculate upper surface, slender, elongated bulbate cilia which are commonly branched subapically and dichotomously branched rhizines. Currently this rare endemic species is only known from the type locality. Canoparmelia herveyensis Elix sp. nov. Fig. 2 Species cum thallo ut in Canoparmelia raunkiaeri sed ab hac specie medulla inferiore pigmento ochraceo et euplectino continente differt. Type: Australia. Queensland. On granite rocks in dry sclerophyll forest, Hervey Range, 45 km south-west of Townsville, 19°26'S, 146°24’E, 350 m, J. A. Elix 20427 & H. Streimann, 20.vi.1986; holo: CBG; iso: MEL. Thallus adnate, saxicolous, pale mineral grey, 5-8 cm diam. Lobes subirregular, apically rotund, 2-4 mm wide. Upper surface plane, becoming irregularly rugose towards the centre, pustulate-sorediate, the pustules coalescing in a coarsely sorediate mass in the older portions of the thallus. Medulla white for the most part but pigmented yellow-orange to orange- brown adjacent to lower surface. Lower surface black, sparsely rhizinate, 103 rhizines simple, with a narrow bare, brown zone at the margins. Apothecia and pycnidia not seen. Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P+ orange-red, lower medulla K+ violet; containing atranorin, chloroatranorin, protocetraric acid (major), virensic acid (trace), euplectin (minor). This new species resembles Canoparmelia raunkiaeri (Vainio) Elix & Hale as both develop pustulate soredia towards the centre of the thallus and contain medullary protocetraric acid. However, C. herveyensis, is clearly distinguished by the partially orange-yellow pigmented lower medulla due to the presence of the anthraquinone, euplectin. At present this new species is only known from the type locality where it appears quite common. Canoparmelia subarida Elix sp. nov. Figs 3 Species cum thallo ut in Canoparmelia macrospora sed ab hac specie sporis brevioribus et acido lecanorico continente differt. Type: Australia. Western Australia. On twigs, South Stirling sand plain, 4 km off the Chillinup road, N. Sammy UWA8, 12.viii.1970; holo: PERTH. Thallus corticolous, adnate, pale mineral grey, 1-3 cm in diam. Lobes subirregular, crowded and imbricate, 1.0-2.0 mm wide. Upper surface smooth and shiny at the apices, plane or weakly foveolate at first, becoming wrinkled and rugose with age, lacking soredia and isidia. Medulla white. Lower surface ivory to pale brown, moderately to densely rhizinate, rhizines concolorous with the lower surface, simple or tufted at the tips. Pycnidia numerous, punctiform, black; conidia cylindrical 5-6 x 1 um. Apothecia common, sessile, 0.5-2 mm in diam., disc concave at first but becoming flattened with age, chestnut brown to dark brown, epruinose; margin thin, entire, smooth; spores (8), colourless, ellipsoid, 7-9 x 5-6 um. Chemistry. Cortex K+ yellow, medulla K-, C+ red, P- ; containing atranorin, chloroatranorin, lecanoric acid (major), orsellinic acid (trace), unknown (trace). This species is one of the few species of Canoparmelia with a pale lower surface. In gross morphology C. subarida closely resembles C. macrospora Elix & Johnston, but the latter species is distinguished by the maculate apical lobes which are densely pruinose, the large (16-20 x 5-6 um) often curved spores and medullary scabrosin 4,4'-diacetate. C. subarida is chemically identical to C. pruinata, but the latter species is distinguished by the much larger (4-15 mm), pruinose apothecia and a black lower surface. C. subarida appears to be a rare species in the subarid inland areas of southern Australia. 104 Figures 1-4. New species of Parmeliaceae: 1, Bulbothrix microscopica (holotype in CBG); 2, Canoparmelia herveyensis (holotype in CBG); 3, Canoparmelia subarida (holotype in PERTH); 4, Canoparmelia whinrayi (holotype in MEL). Scale bar = 5 mm. 105 Specimens Examined VICTORIA. On twigs, W side of Wyperfeld National Park, along fire access trail, c. 31.55 km W of Wonga Hut, R. Filson 16024, 29.viii.1978 (MEL). SOUTH AUSTRALIA. On bark of Acacia aneura, 20 km W of Vokes Corner, 28°34'S, 130°29'E, N. N. Donner 7376, 23.viii.1980 (AD). Canoparmelia whinrayii Elix sp. nov. Fig. 4 Species cum thallo ut in Parmelina labrosa sed ab hac specie thallo subto pallido et cilia desunt differt. Type: Australia. Tasmania. On twigs, north-west of Trig Point, Flinders Island, John Whinray, 1.i1.1969; holo: MEL. Thallus corticolous, adnate, olive-grey to mineral grey, c. 7 cm diam. Lobes subirregular and apically subrotund, 3-5 mm wide, lacking cilia. Upper surface plane to wrinkled, foveolate, shining or rather waxy, sorediate; soralia laminal or marginal, white, becoming grey-brown with age, originating from coarse pustular ridges, spreading over the thallus. Medulla white. Lower surface ivory to pale tan, moderately rhizinate, rhizines simple, concolorous with lower surface. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow, medulla K-, C+ red, P-; containing atranorin, chloroatranorin, lecanoric acid (major). This unique new species is distinguished by the broad lobes with a pale lower surface, the pustular soralia, the medullary lecanoric acid and the absence of pseudocyphellae and cilia. It is the only sorediate Canoparmelia with a pale lower surface so far recorded for Australia. In overall growth form and in the development of soralia, this new species resembles Parmelina labrosa (Zahlbr.) Elix & Johnston, but the latter is distinguished by the sparse marginal cilia and the black lower surface. At present this species is only known from the type collection. Flavoparmelia kantvilasii Elix sp. nov. Fig. 5 Species cum thallo ut in Flavoparmelia euplecta sed ab hac specie thallo diminuto, medulla albida, inferiore nonpigmentifera et euplectino noncontinente differt. Type: Australia. New South Wales. On twigs of Lomatia arborescens in Eucalyptus pauciflora woodland, Thunderbolt Lookout, Barrington Tops National Park, 25°54'S, 153°01'E, 1450 m, G. Kantvilas 426/88, 4.vii.1988; holo: HO, iso: NSW, ANUC. 106 Thallus corticolous, adnate, green to yellow-green, 2-4 cm in diam. Lobes irregularly rotund, incised, margins crenulate, 2.0-5.0 mm wide; secondary lobes narrower, overlying the marginal lobes. Upper surface plane, matt, smooth at the marginal lobes becoming slightly rugulose towards the centre of the thallus, isidia absent, pustulate-sorediose; soralia sometimes developing from cracks or pustules which burst to form granular soredia. Medulla white. Lower surface jet black, with a pale brown bare zone at the lobe ends, sparsely rhizinate; rhizines simple or fasciculate. Pycnidia and apothecia not seen. Chemistry. Cortex K-, medulla K-, C-, P+ red; containing usnic acid, protocetraric acid (major), atranorin (+), caperatic acid, virensic acid (¢ trace), unknown (+ trace). Morphologically this species closely resembles Flavoparmelia euplecta (Stirton) Hale as both species have adnate thalli with a pustulate sorediate upper surface and contain protocetraric acid as a major medullary constituent. However the thalli of F. kantvilasii are smaller (2-4 cm cf. 4-12 cm) and more fragmented, not only when growing on its normal twig habitat but even when found on broader branches. Furthermore F. kantvilasii lacks the orange pigmented lower medulla (K+ purple, containing euplectin) so characteristic of F. euplecta. F. kantvilasii is found in the temperate forests of northern New South Wales. Specimens Examined NEW SOUTH WALES. On Ficus in coastal Eucalyptus forest, Telegraph Point, 25 km NNW of Port Macquarie, 31°19'S, 152°48'E, K. L. Gaul, viii.1986 (ANUC); on canopy twigs of Nothofagus moorei in rainforest, Burraga Swamp, Mt Allyn Forest Park, Chichester State Forest, 32°06'30"S, 151°25'30"E, 1000 m, G. Kantvilas 187/88, 29.vi.1988 (ANUC, HO, NSW); on canopy twigs of Nothofagus moorei in rainforest, Barrington Tops National Park, Mt William, 1400 m, G. Kantvilas 307/88, 30.vi.1988 (HO, NSW); on canopy twigs of Nothofagus moorei in rainforest, Gloucester Tops, 1150 m, G. Kantvilas 414/88A; 2.vii.1988 (HO, NSW); on twigs of Nothofagus moorei in rainforest, Tweed Range, Wiangerie Forest Drive, Antarctic Beech Walk, 1000 m, G. Kantvilas 703/88, 3.viii.1988 (HO, NSW); on canopy twigs of Nothofagus moorei, New England National Park, Cascade Creek near Wright's Lookout, G. Kantvilas 705/88, 6.viii.1988 (HO, NSW). Hypotrachyna booralensis Elix sp. nov. Fig. 6 Species cum thallo ut in Hypotrachyna densirhizinata sed ab hac specie thallo multo parviore et lobi angustiore differt. Type: Australia. New South Wales. On bark, in a bushland gully, 3 km west of Booral, Rex Filson 16764, 1.ii.1969; holo: MEL. 107 Thallus corticolous, adnate, whitish-grey to mineral grey, c. 2.5 cm diam. Lobes sublinear, 0.5-0.8 mm wide, dichotomously branched, separate, the margins subcrenate. Upper surface shiny on younger lobes, dull and irregularly cracked on older lobes, plane, emaculate, sorediate; soralia orbicular, mostly subterminal, capitate. Medulla white. Lower surface black, densely rhizinate, rhizines black, moderately dichotomously branched. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow, medulla K-, C-, KC+ rose, P-; containing atranorin, chloroatranorin, alectoronic acid (major). This new species resembles the American-African species Hypotrachyna densirhizinata (Kurok.) Hale in that both are sorediate with capitate soralia and contain alectoronic acid. However H. densirhizinata is clearly distinguished by the size of the thallus (6-20 cm diam.), the breadth of the lobes (2-6 mm wide) and the presence of additional a-collatolic acid. At present this new species is only known from the type collection. Hypotrachyna proserpinensis Elix sp. nov. Fig. 7 Species cum thallo ut in Hypotrachyna heterochroa sed ab hac specie thallo laxe adnato, medulla albida, inferiore nonpigmentifera et acido gyrophorico continente differt. Type: Australia. Queensland. On granite rocks on edge of escarpment in Eucalyptus-Casuarina dominated woodland, Clarke Range, 46 km south of Proserpine, 20°50'S, 148°32'E, 800 m, J. A. Elix 20929 & H. Streimann, 29.vi.1986; holo: CBG; iso: MEL. Thallus saxicolous, loosely adnate, whitish mineral grey, 5-9 cm diam. Lobes sublinear, contiguous, 1-2 mm wide, subdichotomously branched, apices incised, margins entire, axils sinuous. Upper surface plane, continuous, soredia and isidia lacking. Medulla white. Lower surface black, moderately to densely rhizinate, rhizines black, densely dichotomously branched. Pycnidia scattered, conidia bifusiform 5-7 x 1 um. Apothecia common, substipitate, 2-4 mm diam., thalline exciple smooth to undulate; spores (8), colourless, ellipsoid, 8-11 x 5-7 um. Chemistry: Cortex K+ yellow; medulla K-, C+ pink, P-; containing atranorin, chloroatranorin, gyrophoric acid (major), 3-methoxy-2,4-di-O- methygyrophoric acid (trace), 2,4-di-O-methygyrophoric acid (trace), unknown (minor). Morphologically this new species resembles Hypotrachyna heterochroa (Hale & Kurok.) Elix, as both have narrow sublinear lobes which are commonly fertile and lack soredia and isidia. However whereas the thalli of H. heterochroa are adnate, have a pigmented lower medulla 108 and contain protocetraric acid and skyrin, H. proserpinensis has loosely adnate thalli with a white medulla containing gyrophoric acid. At present this new species is known only from the type collection. Hypotrachyna subpustulifera Elix sp. nov. Fig. 8 Species cum thallo ut in Hypotrachyna pustulifera sed ab hac specie thallo saxicolo, multo parviore et lobis angustioribus differt. Type: Australia. New South Wales. On exposed sandstone rocks in dry sclerophyll forest, trail to Pigeon House Mountain, 19 km west of Ulladulla, 35°20'S, 150°16'E, 460 m, J. A. Elix 21305 & H. Streimann, 2.xii.1986; holo: CBG; iso HO, MEL. Thallus corticolous, adnate to tightly adnate, whitish mineral-grey, 2.5-4.0 cm diam. Lobes linear-elongate, separate to subimbricate, dichotomous to subirregularly divided, 0.5-1.0 mm wide, margins entire. Upper surface shiny, continuous, plane to slightly convex, emaculate, pustulate; pustules subapical, bursting open, eroding and becoming coarsely granular-sorediate with age and exposing black lower cortex. Medulla white. Lower surface black, moderately to densely rhizinate below and at the margins, rhizines black, shiny, dichotomously branched. Pycnidia and apothecia not seen. Chemistry. Cortex K-, medulla K- or K+ pale reddish, C- KC+ rose, P-; containing atranorin, chloroatranorin, colensoic acid (major), 4-O- methylphysodic acid (major), lividic acid (trace), physodic acid (minor), oxyphysodic acid (trace). Superficially this new species resembles Hypotrachyna pustulifera (Hale) Hale as both species have erumpent pustules on the upper surface and contain components of the lividic acid-physodic acid chemosyndrome. However H. pustulifera is corticolous and has much larger thalli (4-6 cm diam.), broader lobes (2-3 mm wide) and contains lividic acid as the major secondary metabolite. H. subpustulifera by contrast is saxicolous, has smaller thalli (2.5-4.0 cm diam.), narrower lobes (0.5-1.0 mm wide) and contains 4-O-methylphysodic acid and colensoic acid as major secondary substances. This new species is known from several localities in New South Wales, growing on Hawksbury sandstone. Specimens Examined NEW SOUTH WALES. On sandstone rocks in heath, Peckmans Plateau, Katoomba, 980 m, J. A. Elix 3211, 24.iv.1977 (ANUC); on sandstone rocks in open Eucalyptus woodland, Morton National Park, 8 km north-east of Nerriga, 35°07'S, 150°08'E, 750 m, J. A. Elix 11598 & P. W. James, 20.i.1984 (ANUC). 109 Figures 5-8. New species of Parmeliaceae: 5, Flavoparmelia kantvilasii (isotype in ANUC); 6, Hypotrachyna booralensis (holotype in MEL); 7, Hypotrachyna_ proserpinensis (holotype in CBG); 4, Hypotrachyna subpustulifera (holotype in CBG). Scale bar (Fig. 5,7) = 10 mm; (Fig. 6,8) = 5 mm. 110 Neofuscelia kondininensis Elix sp. nov. Fig. 9 Species cum thallo ut in Neofuscelia incantata sed ab hac specie subto pallido differt. Type: Australia. Western Australia. On exposed rocks in open scrubland, with scattered Acacia and Eucalyptus, 29 km east of Kondinin on road to Hyden, J. A. Elix 21680 & M. V. Sargent, 19. viii.1987; holo-CBG. Thallus saxicolous, tightly adnate, yellowish brown to dark brown, 4-5 cm in diam. Lobes sublinear-elongate to subirregular, barely imbricate, 0.5- 1.5 mm wide. Upper surface dull or slightly shining, smooth at the margins, becoming rugulose on older lobes, isidiate; isidia cylindrical, simple for the most part, seldom branched. Lower surface ivory to pale brown, moderately rhizinate, rhizines simple, to 0.3 mm long. Pycnidia rare, conidia not seen. Apothecia not seen. Chemistry. Cortex K-, HNO3+ dark blue-green; medulla K-, C-, KC+ rose- red, P-; containing physodic acid. This new species closely resembles Neofuscelia incantata (Essl.) Essl. both morphologically and chemically, but differs in the colour of the lower surface. In N. incantata the underside is dark brown or black, but often paler at the apices whereas in N. kondininensis the lower surface is ivory to pale brown, but becoming darker at the apices. At present this new species is known only from the type collection. Neofuscelia imatatricoides Elix sp. nov. Fig. 10 Species cum thallo ut in Neofuscelia imitatrix simili sed ab hac specie lobis accessoriis saepe subcylindrico et acido 4-O-methylphysodico, acido a- collatolico et acido alectoronico continente differt. Type: Australia. New South Wales. On sandstone rocks in open Eucalyptus forest, Morton National Park, 8 km north-east of Nerriga, 35 07'S, 150 08'E, 760 m, J. A. Elix-3062, 30.iii.1977; CBG-holotype, MEL-isotype. Thallus foliose, saxicolous, moderately adnate, dark brown to black- brown, paler at the periphery, 3-4 cm diam. Lobes sublinear-elongate, 0.6-1.0 (-1.2) mm wide, flat or slightly convex at the apices, becoming more strongly convex, cracked and areolate within the thallus, developing very narrow (0.1-0.4 mm wide), elongate, dichotomously divided secondary lobes at the periphery and within the thallus. The secondary lobes subfruticose to fruticose in habit, circular to subcircular in cross section and for the most part unattached to the substrate. Upper surface dull to slightly shiny at the apices, smooth at the apices but becoming markedly rugose within the thallus, lacking soredia and isidia. Medulla white. Lower surface jet black, 111 shining, sparsely rhizinate, rhizines black, short, simple or occasionaly tufted, slender. Pycnidia common, conidia bifusiform, 6-8 x 1 um. Apothecia common, short stipitate, 1-3 mm in diam., disc black-brown, concave at first but becoming more or less flat or undulating, margin thin, entire; spores (8), colourless, ellipsoid, 7-9 x 3.5-5.5 um. Chemistry. Cortex K-, HNO+ dark blue-green, medulla K-, C-, P-; containing alectoronic acid (major), a-collatolic acid (major), 4-O-methylphysodic acid (major) and physodic acid (minor). This new species appears to be related to Neofuscelia imitatrix (Taylor) Essl. since both species often develop distinctly pulvinate thalli with a black lower surface, and give similar chemical reactions. However they can be separated clearly by their morphology since the secondary lobes of N. imitatricoides are distinctly circular to subcircular in cross section while those of N. imitatrix are flat to weakly convex. The chemical profile of these two species also differs consistently since N. imitatricoides contains alectoronic acid, a-collatolic acid and 4-O-methylphysodic acid (all in substantial proportions) and physodic acid (traces or minor) while N. imitatrix contains physodic acid (major) together with occasional traces of 4- O-methylphysodic acid. Thus these two species appear to have overlapping chemistries (i.e. they belong to the same chemosyndrome) but maintain their chemical integrity even when sympatric and are considered distinct species (Elix 1982). Neofuscelia parasitica Elix sp. nov. Fig. 11 Species cum thallo ut in Neofuscelia pulla sed ab hac specie acido glomelliferonico, acido loxodellonico et acido glomellonico continente differt. Type: Australia. Australian Capital Territory. On sandstone rocks, Kowen forest, 16 km east of Canberra, 35°15'S, 149°16'E, 730 m, J. A. Elix 1329, 21.xi.1975; CBG-holotype. Thallus small foliose, growing over thalli of Neofuscelia verisidiosa (Essl.) Essl. or Neofuscelia loxodella (Essl.) Essl., moderately adnate, olive- brown to reddish or dark brown, often paler at the periphery, to 2.0 cm diam. Lobes irregular to sublinear-elongate, 0.8-1.5 mm wide, contiguous to imbricate in part. Upper surface dull to slightly shiny at the apices, smooth, lacking soredia and isidia. Medulla white. Lower surface black, sometimes dark brown towards the apices, moderately to densely rhizinate, rhizines slender, simple, black. Pycnidia and apothecia not seen. Chemistry. Cortex K-, HNO3+ dark blue-green, medulla K-, C-, KC+ rose, P-; containing glomelliferonic acid (major), loxodellonic acid (major), and glomellonic acid (trace). iy rd This unique new species could be confused with Neofuscelia delisea (Duby) Essl. since both species lack isidia, have a black lower surface, and give comparable medullary spot tests. However, N. delisea contains glomelliferic acid, loxodellic acid and glomellic acid whereas N. parasitica contians the isostructural depsidones, glomelliferonic acid, loxodellonic acid and glomellonic acid. This chemistry has been observed previously in N. subincerta (Essl.) Essl. (an isidiate species) and N. pyrenaica (Essl.) Essl. (Elix, Jenkins & Jenie 1987). The latter species differs from N. parasitica in the much larger thalli (2-9 cm diam.), pale brown to black-brown lower surface and saxicolous substrate. At present N. parasitica has only been found growing over thalli of the common isidiate Neofuscelia species, N. verisidiosa and N. loxodella. Interestingly the former species produces medullary divaricatic acid, a depside unrelated structurally to the depsidones present in N. parasitica. At present this new species is only known from the type locality where it appears quite common. Specimens Examined AUSTRALIAN CAPITAL TERRITORY. Type locality, J. A. Elix 921, 13.vi.1975, J. A. Elix 21807A, 25.x.1987 (ANUC). Neofuscelia scabrosina Elix sp. nov. Fig. 12 Species cum thallo ut in Neofuscelia luteonotata sed ab hac specie acido constipatico, acido protoconstipatico, scabrosini 4,4'-diacetate, scabrosini 4-acetate-4'- sputyzate, scabrosini 4,4'-dibutyrate et acido ursolico continente differt. Type: Australia. Western Australia. On soil and termite mounds at base of rock outcrop, Camels Peak, N. Sammy, 24.v.1972 ; holo-PERTH 810191. Thallus foliose, terricolous, moderately adnate to the substrate, olive- brown or dark brown to black-brown, to 8 cm diam. Lobes irregular to sublinear-elongate, 1.0-3.0 mm wide, discrete to imbricate in part. Upper surface dull to slightly shiny and smooth at the apices but becoming markedly rugose within the thallus, lacking soredia and isidia. Medulla white. Lower surface pale tan to light brown, sometimes darkening towards the lobe apices, moderately to densely rhizinate, rhizines short, simple, concolorous with the lower surface or darkening. Pycnidia common, conidia bifusiform, sometimes only weakly so, 6-8 x 1 um. Apothecia common, short stipitate, 3-5 mm in diam., disc black-brown, concave at first but becoming more or less flat or undulating, margin thin, entire; spores (8), colourless, ellipsoid, 6-9 x 3.5-5.5 um. Chemistry. Cortex K-, HNO+ dark blue-green, medulla K-, C-, P-; containing constipatic acid (major), protoconstipatic acid (major), scabrosin 4,4'- 113 ‘ee 3 = B: Figures 9-12. New species of Parmeliaceae: 9, Neofuscelia kondininensis (holotype in CBG); 10, Neofuscelia imatatricoides (holotype in CBG), 11, Neofuscelia parasitica (holotype in CBG); 12, Neofuscelia scabrosina (holotype in PERTH). Scale bar = 5 mm. 114 diaceatate, scabrosin 4,4'-dibutyrate, scabrosin 4-acetate-4'-butyrate and ursolic acid. This new species could be confused with Neofuscelia luteonotata (J. Stein.) Essl. since both species often occur on soil, have similar foliose thalli with a pale tan lower surface, and give completely negative medullary spot tests. The two species have slightly different general appearance which is difficult to define and positive identification must rely on chemical analysis. Thus N. scabrosina contains the fatty acids constipatic acid and protoconstipatic acid and three scabrosin derivatives while N. luteonotata contains stenosporic acid and/or divaricatic acid, sometimes with accessory gyrophoric acid. Furthermore N. scabrosina has a very limited distribution (Western Australia) while N. luteonotata is widely distributed in southern Europe, north and south Africa as well as Australia. Given the divergent distributions and unrelated chemistry these two taxa are considered distinct species (Elix 1982). Chemically this new species most closely resembles N. scabrella (Essl.) Essl. from South Africa, but the latter species lacks the fatty acids, and is distinguished morphologically by the elongate, dichotomously branched, loosely entangled lobes. At present this new species is only known from the type locality. Specimen Examined WESTERN AUSTRALIA. Type locality, N. Sammy, 24.v.1972 (PERTH). Neofuscelia subbarbatica Elix sp. nov. Fig. 13 Species cum thallo ut in Neofuscelia parviloba sed ab hac specie thallo crassiore et acido barbatico, acido 4-O-demethylbarbatico et acido norobtusatico continente differt. Type: Australia. Western Australia. On low granite rocks in sclerophyll forest, 40 km south-east of Armadale, beside the Albany highway, G. Rambold 5276, 1.iv.1986 ; holo-M. Thallus small foliose or becoming subcrustose in the centre, tightly adnate, dark brown, 1-4 cm in diam. but coalescing into larger patches. Lobes sublinear to irregular, 0.2-0.8 mm wide. Upper surface more or less smooth at the periphery, convex within and becoming fissured and rugose to subareolate, dull throughout or slightly shiny at the apices, without isidia. Medulla white. Lower surface pale tan to pale brown, smooth and more or less dull, in part attached directly to the substrate, elsewhere sparsely rhizinate, the rhizines concolorous or darkening, small and fine, to 0.2 mm long. Pycnidia common, conidia weakly bifusiform, 5-6 x 1 um. Apothecia common, sessile or short stipitate, concave at first but becoming more or less flat or undulating, to 0.5-2.0 mm in diam., margin entire; spores (8), colourless, ellipsoid, 7-9 x 4-6 um. 115 Chemistry. Cortex K-, HNO3+ dark blue-green; medulla K-, C-, KC+ yellow, P-; containing barbatic acid (major), norobtusatic acid (minor), 4-O- demethylbarbatic acid (trace). Morphologically this new species closely resembles Neofuscelia parviloba (Essl.) Essl. in the diminutive subcrustose thalli, the pale lower surface and the absence of isidia. However N. parviloba has a flatter, thinner thallus and contains fumarprotocetraric acid and protocetraric acid. N. melanobarbatica (Essl.) Essl. the only other barbatic acid containing Neofuscelia with a pale lower surface is distinctly foliose and loosely adnate. At present this new species is only known from the type collection. Paraparmelia yamblaensis Elix sp. nov. Fig. 14 Species cum thallo ut in Paraparmelia lithophiloides sed ab hac specie thallo laxe adnato, lobi latioribus et acido norstictico et acido salazinico noncontinente differt. Type: Australia. New South Wales. On rocks in grassland, Great Yambla Ridge, 17 km south-south-east of Culcairn, 35°50'S, 147°04'E, 580 m, J. A. Elix 23095, 16.xi.1989; holo: CBG, iso: MEL. Thallus foliose, loosely adnate on rocks, pale mineral grey but darkening with age, irregularly lobate, 6-8 cm in diam. Lobes variable, irregular to sublinear-elongate, moderately to sparingly imbricate, 2.0-3.0 (- 5.0)mm wide, the apices crenulate, more or less rotund, occasionally developing narrower (0.5-1.5 mm), subdichotomously divided secondary lobes. Upper surface dull, emaculate, lacking soredia and isidia, darkening with age, the margins commonly blackened. Medulla white. Lower surface jet black except for a narrow brown marginal zone, very sparsely rhizinate, rhizines concolorous with the lower surface, simple, short, slender. Pycnidia rare; conidia bifusiform, 6-9 x 1 um. Apothecia scattered, adnate, 3- 8 mm in diam.; disc concave then undulate-distorted, brown to black- brown, margin thick and involute at first, then thin, persistent; spores (8), colourless, ellipsoid, 7-8 x 5-6 um. Chemistry: Cortex K+ yellow; medulla K-, C-, KC-,.P-; containing atranorin. This new species is characterised by the large, loosely adnate thalli, the broad sublinear lobes with a black lower surface which lack isidia, and the absence of medullary chemistry. The more adnate portions of the thallus resemble P. lithophiloides (Kurok.) Elix & Johnston, but that species has narrower lobes (1-2 mm wide), denser rhizines and contains the medullary depsidones, norstictic acid and salazinic acid. P. yamblaensis is only known from the type locality where it is particularly common. 116 Specimens Examined NEW SOUTH WALES. Type locality, J. A. Elix 23106, 16.xi1.1989 (ASU, PRE); Ju A, ElpeZ3T 22h SAGX 23123 MiG xt 989K ANDS) Parmelia crowii Elix sp. nov. Fig. 15 Species cum thallo ut in Parmelia erumpens sed ab hac specie isidiis coralloidibus et esorediatis differt. Type: Australia. New South Wales. Brigadoon Farm, Bunga, 18 km south of Bermagui, 36°34'S, 150°03'E, 30 m, J. A. Elix 23373, 2.xii.1989; holo: CBG; iso ANUC, HO. MEL, PRE. Thallus saxicolous, adnate to loosely attached, pale mineral-grey, 8-20 cm in diam. Lobes short, subirregular to apically rotund, imbricate, 2-5 mm wide. Upper surface shiny, plane, white-reticulate at first but becoming conspicuously cracked to the margin, pseudocyphellae effigurate, somewhat raised, dense, fusing into a reticulate network over the whole surface, isidiate; the isidia epicorticate, coarse, cylindrical at first then coralloid, rarely bursting apically and not becoming granular and sorediose. Lower surface moderately rhizinate, rhizines simple or squarrosely branched, 1-2 mm long. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow; medulla K+ yellow then deep red, C-, P+ red- orange; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (minor), norstictic acid (trace). This new species closely resembles P. erumpens Kurok. except in the develpment of asexual propagules. Thus P. erumpens is sorediate-isidiate where the coarse granular soredia become in part corticate, forming dense marginal and laminal soralia and/or extended, granular, coralloid-isidioid outgrowths while in P. crowii the coarse cylindrical isidia become densely coralloid branched, but only very rarely burst open and then do not become sorediate. Parmelia pseudotenuirima Gyelnik, another Australian species with cylindrical isidia, differs in having much thinner (less than 0.05 mm vs. 0.2-0.3 mm thick), syncorticate isidia. This new species is named in honour of Dr. W. D. Crow, friend, colleague and owner of Brigadoon Farm. Parmelina euplectina Kurok. ex. Elix sp. nov. Fig. 16 Species cum thallo ut in Parmelina pseudorelicina sed ab hac specie medulla inferiore pigmento ochraceo et euplectino continente differt. 117 Figures 13-16. New species of Parmeliaceae: 13, Neofuscelia subbarbatica (holotype in M); 14, Paraparmelia yamblaensis (holotype in CBG); 15, Parmelia crowii (holotype in CBG); 16, Parmelina euplectina (holotype in MEL). Scale bar = 5 mm. 118 Type: Australia. New South Wales. On Casuarina species, Raymond Terrace to Buladelah road, 30 km north of Karuah, R. Filson 7176, 9.v.1965; holo: MEL. Thallus corticolous, adnate to tightly adnate, pale grey to grey-green, 2-4 cm diam. Lobes sublinear-elongate, generally narrow but occasionally more robust, 1.0-3.0 mm wide, irregularly branched, imbricate, margins sinuous, shining, black, ciliate, the cilia sparse, black, simple, short, 0.1-0.3 mm long, mainly in the axils of the lobes. Upper surface shiny, plane or weakly foveolate at first, becoming rugulose with age, emaculate, lacking soredia and isidia. Upper medulla white, lower medulla orange-yellow. Lower surface black, with a narrow, brown, naked marginal Zone, moderately to sparsely rhizinate, rhizines simple, black. Pycnidia numerous, conidia cylindrical to weakly fusiform, 6-7 x 0.7 um. Apothecia common, sessile to substipitate, 1-2.5 mm diam., disc concave, margin thin, entire, not crenulate or inflexed, thalline exciple smooth; spores (8), colourless, ellipsoid, 9-13 x 6-7 um. Chemistry. Cortex K+ yellow, medulla K-, C+ red, P-, lower medulla K+ purple; containing atranorin, chloroatranorin, lecanoric acid (major), euplectin. Although this new species closely resembles Parmelina pseudorelicina (Jatta) Kantvilas & Elix, it is clearly distinguished by the partially orange-yellow pigmented lower medulla due to the presence of the anthraquinone, euplectin. To date this is the only species of Parmelina known with a pigmented medulla. This species was formerly referred to as Parmelia jejunga Kurok. but the latter is a nomen nudum (Filson 1983). P. euplectina is a very rare endemic species, currently known only from the type locality. Parmelina johnstoniae Elix sp. nov. Fig..17 Species cum thallo ut in Parmelina conlabrosa sed ab hac specie thallo brunneo-cinerascens, fragiliore et acido protolichesterinico et acido lichesterinico continente differt. Type: Australia. New South Wales. On trunk of Acacia, Buckenbowra River Estuary, 7.5 km W of Batemans Bay, 34°42'S, 150°06'E, 5 m, J. Johnston 2610; holo: CBG. Thallus corticolous or lignicolous, adnate, brownish-grey to pale mineral grey, 2-6 cm in diam. Lobes variable, sublinear-elongate to subirregular and apically subrotund, 1-3 mm wide, sometimes subdichotomously branched, marginal cilia moderate to sparse and concentrated in the lobe axils, simple. Upper surface shiny, emaculate, more or less plane, moderately to densely isidiate, the isidia cylindrical, 119 erect, often branched, less than 0.5 mm high. Medulla white. Lower surface black, moderately rhizinate, the rhizines black, shiny simple. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P-; containing atranorin, chloroatranorin, lichesterinic acid, protolichesterinic acid. This new species is distinguished by the cylidrical isidia and medullary chemistry. Morphologically this species closely resembles the common P. conlabrosa (Hale) Elix & Johnston but differs in the browner, more fragile thallus and the negative medullary reactions. P. conlabrosa contains medullary lecanoric acid. P. johnstoniae is probably the isidiate counterpart of P. endoleuca (Taylor) Hale. At present this new species is only known from the type locality where it appears quite common. This species is named in honour of my former colleague, Ms. Jen Johnston. Specimens Examined NEW SOUTH WALES. Type locality, J. A. Elix 10978, 29.v.1983, J. A. Elix 22618, 4.vili.1988 (ANUC). Parmelinopsis protocetrarica Elix sp. nov. Fig. 18 Species cum thallo ut in Parmelinopsis minarum sed ab hac specie subtus pallido et acido protocetrarico continente differt. Type: Australia. Queensland. On mossy trunk of felled tree in rainforest, Walter Hill Range, 26 km south-east of Ravenshoe, 17°46'S, 145°41'E, 800 m, J. A. Elix 17041 & H. Streimann, 2.vii.1984; holo: CBG. Thallus corticolous, adnate, whitish to pale greenish mineral grey, 2-4 cm diam. Lobes sublinear-elongate, contiguous, 0.8-1.2 mm wide, the marginal cilia irregularly dispersed, simple, to 0.7 mm long. Upper surface shiny, sparingly maculate at the apices, plane, densely isidiate, the isidia cylindrical, erect, often branched, to 0.5 mm high, becoming procumbent and flattened with age. Medulla white. Lower surface ivory to pale brown, black-brown at the apices, moderately rhizinate, the rhizines black, shiny, simple or squarrose branched. Pycnidia and apothecia not seen. Chemistry. Cortex K+ yellow medulla K-, C+ rose, KC+ red, P+ orange; containing atranorin, gyrophoric acid (minor), protocetraric acid (major), unknown (minor). Morphologically this new species superficially resembles Parmelinopsis minarum (Vainio) Elix & Hale with fragile thalli with densely isidiate narrow lobes. However P. protocetrarica is clearly distinguished by the pale lower surface of the lobes (black in P. minarum) 120 and the presence of additional protocetraric acid in the medulla. At present this new species is only known from the type collection. Parmotrema submerrillii Elix sp. nov. Fig. 19 Species cum thallo ut in Parmotrema merrillii sed ab hac specie lobis sine laciniis marginalibus differt. Type: Australia. Queensland. On tree in picnic area adjacent to rainforest, Mt. Spec National Park, ridge on the Loop, on the Paluma road, WNW of Townsville, 1000 m, M. E. Hale 64052, 28.vii.1983; holo: CBG. Thallus corticolous, loosely adnate, coriaceous, pale mineral grey or darkening with age, 8-10 cm diam. Lobes irregular, 5-15 mm wide, with rotund apices, margins ciliate, cilia prominent, 1.0-3.5 mm long. Upper surface emaculate, lacking soredia and isidia. Medulla white. Lower surface black, marginal rim brown, sparsely rhizinate, rhizines simple, black. Pycnidia common, conidia sublageniform, 5-6 x 1 um. Apothecia common, stipitate, 4-8 mm diam., imperforate, the exciple dentate. ciliate, amphithecium rugose, white maculate; spores (8), colourless, ellipsoid, 25- 34 x 12-16 um. Chemistry. Cortex K+ yellow, medulla K-, C-, KC-, P+orange-red; containing atranorin, chloroatranorin, protocetraric acid (major). This new species resembles the south-east Asian species, P. merrillii (Vainio) Hale both chemically and morphologically. However where P. merrillii develops long narrow laciniae on the lobes and frequently on the exciple of the apothecia, these are absent in P. submerrillii, although the exciple is dentate in the latter. This species may also be confused with P. zollingeri (Hepp) Hale as they have similar thalli and contain protocetraric acid. However P. zollingeri has very sparse, short (less than 1 mm) cilia, smaller spores (18-22 x 8-10 um) and an eciliate exciple. At present this new species is only known from the type locality. Xanthoparmelia austrocostrictans Elix sp. nov. Fig. 20 Species cum thallo ut in Xanthoparmelia constrictans sed ab hac specie superne emaculata et rhizinis modicis differt. Type: Australia. New South Wales. On sandstone rocks in open Eucalyptus woodland, Morton National Park, 8 km north-east of Nerriga, J.A. Elix 3070; CBG-holotype. Thallus foliose, moderately to loosely adnate to rocks or soil, yellow- green in colour darkening with age, 5-7 cm in diam. Lobes linear- elongate, dichotomously to subdichotomously branched, moderately to 121 Figures 17-20. New species of Parmeliaceae: 17, Parmelina johnstoniae (J. A. Elix 10978 in ANUC); 18, Parmelinopsis protocetrarica (holotype in CBG); 19, Parmotrema submerrillii (holotype in CBG); 20, Xanthoparmelia austroconstrictans (holotype in CBG). Scale bar = 5 mm 122 sparingly imbricate, 0.4-1.5 mm wide but often irregularly widened and constricted. Upper surface smooth, shiny, emaculate, lacking soredia and isidia but with numerous pycnidia, margins black; medulla white. Lower surface jet black but brown at the tips of the lobes, moderately rhizinate, rhizines simple or tufted. Pycnidia common, conidia bifusiform, 5-8 x 0.7 um. Apothecia not seen. Chemistry. Cortex K-, medulla K+ yellow becoming deep red, C-, P+ orange; containing usnic acid, salazinic acid and consalazinic acid. X. austroconstrictans is distinguished by the loosely imbricate, linear elongate lobes which are irregularly constricted and the production of salazinic acid in the medulla. Chemically X. austroconstrictans is identical with X. rubrireagens - a morphologically similar species. However the lobes of X. rubrireagens tend to be much more uniform in width (i.e. not irregularly constricted) and the underside is sparsely rhizinate and often erhizinate towards the apices, the lower surface rugose and wrinkled, and the rhizines themselves short, robust and + tufted. By contrast X. austroconstrictans is moderately rhizinate, with + subapically grouped rhizines and the lower surface is more or less plane. This new species also resembles the South African X. constrictans (Nyl.) Hale - a species which differs in having a maculate upper surface and dense, long, + branched, curved rhizines commonly projecting beyond the margins of the lobes. X. austroconstrictans is a rare species in Australia, being restricted to the mountains of south-eastern Australia (New South Wales and Victoria) and Tasmania. Specimen Examined VICTORIA. Bullock Creek, 16 km south of Bendigo along Calder Highway, J. Johnston 1162A (ANUC). TASMANIA. Royal George Mine area, G.C. Bratt 345 (HO). Xanthoparmelia canobolasensis Elix sp. nov. Fig. 21 Species cum thallo ut in Xanthoparmelia glareosa sed ab hac specie thallo sparsim rhizinato, rhizinis simplicibus et acido hypostictico et acido hyposalazinico continente differt. Type: Australia. New South Wales. On volcanic rock ledge in open Leptospemum scrub, west slope of Mt. Canobolas, 8 km south-west of Orange, 32°21'S, 148°59'E, 1300 m, J. Johnston 2909, 6.xii.1989; holo: CBG. Thallus foliose, adnate to tightly adnate, saxicolous or terricolous, yellow-green but becoming greyish with age, 5-10 cm in diam. Lobes subirregular, elongate, imbricate, 1.0-3.0 mm wide, with distinct black margins; secondary lobes 0.4-1.0 mm wide, subdichotomously branched, 123 overlying the centre of the thallus but sometimes becoming dominant. Upper surface dull or commonly shiny, emaculate, more or less smooth, lacking soredia and isidia; medulla white. Lower surface black, brown-black at the apices, sparsely rhizinate, rhizines simple, concolorous with the lower surface. Pycnidia common, conidia bifusiform 5-7 x 1 um. Apothecia sessile, 2.0-4.0 mm in diam., concave; disc cinnamon-brown, margin thin, strongly involute at first but becoming crenulate and lacerate; spores (8), colourless, ellipsoid, 7-10 x 5-6 um. Chemistry. Cortex K-; medulla K+ yellow becoming red, C-, KC-, P+ yellow- orange; containing usnic acid, norstictic acid (major), connorstictic acid (minor), hypostictic acid (minor/trace), and hyposalazinic acid (trace). This new species resembles X. glareosa (Kurok. & Filson) Elix & Johnston in growth form and chemistry, but X. glareosa is moderately rhizinate with simple and dichotomously branched rhizines and contains medullary salazinic acid in addition to norstictic and connorstictic acids whereas X. canobolasensis is sparsely rhizinate, has simple rhizines and contains hypostictic and hyposalazinic acids in addition to norstictic and connorstictic acids. Specimen Examined NEW SOUTH WALES. On soil in grassy area, Orange View, Pinnacles Road, Mt. Canobolas, 1100 m, M. E. Hale 59947, 25.11.1984 (ANUC). Xanthoparmelia hyposalazinica Elix sp. nov. Fig. 22 Species cum thallo ut in Xanthoparmelia subnuda sed ab hac specie lobis sublineariter elongatis et acidum hyposalazinicum continente differt. Type: Australia. Australian Capital Territory. On porphyry rocks in pasture, 0.8 km west of Coppins Crossing, 8 km west of Canberra, 35°17'S, 149°03'E, 520 my fA Elix Al/753,17 41992) holo: CBG: Thallus foliose, loosely adnate on rocks, yellowish-green, 3-5 cm in diam. Lobes sublinear-elongate, di- or trichotomously branched, imbricate, often ascending at the apices, 0.5-1.5 mm wide. Upper surface smooth, glossy or opaque, emaculate, often blackened at the margins, lacking soredia and isidia; medulla white. Lower surface black, nude or sparsely rhizinate, rugulose, rhizines simple, black, robust, 0.2-1.0 mm long. Pycnidia common; conidia bifusiform, 5-7 x 1 um. Apothecia sessile to substipitate, 2- 4 mm in diam., margins undulate or not, disc dark brown, thalline exciple smooth, emaculate; spores (8), colourless, ellipsoid, 9-12 x 5-7 um. Chemistry. Cortex K-, medulla K+ yellow then red, C-,KC-, P+ yellow; containing usnic acid and hyposalazinic acid. 124 Its narrow, sublinear-elongate lobes, lack of maculae, soredia and isidia together with the black lower surface and medullary hyposalazinic acid distinguish this species. X. subnuda (Kurok.) Elix, a related species, has more linear-elongate lobes and contains medullary norstictic acid and connorstictic acid. Currently this rare endemic species is only known from the type locality. Xanthoparmelia masonii Elix sp. nov. BigaZ3 Species cum thallo ut in Xanthoparmelia mougeotina sed ab hac specie thallo subtus brunneo differt. Type: Australia. New South Wales. On granite outcrops in pasture with Eucalyptus, M. E. Hale 59,276, 10.i.1982; holo: CBG. Thallus small foliose to subcrustose, tightly appressed on rocks, greenish-yellow in colour, subirregularly lobate or becoming almost subcrustose towards the centre, 3-6 cm in diam. Lobes sublinear, discrete or contiguous, 0.3-1.0 mm wide, older lobes irregularly fractured and areolae commonly formed. Upper surface opaque, emaculate, often tangentially cracked, moderately to sparsely isidiate, the isidia cylindrical, mainly simple; medulla white. Lower surface ivory to pale brown but often straw-yellow near the apices of the lobes, moderately rhizinate, the rhizines simple, pale brown. Pycnidia and apothecia not seen. Chemistry. Cortex K-, medulla K+ yellow, C-, KC-, P+ orange; containing usnic acid, stictic acid, constictic acid, norstictic acid (minor), cryptostictic acid (trace), rarely menegazziaic acid ( trace), and associated unknown (+ trace). Morphologically this species closely resembles X. mougeotina (Nyl.) D. Gall. in that both lichens have tightly adnate thalli, contiguous lobes, cylindrical isidia and medullary stictic acid. However X. mougeotina is readily distinguished by the black lower surface. Another related species, X. victoriana Elix & Johnston has a pale lower surface, but is distinguished by the inflated, erumpent isidia. At present this new species is known only from the type collection. Xanthoparmelia trirosea Elix sp. nov. Fig. 24 Species cum thallo ut in Xanthoparmelia multipartita sed ab hac specie lobi lineariter angustatis, modice imbricatis et acido hypoconstictico adjectus continente differt. 125 Figures 21-24. New species of Parmeliaceae: 21, Xanthoparmelia canobolasensis (holotype in CBG); 22, Xanthoparmelia hyposalazinica (holotype in CBG); 23, Xanthoparmelia masonii (holotype in CBG); 24, Xanthoparmelia trirosea (isotype in ANUC). Scale bar (Figs. 21, 22, 24) =5 mm; (Fig. 23) = 2 mm. 126 Type: Australia. Australian Capital Territory. On granite rocks in open woodland, Gudgenby River Gorge, 4.5 km south of Tharwa, 35°34'S, 149°04'E, 620 m, J. A. Elix 10165, 12.iv.1982; hoio: CBG, iso: ANUC, MEL JJ. A. Elix : Lichenes Australasici Exsiccati, No. 19, distributed as Parmelia multipartita (R. Br. ex. Crombie) Elix]. Thallus foliose, very loosely adnate on rocks, yellow to yellow-green but blackening with age, 6-9 cm in diam. Lobes _ linear-elongate, subdichotomously branched, divaricate in places, moderately imbricate, becoming suberect and exposing the black lower surface, 0.5-1.0 mm wide. Upper surface smooth, shiny, emaculate, lacking soredia and isidia but with black margins; medulla white. Lower suface jet black, wrinkled, with very sparse rhizines or rhizines absent, rhizines simple, short, robust. Pycnidia common; conidia bifusiform, 5-7 x 1 um. Apothecia adnate, 1.0-3.0 (-10.0) mm in diam., disc chestnut brown at first but darkening with age, margin entire, thick and involute at first but becoming thin and subentire with age; spores (8), colourless, ellipsoid, 6-10 x 3.5-5.5 um. Chemistry. Cortex K-, medulla K+, pale reddish, C-, KC-, P-; containing usnic acid, hypostictic acid, hyposalazinic acid, hypoconstictic acid . This species has previously been confused with Xanthoparmelia multipartita (Crombie) Hale, but these two taxa show consistent morphological and chemical differences. xX. multipartita differs morphologically in having more densely imbricate lobes and marginal lobes with a brown lower surface, whereas the thalli of X. trirosea are only moderately imbricate and the lower surface is jet black to the margins. Chemically X. trirosea contains hypoconstictic acid in addition to hypostictic acid and hyposalazinic acid observed in X. multipartita. The name of this species refers to the three rose-red spots which develop on a TLC plate after spraying with 10% sulfuric acid and charring. The latter species also contains minor or trace amounts of 4-O-methylhypoprotocetraric acid which is absent in X. trirosea. This new species occurs on exposed, acidic rocks in areas of moderate rainfall in coastal and hinterland areas of south- eastern Australia (N.S.W., A.C.T., Tas.). Specimens Examined NEW SOUTH WALES: On granite rocks, 10 km N of Jindabyne along the Cooma road, J. A. Elix 1653, 1654, 1818, 21.1.1976 (ANUC). TASMANIA: On granite rocks along the foreshore, Bicheno, J. A. Elix 5531, 13.1.1979 (ANUC). New Combinations and Name Bulbothrix queenslandica (Elix & Stevens) Elix, comb. nov. Basionym: Parmelia queenslandica Elix & Stevens, Aust. J. Bot. 27: 873 (1979). 127, Bulbothrix subtabacina (Elix) Elix, comb. nov. Basionym: Parmelia subtabacina Elix, in Elix & Stevens, Aust. J. Bot. 27: 875 (1979). Canoparmelia owariensis (Asah.) Elix, comb. nov. Basionym: Parmelia owariensis Asah., J. Jap. Bot. 28: 135 (1953). Canoparmelia pustulescens (Kurok.) Elix, comb. nov. Basionym: Parmelia pustulescens Kurok. in Hale & Kurok., Contrib. U.S. Nat. Herb. 36: 156 (1964). Canomaculina melanochaeta (Kurok.) Elix, comb. nov. Basionym: Parmelia melanochaeta Kurok. in Hale & Kurok., Contrib. U.S. Nat. Herb. 36: 133 (1964). Imshaugia evernica (Elix & Johnst.) Elix, comb. nov. Basionym: Parmeliopsis evernica Elix & Johnst., Mycotaxon 31: 495 (1988). Parmotrema pseudovirens (Gyelnik) Elix, comb. nov. Basionym: Parmelia pseudovirens Gyelnik, Fedde, Repert. Spec. Nov. 29: 288 (1931). Rimeliella haitiensis (Hale) Elix, comb. nov. Basionym: Parmelia haitiensis Hale, Bryologist, 62: 20 (1959). Xanthoparmelia xanthofarinosa Elix , nom. nov. Thallus adnatus vel arcte adnatus, saxicola, flavovirens, 3-5 cm diametro; lobi sublineares vel subirregulares, 0.5-2.0 mm lati. Superfices nitida, emaculata, dense sorediata, soraliis globoso-capitatis, confluentibus, et sorediis farinosis; superfices inferior castanea sed apices loborum versus saepe atrobrunnescens, modice rhizinata. Thallus acidum usnicum, acidum constipaticum, acidum_ protoconstipaticum, loxodinum, conloxodinum, norlobaridonum, conorlobaridonum et substantia ignota continens. Type: Australia. Australian Capital Territory. On sandstone rocks, Kowen Forest, 16 km east of Canberra, 730 m, J. A. Elix 1830; holo: MEL; iso: CBG. = Parmelia xanthosorediata Elix (nomen nudum, Chester & Elix 1979). 128 = Xanthoparmelia xanthosorediata (Elix) Elix & Johnston, Bull. Br. Mus. nat. Hist. (Bot.) 15: 357 (1986) (nomen nudum, Begg, Chester & Elix 1979). ACKNOWLEDGEMENTS I thank the Australian Research Council for generous financial support of this project and the late Dr. M. E. Hale for his cooperation, generous advice and assistance in obtaining critical type material in the early part of this project. I also wish to thank the staff of the Science Photographic Unit, ANU, for preparing the photographs, Mr. D. Verdon for checking the Latin descriptions and Ms. C. E. Barclay and S. A. Elix who determined the chemistry of many specimens. I thank the following colleagues and herbaria for making collections available: Dr. R. B. Filson (Booral), Ms. J. Johnston (ANUC), Dr. G. Kantvilas (HO), Dr. P. M. McCarthy (MEL), Mr. N. Sammy (PERTH), Dr. G. Rambold (M) and Mr. J. Whinray (MEL). LITERATURE CITED Begg, W. Rv Chester,’ D.’ Orie PEiix, J. A: (1979).. The structure of conorlobaridone and conloxodin. New depsidones from the lichen Xanthoparmelia xanthosorediata. Aust. J. Chem. 32: 927-929. Chester, D. O. & Elix, J. A. (1979). Three new aliphatic acids from lichens of genus Parmelia (subgenus Xanthoparmelia). Aust. J]. Chem. 32: 2565- 2569. Culberson, C. F. (1972). Improved conditions and new data for the identification of lichen products by a standardized thin-layer chromatographic method. J. Chromatogr. 72: 113-125. Culberson, C. F.. Culberson, W. L. and Johnson, A. (1981). A standardized TLC analysis of B-orcinol depsidones. Bryologist 84: 16-29. Culberson, C. F. and Johnson, A. (1982). Substitution of methyl tert.-butyl ether for diethyl ether in the standardized thin-layer chromatographic method for lichen products. J. Chromatogr. 238: 483-487. Elix, J. A. (1982). Peculiarities of the Australasian lichen flora: accessory metabolites, chemical and hybrid strains. J. Hattori Bot. Lab., 52: 407- 415. Elix, J. A., Jenie, U. A. & Jenkins, G. A. (1987). Three new depsidones from the lichen Neofuscelia subincerta. Aust. J. Chem. 40: 2031-2036. Elix, J. A. & Johnston, J. (1986). New Species of Relicina (Lichenized Ascomycotina) from Australasia. Mycotaxon, 27: 611-616. 129 Elix, J. A. & Johnston, J. (1987a). New Species of Paraparmelia (Lichenised Ascomycotina) from Australia and New Zealand. Brunonia 1986, 9: 139-193. Elix, J. A. & Johnston, J. (1987b). New species of Parmelina (Lichenised Ascomycotina) from Australia and New Zealand. Brunonia 1986, 9: 155-161. Elix, J. A. & Johnston, J. (1987c). New species and new records of Xanthoparmelia (Lichenized Ascomycotina) from Australia. Mycotaxon 29: 359-372. Elix, J. A. & Johnston, J. (1988a). New species in the lichen family Parmeliaceae (Ascomycotina) from the Southern Hemisphere. Mycotaxon 31: 491-510. Elix, J. A. & Johnston, J. (1988b). New species in the lichen genus Paraparmelia (Ascomycotina) from the Southern Hemisphere, Mycotaxon 32: 399-414. Elix, J. A. & Johnston, J. (1988c). New Species and new reports of Flavoparmelia (lichenized Ascomycotina) from the Southern Hemisphere, Mycotaxon 33: 391-400. Elix, J. A., Johnston, J. & Armstrong, P. M. (1986a). A revision of the lichen genus Xanthoparmelia in Australasia. Bull. Br. Mus. nat. Hist. (Bot.) 15: 163-362. Elix, J. A., Johnston, J. and Parker, J. L. (1987). A Catalogue of Standardized Thin Layer Chromatographic Data and Biosynthetic Relationships for Lichen Substances (Aust. Nat. University, Canberra). Elix, J. A., Johnston, J. and Parker, J. L. (1988). A Computer Program for the Rapid Identification of Lichen Substances. Mycotaxon 31: 89-99. Filson, R. B. (1983). Checklist of Australian Lichens, Ist Edn. National Herbarium of Victoria, Public Lands and Forests Division, Department of Conservation, Forests and Lands, p 70. Lumbsch, H. T. and Elix, J. A. (1985). A new species of the lichen genus Diploschistes from Australia. Pl. Syst. Evol., 150: 275-279. Nash, T. H. III & Elix, J. A. (1987). New species and new reports in the Parmeliaceae (Lichenized Ascomycotina) from South Africa and Australia. Mycotaxon, 29: 467-476. 4 J ae i iva aX i eV Se , ' i ina,” p PI Oe * i oe oe Oe ay : q rar 7 a ay ¥ MES a Wee SHE RLEN Mee Ss ee eee. ee ei bh Oy ited 2s iS COR a OTC vet aang a as | wae | ee ee ey ee SR ee ae Re ce : j eG by! me 4 2 » i YW 1s) i ALS on ot eh Ae ity avs 5 f a” 7 Meh ‘ae y dy & pata |... Mot eer? al t iy ; ay i‘, MYCOTAXON Volume XLVII, pp. 131-146 April-June 1993 TAXONOMIC STUDIES IN THE GENUS MYCOSPHAERELLA. 2. NOTES ON SOME ADDITIONAL SPECIES OCCURRING ON BRASSICACEAE. MICHAEL CORLETT Centre for Land and Biological Resources Research Agriculture Canada, Research Branch C.E.F., Ottawa, Ontario, Canada K1A OC6 Abstract Comments, descriptions and illustrations of additional non Canadian species of Mycosphaerella occurring on Brassicaceae are provided. Type specimens of four species were examined: M. denigrans, M. pachyasca, M. pashkiensis and M. pyrenaica. Several species are only known from their original descriptions as type or other specimens were unavailable for study or not located; some species have never been illustrated. Introduction A publication (Corlett, 1988) on the species of Mycosphaerella occurring on Cultivated, wild and weedy members of the family Brassicaceae in Canada treated four species, M. brassicicola (Duby) Lindau, M. cruciferarum (Fr.) Lindau, M. densa (E. Rostrup) Lind and M. tassiana (de Not.) Johans. There are some additional species described from Brassicaceae by various authors which were not dealt with in that publication. | have not seen Canadian collections of these species and they are presently presumed to be extralimital. some of the names are known to me only from the literature. Type collections of four species have been examined. | have attempted to borrow types of several others for examination but they were 132 unavailable. Nevertheless, it was considered worthwhile to discuss or provide notes or brief descriptions, from personal observations or from the literature, and where possible to provide illustrations. Unfortunately for several species known to me only from the literature and for which types or other collections have not been examined, there are no published illustrations available. Species described on Brassicaceae Mycosphaerella alyssi (Hollés) Moesz, Balkan-Kutat. Tud. Eredm. 3: 132 & 136. 1926. Sphaerella alyssi Hollés, Ann. Hist.-Nat Mus. Hung. 5: 44. 1907. This species was described by Hollds (op. cit., 1907, p. 44) on Alyssum tortuosum Waldst. & Kit. from Hungary. Hollés provided the following description without illustrations: “Peritheciis gregariis, epidermide velatis, depresso-globosis, fuscis, punctiformibus, pertusis, 50-70 p diam., contextu parenchymatico; ascis ovoideis vel Clavatis, breve st[i]pitatis, octosporis, 24-30 x 9-10 yw aparaphysatis; sporidiis 2-3 stichis obiongis, medio 1-septatis, non constrictis, hyalinis, 9-10 x 3-4 yw". Moesz (op. cit., 1926, p. 136), in making the combination to Mycosphaerella listed Alyssum scardicum Wettst. as a host from Albania. | only know this species from the literature and have not seen type or any other collections, nor have | seen any illustrations of this fungus. M. alyssi was described in Tomilin (1979, p. 75). He cited the measurements given by Hollés and did not provide any discussion or illustrations of the asci and ascospores. Mycosphaerella brassicicola (Duby) Lindau in Engler & Prantl, Die Natutrlichen Pflanzenf. 1(1): 424. 1897. Sphaeria brassicicola Duby in de Candolle, Botanicon Gallicum 2: 712. 1830, as brassicaecola, non Sphaeria brassicicola Berkeley & Broome in Berkeley, Outlines Brit. Fung., p. 401. 1860. _ Sphaerella brassicicola (Duby) Cesati & de Notaris, Comment. Soc. Crittog. Ital. 1(3): 238. 1963. Anamorph: Asteromella brassicae (F. Chevallier) Boerema & van Kesteren, Persoonia 3: 18. 1964. 133 This important pathogen causing conspicuous ‘ring spot" lesions on the leaves of Brassica species has been redescribed, illustrated and discussed by Punithalingam and Holliday (1975) and by Corlett (1988, p. 63). Mycosphaerella carniolica (Niessl) Lindau, Hilfsb. Sammeln Ascomyceten, p. 38. 1903. Sphaerella carniolica Niessl, Oesterr. Bot. Z. 25(8): 85. 1875; Hedwigia 14(10): 147. 1875. Sphaerella carniolica originally was described by Niessl (op. cit., 1875) from Draba ciliata Scop. without illustrations as follows: "Epi- rarius hypophylla. Perithecia dense disseminata, minuta, punctiformia, globosa, papillata, tandem vertice umbilicata, atra; ascis fasciculatis 8sporis, obovatis vel oblongis, sessilibus 30-42 x 14-18; sporidiis farctis, Ccuneato-oblongis, rectis, medio septatis vix constrictis, dilute virescentibus 15-19 x 4-5". This name is known to me only from the literature; | have not seen the type or any other collections. Tomilin (1979, p. 76) accepted carniolica as a distinct species of Mycosphaerella but provided no illustrations. Lind (1934, p. 66) considered Sphaerella carniolica to be a synonym of Mycosphaerella confinis (Karsten) Dearness (as confinis (Karsten) Lind). Von Arx (1949, p. 40) considered both confinis and carniolica to be synonyms of Mycosphaerella tassiana (de Notaris) Johanson. Niessl’s measurements for the asci and ascospores of S. carniolica are rather low for a mature specimen of M. tassiana. | am unable to comment on the taxonomic status of this species. Mycosphaerella confinis (P.A. Karsten) Dearness, Rep. Can. Arctic Exped. 1913-1918 4(C): 6. 1923. Sphaerella confinis P.A. Karsten, Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 1872(2): 106. 1872; Hedwigia 11: 187. 1872. This species was described by Karsten (op. cit., 1872) from dead foliage of Braya purpurascens Bunge, Draba martinsiana Gay and D. wahlenbergii Hartm. from Svalbard, Norway without illustrations as follows: “Perithecia dense gregaria, amphigena, primitus epidermidi innata, dein protuberantia, spohaeroidea, interdum vertice obtusissime 134 conoidea, dein ostiolo rotundato hiantia, laevia, glabra, nigra, latit. circiter 150 mmm. Asci subsessiles, fusoideo-elongati vel elongato- clavati, vulgo inaequilaterales, longit. 50-65 mmm., crassit. 11-13 mmm. Sporae 8:nae, di- vel subtristichae, aciculari- vel fusoideo- elongatae, ut plurimum leviter Curvulae, uniseptatae, medio non constrictae, hyalinae, longit. 16-24 mmm., crassit. 3-5 mmm. Paraphyses nullae". Measurements given by Saccardo (Syll. F. 1: 507-8. 1882) for S. confinis Karsten differ somewhat from those given by Karsten and are as follows: perithecia 100-120um, asci 45-52 x 10-13um, ascospores 14-19 x 3-5um. Dearness (op. cit., 1923, p. 6) listed a collection on Anemone parviflora Michx. [Ranunculaceae]. A collection identified as M. confinis in the Dearness herbarium (no. 6018) in DAOM, is on Antennaria margaritacea R. Br. [Asteraceae]. Number 6018 does not agree with Karsten’s description of S. confinis; the asci of the Dearness specimen are cylindrical and narrow and measure 45-50 x 6-9um and the ascospores at 14 to 15um in length likewise do not match Karsten’s description. Von Arx (1949, p. 40) considered Sphaerella confinis Karsten to be a synonym of Mycosphaerella tassiana (de Notaris) Johanson. Mycosphaerella cruciferarum (Fries) Lindau in Engler & Prantl, Die Naturlichen Pflanzenf. 1(1): 324. June 1897. Sphaeria cruciferarum Fries, Systema Mycol. 2: 525. 1823. Sphaerella cruciferarum (Fries) Saccardo, Michelia 2: 315. 1881. A presumably authentic collection of Sphaeria cruciferarum Fr. from the Fries herbarium at UPS on Cheiranthus L. from France was seen by me. A microscope slide prepared from this collection possessed a single fascicle of relatively mature aparaphysate asci, 60-65 x 14-16um, unfortunately without ascospores. While these asci have the morphology and dimensions of asci of M. cruciferarum (Corlett, 1988), a positive identification cannot be made. WM. cruciferarum was recently redescribed and illustrated by Corlett (1988, p. 64). Despite not having observed type or good authentic material, this species appears to be well characterized (Barr, 1959, p. 15; Corlett, op. cit.) by its relatively broad fusoid ascospores and the presence of a network of dark subepidermal hyphae connecting individual closely grouped ascomata. 135 Mycosphaerella denigrans (Kirschstein) Tomilin, Opredelitel’gribov roda Mycosphaerella Johns., p. 77. 1979. Sphaerella denigrans Kirschstein, Ann. Mycol. 37: 103. 1939. Mycosphaerella phlomidicola Tomilin, Nov. *Sist. Niz. Rast. 1966, p. 150. 1966. FIG. 1 Sphaerella denigrans was described by Kirschstein (op. cit., 1939, p. 103) from Erysimum hieraciifolium [given without authority] without illustrations and described as follows: “Peritheciis dense _ stipatis, substratum denigratum saepe longe, interdum ex toto inducentibus, sub peridermio convexo-sublato orientibus, demum plus minus erumpentibus, ostiolo plano, pertuso, rotundulo instructis, nigris, globosis, membranaceis, 80-100 yp diam.; contextu fuliginoso, e cellulis minutis Composito. Ascis aparaphysatis, late cylindraceis vel clavatis, apice plane rotundatis et incrassatis, vix pedicellatis, 8- sporis, 25-30 x 7-9 wu. Sporidiis oblique monostichis vel longitudinaliter di- vel tristichis, hyalinis, medio uniseptatis, oblonge ellipsoideis, rectis et [subinaequalibus], non constrictis 10-11 x 3 p". Tomilin (op. cit., 1979, p. 77 & fig. 39) has examined the holotype collection of Sphaerella denigrans, ex herb B, and has redescribed FIGURE 1, asci and ascospores of Mycosphaerella denigrans ex type: left side, redrawn from Tomilin (1979, fig 39). (scale = 10um) 136 and illustrated this species. The dimensions given by Tomilin for M. denigrans are as follows: ascomata 80-100um diam.; asci 30-34 x 10-11pm; ascospores 10-12 x 2.5-3um. | likewise have examined the holotype of Sphaerella denigrans and found asci and ascospores which agree with Kirschstein’s description and match the dimensions and_ illustrations provided by Tomilin for this species. My measurements for the asci and ascospores were 30-37 x 9-11um and 12-14 x 3-3.5um_ respectively. The ascomata were subepidermal, medium brown in colour and 99-110”um wide. The beak was 30-40jim across and the ostiole ca 10um. M. denigrans is very similar to the earlier published M. pyrenaica (Speg.) Arx. The dimensions and morphology of the ascospores of the two species are very close. Mycosphaerella densa (E. Rostrup) Lind, Rep. Sci. Results Norw. Exped. Novaya Zemlya 1921, 19: 12. 1924. S. densa E. Rostrup, Bot. Tidsskr. 14(3): 225. 1885. This species was redescribed and illustrated by Corlett (1988, p. 65). The description provided was based in part on observations of the type collection, ex herb. C. Mycosphaerella hambergii (Romell & Saccardo) Petrak, Acta Horti Gothob. 17: 137. 1947. Sphaerella confinis P.A. Karsten, Ofvers Forh. Kongl. Svenska Vetensk.-Akad. 1872(2): 106. 1872 subspecies hambergii L. Romell & Saccardo in Saccardo, Syll. Fung. 11: 296. 1895 Romell and Saccardo (op. cit., 1895, p. 296) described Sphaereila confinis var. hambergii on Arabis gerardi Bess. from Sweden without illustrations as follows: "Peritheciis dense gregariis, subsphaeroideis, 100 yp diam., epidermide arcte velatis erumpentibusque, nigris, subastomis; ascis subclavatis sessilibus, 25-50 x 7-10; sporidiis oblongo-fusoideis, obscure uniseptatis, plerumque 4-guttatis, rectis curvulisve, 12-15 x3-4, ex hyalino chlorinis". The dimensions given for subspecies hambergii are slightly less than those given by Karsten (op. cit., 1872) for Sphaerella confinis Karsten (= M. tassiana (de Notaris) Johans.). Tomilin (1979) did not discuss M. hambergii. | have not seen any material of this species and am unable to comment on its taxonomic status. 137 Mycosphaerella isatidis Kalymbetov, Trudy Inst. Bot.7: 326. 1959. M. isatidis was described by Kalymbetov (op. cit., 1959, p. 326) on /satis tinctoria L. from Kazakhstan. Kalymbetov provided the following description without illustrations: “Perithecia globosa, nigra, primo, immersa, dein emergentia. Asci cylindrici, basi fasciculati, hyalini, aparaphysati, 12,3-21,5 x 34,4-51,6 yu. Sporae hyalinae, bicellulares, cylindricae, 8,6-12,3 x 4,3-6,4 yu". Tomilin (1979, p. 291) provided a brief description of Kalymbetov’s species without illustrations: ascomata 70-100um; asci 34-52 x 12-21~7m; ascospores 10-13 x 4-5(6)um; host /satis tinctoria L. This species is known to me only from the literature. | have not seen type or any other material of M. isatidis and am unable to comment on its taxonomic status. Sphaerella napicola Fautrey, Rev. Mycol. 12(46):64. April 1890. S. napicola was described by Fautrey (op. cit, 1890, p. 64) on dead stems of Brassica napus L. [var. oleifera?]. Fautrey provided the following description without illustrations: “Taches grandes indéterminées noires; Péritheces nombreux, pressés sur la tache et disséminés au dehors, arrondis, noirs; Theques ventrues; Spores hyalines, uniseptées, a 4 gouttes, a loge supérieure plus longue et plus large, 18, 28 x 4, 6 [18.28 x 4.6]". Tomilin (1979, p. 77) listed S. napicola as a synonym of Mycosphaerella cruciferarum (Fries) Lindau. Certainly, the ascospore measurements given by Fautrey are within the range of those of M. cruciferarum. | have not seen the type or any other collections of this species. Mycosphaerella pachyasca (E. Rostrup) Vestergren, Bih. Kong. Svenska Veten.-Akad. Handl. 26(3, 12): 7. 1900. Sphaerella pachyasca E. Rostrup, Meddl. Groenland 3(2): 552. 1888. FIG. 2 Rostrup (op. cit., p. 552-3) stated that Sphaerella pachyasca Rostrup was common and widespread, occurring on dicotyledonous 138 members of numerous plant families in Greenland, including Draba and Arabis, members of the Brassicaceae. He considered it to be the morphological equivalent of Sphaerella tassiana de Notaris, the latter being restricted to monocot hosts by Rostrup. Further, Rostrup stated that these two species or species complexes were probably converging and in many cases could only be distinguished on the basis of whether the host was a dicot or a monocot [English translation of Rostrup’s discussion provided by Dr. H. Knudsen, Keeper of Mycology at the Botanical Museum (C)]. Rostrup described S. pachyasca without illustrations as follows: “Perithecia Sparsa v. gregaria, foliicola v. cladogena. Asci crasse ovoideo- oblongati, inaequilaterales, long. 40-50 p/, crass. 15-24 py, vulgo apice tunicati. Sporae conglobatae, conoideo- v. ovoideo-oblongatae, long. 16-20 py, crass. 5-6 yp. Von Arx (1949, p. 40) considered this species to be a synonym of Mycosphaerella tassiana (de Notaris) Johanson. | have FIGURE 2, asci and ascospores of Mycosphaerella pachyasca from one of several collections labelled "Typus", Disco |., 20-7-84[1884]. (scale = 10um) 139 examined collections of S. pachyasca cited by Rostrup (op. cit., p. 552) ex Herb. C and all labelled as “TYPUS". | found asci and ascospores representative of S. pachyasca in only a few of these; the asci and ascospores measured 56-60 x 17-17.5um and 15-20 x 5-6.2um respectively. These collections are morphologically indistinguishable from material | consider to be representative of M. tassiana (Corlett, 1988). | agree with the use of M. tassiana sensu von Arx as a single broad unspecialized species or species complex perhaps, widespread on many monocot and dicot plant families. If M. pachyasca were to be maintained as a distinct species, a lectotype would have to be selected from among the collections listed by Rostrup with his description of S. pachyasca. Collections examined and yielding ascospores: On Plantago borealis, Itivnek, W. & H., 13-7-84. On Diapensia lapponica, Disco |., W. & H., 20-7-84. On Draba hirta, Upernivik, 8-84. Mycosphaerella pashkiensis Petrak, Sydowia 16(1-6): 340. 1962 [May 1963]. FIG. 3 M. pashkiensis was described by Petrak (op. cit., p. 340) on dead leaves of Draba species from Afghanistan. Petrak provided the following description without illustrations: "Perithecia amphigena sed plerumque hypophyila, per totam folii sSuperficiem irregulariter dispersa, solitaria, raro bina complurave subaggregata, subepidermalia, depresso-globosa vei late ellipsoidea, 70-100 p diam., raro etiam paulo majora, omnino clausa, nec ostiolata; pariete membranaceo, ca. 10 pt crasso, contextu pseudoparenchymatico e cellulis irregulariter angulosis vel fere globosis, vix Ccompressis, obscure atro-brunneis, 3,5-6 pj diam. metientibus Composito; asci sat numerosi, rosulati, cylindracei, antice late rotundati, postice plus minusve sed plerumque parvum saccati, subsessiles vel brevissime et crassiuscule noduloso-stipitati, 8-spori, 42-55 x 9-10,5 ul; sporae di-vel indistincte tristichae, clavato-cylindraceae vel bacillari-clavatae, utrinque obtusae, antice vix vel parum., postice plerumque distincte et paulatim attenuatae, rectae, raro inaequilaterae, circa medium septate nec constrictae, guttulis plerumque duabus in quaque cellula praeditae, hyalinae, 10-13 y, raro usque ad 14,5 py longae, 2,5-3,5 pi latae; paraphysoides paucae, indistincte fibrosae, mox omnino mucosae". Mycosphaerella pashkiensis Petrak was accepted by Tomilin (1979, p. 76) who published a brief description without 140 illustrations: ascomata 70-100um; asci 42-55 x 9-10.541m; ascospores (10)12-14 x 2.5-3(3.5)um; host Draba. | have examined the type collection of M. pashkiensis ex herb. W. My dimensions of the asci (specifically ascus width) and ascospores from the type collection exceed the measurements given by Petrak for M. pashkiensis. My description of the type material is as follows: ascomata subepidermal, medium brown, 78-108um diam., ostiole, pale brown, ca 10um diam. Asci bitunicate, thick-walled, saccate when young, clavate to somewhat cylindrical, widest above or below, 8-spored, 42.5-51 x 12.5-14um [Petrak: 42-55 x 9-10.5]. Ascospores hyaline, mostly cylindrical-clavate, straight or curved, 15- 18.5 x 4-5.25um [Petrak: 10-13(14.5) x 2.5-3.5], septate at or near the middle, not or hardly constricted at the septum, upper cell wider and ascospores distinctly tapering to the base, ends rounded but not broadly so, bi- to triseriate in the ascus. | believe that both Petrak and | observed the same fungus but | cannot account for the discrepancy in our dimensions of the asci and ascospores. My measurements approach those of a smail or immature Mycosphaerella tassiana. FIGURE 3, ascoma, asci and ascospores of Mycosphaerella pashkiensis ex type. (scales = 100um & 10um respectively) 141 Mycosphaerella pyrenaica (Speg.) Arx, Sydowia 3:64. 1959. Sphaerella pyrenaica Speg., Rev. Mycol. 4(14): 78. 1882. FIG. 4 Spegazzini (op. cit. 1882, p. 78) described S. pyrenaica from dead tissue of Peterocallis [Draba]] pyrenaica from France. He provided the following description without illustrations: "Perithecia amphigena, globoso-lenticularia (90-100), innato-erumpentia, atra, submembranacea, contextu§ parenchymatico, fuligineo; asci cylindraceo-clavati, breve stipitati (40-45 x 8), apice crassiuscule tunicati, octospori, aparaphysati; sporidia disticha, cylindraceo- elliptica, utrinque obtusiuscule rotundata (15 x 3), medio 1- pseudoseptata, non constricta, hyalina". The dimensions given by von Arx (op. cit., p. 64) generally agreed with those of Spegazzini: “Pseudothecien ... 90-130 pu... Asci ... 40-50 x 9-11 yw... Sporen ...14- 19 x 3-4 yp ..". M. pyrenaica was accepted by Tomilin (1979, p. 77) who listed Cardamine bellidifolia L., Cochlearia officinalis L., Parrya nudicaulis (L.) Regel and Petrocallis [Draba] pyrenaica as hosts. | have examined the type collection of S. pyrenaica Speg., ex herb. LP and concur with the description and dimensions provided by Spegazzini and by von Arm. My observations of type material are as_ follows: ascomata subepidermal, 85- 100um diam, medium brown; asci__cylindrical-clavate, 8- spored, 40-48 x 9.5-12um; ascospores hyaline, cylindrical- ellipsoidal, straight FIGURE 4, ascus and ascospores of Mycosphaerella pyrenaica ex type (scale = 10um) or curved, 14.5-18.5 x 3.5-4.5um, septate at or just below the middle, not or hardly constricted at the septum, upper cell sometimes slightly wider than lower cell, ends rounded, bi- to triseriate in the asSCuUS. 142 Mycosphaerella tassiana (de Notaris) Johanson, Ofvers. Férh. Kongl. Svenska Vetensk.-Akad. 41(9): 167. 1884. Sphaerella tassiana de Notaris, Sferiacei Italici 1(1): 87. 1863. Anamorph: Cladosporium herbarum (Pers.) Link: S.F. Gray, Nat. Arr. Brit. Pl. 1:556. 1821. A very common and distinctive species found on numerous host plants in different monocot and dicot families. Lind (1934, p. 60) listed 110 species, a few non-angiosperms but mostly monocots and dicots, as hosts of M. tassiana. This species has been described and illustrated by von Arx (1949, pp. 40-59), M.E. Barr (1959, p. 23- 27; 1972, p. 603), Sivanesan (1984, p. 225) and Corlett (1988, p. 67-72). An extensive list of tassiana synonyms, including M. allicina (Fr.: Fr.) Vestergren and M. pachyasca (E. Rostrup) Vestergren (see above) and a lengthy host list were provided by von Arx (op. Cit.) while de Vries (1952) listed the synonyms of the anamorph. Tomilin (1979, p. 270) also considered tassiana and allicina to be synonyms but unlike von Arx, listed M. tassiana as a synonym of the earlier published M. allicina (basionym: Sphaeria allicina Fr.: Fr., Kongl. Vet.- Akad. Handl. 38:247. 1817). M.E. Barr (1972, p. 600) maintained the morphologically similar M. allicina as distinct, considering it to be specialized and host restricted, parasitic on species of Allium in contrast to the wide unspecialized host range of M. tassiana. Mycosphaereila _vesicariae-arcticae (Hennings) Tomilin, Opredelitle’gribov roda Mycosphaerella Johans., p. 78. 1979. Sphaerella vesicariae-arcticae P. Hennings in Allescher & Hennings, Biblioth. Bot. 42: 45. 1897. This species was described by Hennings (op. cit. 1897, p. 45) on leaves and stems of Vesicaria arctica R. Br. [this authority differs from those of the various homonyms of Vesicaria arctica given in Index Kewensis] var. /ejocarpa Trautw. from Denmark. Hennings provided the following description without illustrations: “Peritheciis Caulicolis et amphigenis .sparsis gregariisve, subglobosis atris; ascis oblongis subclavatis vel subfusoideis 60-75 x 17-21 tu, apice obtusis, 8-sporis; sporidiis subdistichis vel conglobatis, clavatis utrinque obtusis, medio 1 septatis haud constrictis, rectis, hyalinis 20-25 x 7- 8 yu". Tomilin (op. cit., 1979, p.78) transferred Hennings’s species to Mycosphaerella and provided a description without illustrations: 143 ascomata 90-120um; asci 60-65 x 20-24y1m; ascospores 17-19 x (5.5)6-7ym. Tomilin listed Vesicaria and Parrya nudicaulis (L.) Regel. as hosts. | have not seen the type or any other specimens of vesicariae-arcticae although the descriptions provided by both Hennings and Tomilin suggest a small spored Mycosphaerella tassiana. 144 Dimensions of ascomata, asci and ascospores of species of Mycosphaerella on Brassicaceae taken from original authors or from Corlett© (1988) Name Ascomata diameter ASsci Ascospores alysii 50-70 24-30x9-10 9-10x3-4 brassicicola® 90-145 30-47(55)x12-16 16-25x3-4 carniolica 30-42x14-18 15-19x4-5 confinis” 150 50-65x11-13 16-24x3-5 cruciferarum® 84-125 34-55x9-16 13-15x3.5-5 denigrans 80-100 30-37x9-11 12-14x3-3.5 densa® 85-140 36-54x9-11 14-24x3-4.5 hambergii 100 25-50x7-10 12-15x3-4 isatidis 34-51x12-21 8.5-12x4-6 napicola”™” 18.28x4.6 pachyasca" 40-60x15-24 16-20x5-6.5 pashkiensis© 78-108 42-51x(9)12-14 [10-13x2-3.5] 15-18x4-5 pyrenaica 85-100 40-48(50)x9-12 14-19x3-4.5 tassiana® 75-155 45-88x15-29(32) 17-31x4.5-9.5 vesicariae 90-120 60-75x17-24 17-25x6-8 -arcticae " = tassiana ™“ = cruciferarum 145 Host Index Species of Mycosphaerella on genera of Brassicaceae Alyssum alyssi™ Arabis hambergii tassiana Brassica brassicicola cruciferarum napicola (= cruciferarum) Braya confinis™ (= tassiana) densa tassiana Cardamine densa pachyasca (= tassiana) pyrenaica tassiana Cheiranthus cruciferarum Cochlearia densa pyrenaica tassiana Draba carniolica confinis (= tassiana) cruciferarum densa pachyasca (= tassiana) pashkiensis™ tassiana Erysimum cruciferarum denigrans™ Eutrema cruciferarum tassiana isatis isatidis™ Lesquerella tassiana Parrya pyrenaica tassiana vesicariae-arcticae Petrocallis (=Draba) pyrenaica™ Vesicaria vesicariae-arcticae™ ™ = Type Host 146 Acknowledgements We would like to thank Dr. M.E. Barr Bigelow, Sidney, British Columbia and Dr. S.J. Hughes, C.L.B.R.R., Ottawa, for critically reviewing the manuscript. | am grateful to the directors and curators of the following herbaria for the loan of specimens in their keeping: B, C, LP, UPS & W. | wish to thank Dr. H. Knudsen, Keeper of Mycology at the Botanical Museum (C) for kindly providing an English translation of Rostrup’s discussion following the description of Sphaerella pachyasca Rostrup. References Ar, J.A. von. 1949. Beitrage zur Kenntnis der Gattung Mycosphaerella. Sydowia 3: 28-100. Barr, M.E. 1959. Northern Pyrenomycetes I. Canadian Eastern Arctic. Contr. Inst. Bot. Univ. Montréal 73:1-101. Barr, M.E. 1972. Preliminary studies on the Dothideales in temperate North America. Contr. Univ. Mich. Herb. 9(8): 523-638. Corlett, M. 1988. Taxonomic studies in the genus Mycosphaerella. Some species of Mycosphaerella on Brassicaceae in Canada. Mycotaxon 31(1): 59-78. De Vries, G.A. 1952 Contribution to the knowledge of the genus Cladosporium Link ex Fr., 121 pp., Diss. Univ. Utrecht, Baarn. Lind, J. 1934. Studies on the geographical distribution of arctic circumpolar micromycetes. Kongel Danske Videnskabernes Selskab. Biol. Meddel. XI, 2 pp. 54-72. Punithalingam, E. & P. Holliday. 1975. CMI Descript. Path. Fungi & Bacteria No. 468. Mycosphaerella brassicicola. CAB Intern. Mycol. Inst., Kew, Surrey. Sivanesan, A. 1984. The bitunicate Ascomycetes and their anamorphs. J. Cramer, Vaduz, pp. 1-701. Tomilin, B.A. 1979. Opredelitel’gribov roda Mycosphaerella Jonans., Nauka, Leningrad, pp. 319. MY COTAXON Volume XLVI, pp. 147-155 April-June 1993 DEVELOPMENT OF CONIDIOMATA IN THE PHYLLOSTICTA STATE OF GUIGNARDIA MANGIFERAE ROY AND OBSERVATIONS ON THE FINE STRUCTURE OF THE CONIDIUM by J. Muthumary, J.A. Jayachandra and M. Bhagavathy Preetha Centre for Advanced Studies in Botany University of Madras, Madras-600 025, INDIA SUMMARY The development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy on the host and in culture is described. Fine structural details of the sheath around the conidium is provided. INTRODUCTION The developmental stages of the conidiomata in Coelomycetes have only been studied in detail in a few members of this group (Punithalingam 1966;1981;1982; Maas et al., 1979; Muthumary and Vanaja 1986). In this investigation developmental stages of the conidiomata in the Phyllosticta state of Guignardia mangiferae Roy were investigated. This forms part of a programme of work on developmental morphology and taxonomy of coelomycetes which is in progress in this department. An attempt was also made to provide information regarding the fine structure of the conidium. MATERIALS AND METHODS The material used in this study was collected in Vandaloor, near Madras, growing on leaves of Mangifera indica in September, 1991. The fungus was isolated in pure culture on oatmeal agar. The specimen was processed for TEM studies with a view to obtaining information on the ultrastructural details of the conidium and its appendage in relation to the wall layers. Portions of agar with conidiomata were fixed in primary fixative ie. 5.0% Glutaraldehyde (W/V) in 0.1 M phosphate buffer (pH=7.5) for 30 min. at room temperature and for 1 hr,. at 4°C. Similarly pieces of leaf with conidiomata were also processed in the same manner. Three washes were given in the buffer solution at 4°C to remove traces of the GA, before fixing in 1.0% O,O, for 1 hr at 4°C. After washing several times in phosphate buffer the specimen was dehydrated through a graded series of Acetone. Subsequently the specimen was embedded in Araldite (Araldite My 148 753=10ml, DDSA=10 ml, BDMA = 0.4 ml, N.Dibutylphthalate =0.5m1) and polymerized at 60°C for 36 hrs. Gold colour sections were cut with an ultracut ultramicrotome. Semithin sections of about 0.1 » thickness were selected to follow the different stages of development of the conidiomata. The semithin sections were stained with aqueous Toluidine blue for light microscopic observations. Ultrathin sections were collected on copper grids and stained with Uranyl acetate followed by lead citrate (Reynolds, 1963). Sections were observed with a CM 10 Philips Electron Microscope at 40 Kv. RESULTS Formation of conidiomatal primordium. The earliest stage of pycnidial development seen in sectioned material from the culture consists of a small, pseudoparenchymatous mass of tissue, the pycnidial primordium (Fig. 1). The pycnidial primordium is almost spherical, consisting of a compact cluster of brown, thick -walled cells. During further development, the primordium considerably increases in size by continued transverse and longitudinal divisions of the cells. Formation of the pycnidial cavity. As a prerequisite for the formation of the pycnidial cavity, the cells at the periphery and centre of the primordium were arranged as follows. The cells along the outer layers of the primordium became arranged tangentially which subsequently formed the conidiomatal wall (Figs. 2-4). The inner cells of the primordium were thin-walled and hyaline whereas the peripheral cells were brown and thick - walled. There may be more than one method involved in the formation of the central cavity. The presence of ruptured and disorganised cells in the cavity indicates that the cavity might have formed by the rupture of cells at the centre of the primordium (Figs. 5-6). The occurrence of hyphal fragments and a gelatinous material within the Cavity suggest that the cells are later dissolved to form the cavity (Fig.3). From the many sections examined, both these conditions were observed as being present in the fungus. The tangentially arranged outer layers of cells form the pycnidial wall by repeated division, thereby increasing the dimension of the developing conidioma. Simultaneously, the outer wall layers become carbonaceous and at maturity the wall consists of 5-6 tangentically arranged layers, the cells of the inner layers being hyaline and thin-walled (Figs. 10-11). Formation of sporogenous tissue. In conidiomata formed in culture, the formation of conidia from the conidiogenous cells resembles that of Ascochyta spp.. (Punithalingam, 1979) and Macrophomina phaseolina (Tassi) Goid. (Punithalingam, 1982). After the cavity is formed by dissolution and disintegration of central cells of the primordium, the cells lining the cavity commence conidial production by a budding like process. Eventually these conidiogenous cells, along with conidia become detached from the wall of the conidioma (Figs. 5- 8), and remain floating still intact in the conidiomatal cavity (Figs 7-8). The conidiogenous cells which were produced initially inside the cavity are referred to as "temporary conidiogenous cells" by Punithalingam (1979). These temporary conidiogenous cells very occasionally bear an annellation and were normally blastic. Subsequently the cells lining the cavity elongate to form a palisade of permanent conidiogenous cells (Punithalingam, 1979) which then start producing conidia (Fig. 9). 149 The presence of permanent conidiogenous cells were observed even when the cavity filled up with dissolved and disintegrated primordial cells (Fig. 9). Consequently the conidioma becomes considerably larger in size by proliferation of the outer layers of the pseudoparenchyma while the inner cells become arranged tangentially into several layers (Figs. 10-12). Formation of ostiole. When the conidioma becomes fully mature, a papilla-like projection appears at the apex (Figs. 16-18). In vertical section, the papilla shows thick-walled, dark cells towards the outer side and thin walled, hyaline, elongated cells towards the inner side (Figs. 13-15). During dehiscence of the conidioma, these inner hyaline layers in the papilla dissolve to form the ostiole and the outer, thick walled cells of the papilla remain intact and form the papillate structure of the conidioma. The papilla may also exert pressure to rupture the epidermis of the host for releasing the conidia outside. Fine structure of the conidia. Light microscopic studies by Punithalingam (1982) showed that the conidium is enveloped by a thick sheath which remains intact, its outer most layer gradually gelatinizing with the ageing of the conidium, and our studies using TEM confirmed the presence of this thick sheath around young conidia. Our observations showed the presence of the sheath around the conidiogenous cells and the growing conidium (Fig. 20). Transverse sections of a young conidium also revealed the presence of the sheath (Figs, 22-23 & 25), but in TEM micrographs of old conidia the sheath around the conidia was absent (Figs. 19, 21 & 24). It is therefore probable that during the later stage of development, the sheath around the conidium gradually gelatinizes. In observations with the light microscope, conidia within young pycnidia were found with the sheath surrounding them (Fig. 14). The mature conidium is multinucleate (Punithalingam and Woodhams, 1982) and in this study, at least three nuclei were observed (Fig. 21). TEM studies showed that the sheath surrounding the conidium disappears with ageing (Figs. 19, 21). DISCUSSION The orgin and structure of the apical conidial appendage of Phyllosticta spp. had already been fully described by Punithalingam and Woodhams (1982). The cultural characters of the Phyllosticta state of Guignardia mangiferae Roy along with the connection between the ascogenous state has also been thoroughly investigated (Punithalingam 1974). This work deals with the different stages of development of the conidiomata within the natural host and also in artificial culture media. It was observed that the Phyllosticta sp. investigated here produces temporary and permanent conidiogenous cells similar to those of Ascochyta spp. (Punithalingam 1979) and Macrophomina phaseolina (Punithalingam, 1982). Normally the temporary conidiogenous cells produce a single conidium after which they get sloughed off from the wall layers. In the present fungus, the temporary conidiogenous cells occasionally show one annellation indicating that it is capable of producing more than one conidium. Probably after producing one or two conidia, the temporary conidiogenous celis may become detached from the wall layers. 150 Another interesting finding is the presence of the thick sheath around the conidium. This sheath was also observed covering the conidiogenous cell and the conidium initial. The presence of the sheath around the conidium confirms observations made by light microscopy. Both transverse and longitudinal sections of the old conidia failed to reveal a sheath surrounding the conidia suggesting that the sheath gradually gelatinizes and disappears as the conidium becomes old. The dual conidiation process was found as a general feature of Ascochyta species in culture (Punithalingam 1979) and also in Macrophomina phaseolina (Punithalingam 1982). Other Coelomycete species studied by the authers, namely, Botryodiplodia theobromae Pat., Ciliochorella mangiferae Syd., Coleophoma cylindrospora (Desm.) Hohn., Pestalotiopsis palmarum (Cooke) Stey. and Urohendersonia pongamia Naj Raj & Ponnappa also show this type of dual conidiation process in culture. In the present investigation the Phyllosticta species also was found to produce temporary and permanent conidiogenous cells. But Robillarda depazeoides (Welw. et Curr.) Sacc. was found to produce only one type of conidiogenous cells during conidiation in culture. Therefore the present study reveals that necessary data collected based an the developmental morphology of conidiomata for many more Coelomycetes will be useful for segregating genera into differant groups. In addition, the ultrastructural details of the conidia given in the present investigation have provided useful’ information regarding the biological significance of the mucilaginous sheath around the conidium. REFERENCES Maas, J.L., Pollack, F.G. and Uecker, F.A. 1979. Morphology and development of Pilidiella quercicola. Mycologia 71: 92-102. Muthumary, J. and Vanaja,R. 1986. Development of conidiomata in Coniella fragariae. Trans. Br. Mycol. Soc. 87(1): 109-114. Punithalingam, E. 1966. Development of the pycnidium in Septoria. Trans. Br. Mycol. Soc. 49: 19-25, aona----- , 1974. Studies on Sphaeropsidales in culture II. Mycol. Pap. 136: 1-63. wennnnnee , 1979. Graminicolous Ascochyta Species. Mycol. Pap. 142: 1-214. wonnenn-- , 1981. Conidiation and appendage formation in Tiarosporella paludosa (Sacc. & Fiori) Hohnel. Nova Hedwigia 34: 539-567. --------- , 1982. Conidiation and Appendage formation in Macrophomina phaseolina (Tassi) Goid. Nova Hedwigia 36: 249-290. w-------- and Woodhams, J.E. 1982. The conidial appendage in Phyllosticta spp. Nova Hedwigia 36: 151-198. Reynolds, E.S. 1963. The use of lead citrate at high pH as an electron opaque stain in electron microscopy. Journal of cell Biology. 17: 208-212. 151 Fig.1. Transverse section of pycnidial primordium x 300 Figs.2-4. Transverse sections of young primordia showing the formation of central cavity. x 300 Figs.5-6. Transverse sections of young conidiomata showing the central cavities with the disentegrated hyphal fragments. x 300 152 oS Se Figs.7-8. Central cavities showing the sloughed off temporary conidiogenous cells with the conidia. x 528. Fig.9. Section of conidioma showing the formation of permanent conidiogenous cells. Note the presence of detached, temporary conidiogenous cells. x 2646 Figs.10-12. Vertical sections of mature conidioma. Fig.10 x 264; 11 x 132; 12 x 88. Figs.13-15. Vertical sections of conidicmata on leaf showing the papillae piercing the epidermis of the host 13 x 350, 14 x 350, 15 x 183. Figs. 16-18. Vertical sections of the conidiomata on leaf showing the papillae. 16 x 350, 17-18 x 183. Figs. 19-25. Transmission electron micrographs of the Phyllosticta state. Bar = 3um. Fig. 19. Section of mature conidium. Note the absence of the sheath surrounding the conidium. Fig. 20. Section of a young conidiogenous cell and conidium. Note the presence of the sheath around the conidiogenous cell and the conidium initial. Fig. 21. Section of a mature conidium showing three nuclei. Note the absence of a sheath around the conidium. Fig. 22. Section of a you ng conidium showing the sheath around the wall. Fig. 23. Section of more or less a mature conidium showing remnants of the sheath around the wall. 15D Figs. 24-25. Bar=3ym. Fig. 24. Section of a mature conidium showing remnants of the sheath around the conidium. Fig. 25. Section of a young conidium showing the sheath. Mave PHN ae Haye es 2 Dee ~ rey . cs ak Brae Ne ai) Sneha Lath beh ieicha hen ae ? . br eos it den dbenae i. ee AOR NT Bp Md 7 nf ta" v its Lert a ie ty eee ‘ Cag) 7 MY COTAXON Volume XLVI, pp. 157-159 April-June 1993 HYDNANGIUM PILA Pat., AN OLDER NAME FOR MARTELLIA MEDITERRANEA Moreno, Galan & Montecchi G. MORENO & R. GALAN Department of Plant Biology, Alcala de Henares University, 28871 Madrid, Spain In a recent publication (MORENO, GALAN & MONTECCHI, 1991), a new species belonging to the hypogeous fungi was described, Martellia mediterranea (Russulales), having been found in Spain, growing with Quercus suber and Cistus ladanifer. It was compared with Zelleromyces stephensii (Berk.) A. Smith and Martellia mistiformis Matt. It. was very surprising for us to note that at almost the same time in a local bulletin (VIDAL, 1991), a fungus "apparently" cospecific with the refered species was described in detail, identified as Hydnangium pila (PATOUILLARD, 1910) and with the proposal of the new combination: Martellia pila (Pat.) Vidal for it, as well as a list of possible different synonyms. In order to clarify this hypothetical taxonomical controversy, we decided to study a duplicate of Vidal's material (now in the private herbarium of J. M. Vidal, ref. 910523-3 and 901226-6), which was kindly sent to’ us by Mr. Vidal, as well as the type of Hydnangium pila Pat., which was received from the Farlow Herbarium (Harvard University, Cambridge) -Fig.1-, thanks to the kindness of Dr. D. H. Pfister. After carefully comparing those specimens with Martellia mediterranea, we must conclude that our suspicions were certain and all of them represent the same _ species, correctly named Martellia pila (Pat.) Vidal. At the same time we consider convenient to point out that the type of MHydnangium pila, consisting of three basidiocarps (two immature and the other one well preserved) shows a wide peridium -reaching 200-300 pm in diam. - with no sphaerocysts and having a remarkable gelification (Figs. 5 & 6). On the other hand, basidia are tetrasporic -exceptionally bisporic- measuring 28-40 x 14- 17 pm, and cystidia were not observed, probably due to the age of the fructification, according to Vidal's suggestion. ins sspite, sof yithis™ the strongly amiloid and _ spiny basidiospores, 9-11 % 9-10 wpm, are good diagnostic characters (Figs. 2-4). 158 ACKNOWLEDGEMENTS We wish to express our gratitude to Mr. Josep M. Vidal and Dr. Donald H. Pfister for loan of specimens and to Dr. Richard P. Korf for critical review of the manuscript, as well to Dr. F. Esteve for help in preparing the English text. LITERATURE CITED MORENO G., GALAN, R. & MONTECCHI, A. (1991). Hypogeous fungi from Peninsular Spain. II. Mycotaxon 42:201-238. PATOUILLARD, N. (1910).Note sur trois espéces d'Hydnangium de la flore du Jura. Bull. Soc. Mycol. France 26:199-204. VIDAL, J. M. (1991). Contribucién al conocimiento de la flora micolégica del Baix Emporda y zonas'7 limitrofes (Catalunya). IV. Hongos hipogeos (Zygomycotina, Ascomycotina y Basidiomycotina). Butll. Soc. Catalana Micol. 14-15:143- 194. Figs. I-6. Hydnangium pila Pat., type: Fig. 1. Original label handwritten by Patouillard. Figs. 2-4. Basidiospores. Fig.5. Cross-section of a sporocarp under light microscope. Fig.6. Texture of the peridium. ree ee ee Si: Brae it MY COTAXON Volume XLVI, pp. 161-176 April-June 1993 COMPUTER CODING OF STRAIN FEATURES OF THE GENUS RHIZOPUS SHUNG-CHANG JONG! and CANDACE MCMANUS? ‘American Type Culture Collection, 12031 Parklawn Drive, Rockville, MD 20852 USA and 7Microbial Systematics Section, National Institute of Dental Research, National Institutes of Health, Bethesda, MD 20892 USA ABSTRACT Members of the fungal genus Rhizopus play important roles in human, animal, and plant disease and in the fermentation industry. Identification of Rhizopus species is based primarily on the morphology of sporangiosporogenous structures, Zygospores, and asexual reproductive structures. The standardized coding system, the RKC Code, used for computer storage and analysis of microbial strain data was expanded to include features specific to the identification of Rhizopus species. BACKGROUND AND DISCUSSION Members of the genus Rhizopus occur as saprophytes on plant debris in soil. Many are known to cause soft rots or spoilage of vegetables and fruits. Some are etiologic agents of mucormycosis in humans and animals. Species of Rhizopus play important roles in the fermentation industry, especially in the production of enzymes (e.g., glucoamylase and lipase), organic acids, alcohol, and fermented foods such as bongkrek, lao-chao, tauco, and tempeh (Inui et al., 1965; Alexopoulos and Mims, 1979). The genus Rhizopus belongs to the family Mucoraceae under the order Mucorales. The genus was established by Ehrenberg (1820) and its history first reviewed by Zimmermann (1871). Rhizopus taxonomy has been the subject of many well known reports including those of Fischer (1892), Lendner (1908), Hanzawa (1915), Zycha (1935), Zycha etal. (1969), Naumov (1939), Inui et al. (1965), 162 Dabinett and Wellman (1973), Schipper (1984), Schipper and Stalpers (1984), and Liou et al. (1990). The taxonomy of Mucorales has been based primarily on morphological characteristics of sporangiosporogenous structures and zygospores. The genus Rhizopus, based on the type species R. stolonifer (Ehrenberg:Fries) Vuillemin, is characterized by mycelia bearing stolons and rhizoids and by formation of sporangiophores arising from the points of rhizoid attachment. Sporangia are differentiated at the tips of sporangiophores and are spherical (globose) with columellae and apophyses. Sporangiospores are produced within the sporangium outside the columellae and are mostly regular in shape and striated (Hesseltine and Ellis, 1961). Although the zygospore is the major morphological characteristic used for species delineation, asexual reproductive structures are most commonly used in routine identification of Rhizopus cultures. Since the genus Rhizopus was established, more and more differential characters were used for species delineation, and, as a consequence, many species in the genus were described. However, species differentiation was often, based on minor differences. In a numerical taxonomic analysis of data previously published by Inui et al. (1965), Dabinett and Wellman (1973) found that many of the 34 taxa included in the analysis were very closely related and that few of the 133 physiological and 16 morphological characters contributed significantly to the classification. More recently, other techniques have been used to study the taxonomy of the genus Rhizopus, including scanning electron microscopy of spore ornamentation (Schipper and Stalpers, 1984), mating experiments (Schipper, 1984; Schipper and Stalpers, 1984), and DNA homology studies (Ellis, 1985, 1986). Many previously described Rhizopus species are now considered to be synonymous, and the number of recognized species has been reduced considerably. Schipper (1984) and Schipper and Stalpers (1984) recognized only five species. Only two species were retained in the R. stolonifer-group -- R. stolonifer and R. sexualis, each with two varieties. In the R. microsporus-group, two species were also retained -- R. homothallicus and R. microsporus, the latter with four varieties. 163 The fifth species was R. oryzae. An additional species, R. azygosporus, was subsequently described by Yang and Jong (1984). Use of a standardized vocabulary for coding microbial strain data facilitates computer storage, retrieval, analysis, and exchange of such data. Data encoded in this manner can be used to build computer databases for study of taxonomic relationships and construction of identification keys or probability matrices for computer-aided identification of new isolates. In this communication, we present a comprehensive set of features important in the differentiation of Rhizopus species that have been incorporated into an existing computer coding system, the RKC Code. The RKC Code (after the original authors -- Rogosa, Krichevsky, and Colwell, 1971) is an open-ended, statement-oriented controlled vocabulary of descriptors of strain characteristics or features. The RKC Code was developed originally for bacterial strains and later expanded to include features specific for algae, protozoa, and selected groups of fungi and to allow coding of characteristics such as serology, phage typing, properties of nucleic acids, and quantitative antimicrobial susceptibility (Rogosa et al., 1986). More recently, the RKC Code was further expanded to include features used to characterize yeusts (Jong et al., 1988), the saprolegnian fungi (Jong et al., 1991), and the fungal genera Phytophthora (Jong et al., 1989) and Pythium (Jong et al., 1992). The set of characteristics developed for use with Rhizopus species is presented in the list below. Although some of these features were already part of the RKC Code, many of the descriptors are unique to Rhizopus species and were created specifically for this genus. New features added to the RKC Code for the Rhizopus species are marked with an "*" in the list. The features include morphological descriptions of rhizoids, stolons, sporangiophores, sporangia, columellae, sporangiospores, zygospores, azygospores, suspensors, and chlamydospores; colony characteristics; and growth temperatures. The terms used for morphological descriptions are based on the descriptions given in Hawksworth et al., 1983. 164 Rhizoids 008342: 008417: *043192: *043193: *043194: *043195: Stolons 008420: *043196: *043197: *043198: *043199: *(043200: *043201: * (043202: *043203: *043204: *043205: *043206: *043207: *043208: *(043209: *043210: *043211: *043212: *043213: Hyphae are septate. Rhizoids (root-like hyphae) are present. Rhizoid system is complex (highly branched). Rhizoids are pigmented. Rhizoids are brown. Rhizoids are black. Stolons are present. Stolons are pigmented. Stolons are brown. Stolons are 0.1-1.0 uw wide. Stolons are 1.1-2.0 u wide. Stolons are 2.1-3.0 yu wide. Stolons are 3.1-4.0 uw wide. Stolons are 4.1-5.0 u wide. Stolons are 5.1-6.0 u wide. Stolons are 6.1-7.0 « wide. Stolons are 7.1-8.0 4 wide. Stolons are 8.1-9.0 u wide. Stolons are 9.1-10 u wide. Stolons are 11-12 yu wide. Stolons are 13-14 uw wide. Stolons are 15-16 u wide. - Stolons are 17-18 yu wide. Stolons are 19-20 yw wide. Stolons are > 20 x wide. Sporangiophores 008539: Sporangia are on sporophores (sporangiophores). *043214: Sporangiophores occur singly. *043215: Sporangiophores occur in pairs. *043216: Sporangiophores occur in groups of 3-4. *043217: Sporangiophores are straight (erect). 165 008786: 008787: *043218: Sporangiophores are branched. Sporangiophore branching is irregular. Sporangiophore branching is dichotomous. *043219: 008792: 008562: *043220: *043221: *043222: *043223: *043224: *043225: *043226: *043227: *043228: *043229: *043230: *043231: Sporangiophores have swellings. Swellings are present at branch points of sporangiophore. Apophyses are present. Sporangiophores are pigmented. Sporangiophores are brown. Sporangiophores are black. Sporangiophores are < 201 u long. Sporangiophores are 201-300 uw long. Sporangiophores are 301-400 wu long. Sporangiophores are 401-500 yw long. Sporangiophores are 501-600 uw long. Sporangiophores are 601-700 yu long. Sporangiophores are 701-800 wu long. Sporangiophores are 801-900 uw long. Sporangiophores are 901-1000 uw long. Sporangiophores are 1001-1500 wu long. Sporangiophores are 1501-2000 yu long. Sporangiophores are 2001-3000 y long. *043232: *043233: *043234: *043235: *043236: *043237: *043238: *043239: *043240: *043241: *043242: *043243: *043244: *043245: *043246: *043247: *043248: *043249: *043250: Sporangiophores are > 3000 yu long. Sporangiophores are 0.1-1.0 uw wide. Sporangiophores are 1.1-2.0 u wide. Sporangiophores are 2.1-3.0 u wide. Sporangiophores are 3.1-4.0 yu wide. Sporangiophores are 4.1-5.0 wide. Sporangiophores are 5.1-6.0 4 wide. Sporangiophores are 6.1-7.0 u wide. Sporangiophores are 7.1-8.0 u wide. Sporangiophores are 8.1-9.0 u wide. Sporangiophores are 9.1-10 uw wide. Sporangiophores are 11-15 yu wide. Sporangiophores are 16-20 u wide. Sporangiophores are 21-25 yu wide. Sporangiophores are 26-30 u wide. Sporangiophores are > 30 yu wide. 166 Sporangia 043074: 008811: 008558: 008815: 008563: 008199: *043251: 008829: *043252: *043253: *043254: *043255: 008578: 008579: 008580: 008581: 008582: 008583: 008584: 008585: *(043256: *(043257: *043258: *043259: *043260: *043261: *043262: *(043263: *043264: *043265: *043266: *(043267: *043268: *(43269: Sporangia are present. Sporangia are terminal. Sporangia are spherical (length to breadth ratio is 1.0-1.05). Sporangia are prolate spheroidal (length to breadth ratio is 1.06-1.15). Sporangial walls are smooth. Sporangial wall disintegrates concurrently with release of spores. Sporangia are powdery. Sporangia are pigmented. Sporangia are blue. Sporangia are brown. Sporangia are gray. Sporangia are black. Sporangia are 1.0-2.0 uw wide. Sporangia are 2.1-3.0 u wide. Sporangia are 3.1-4.0 uw wide. Sporangia are 4.1-5.0 u wide. Sporangia are 5.1-10 u wide. Sporangia are 11-15 wu wide. Sporangia are 16-20 u wide. Sporangia are 21-30 uw wide. Sporangia are 31-40 mu wide. Sporangia are 41-50 mu wide. Sporangia are 51-60 uw wide. Sporangia are 61-70 su wide. Sporangia are 71-80 u wide. Sporangia are 81-90 u wide. Sporangia are 91-100 u wide. Sporangia are 101-150 u wide. Sporangia are 151-200 yu wide. Sporangia are 201-250 u wide. Sporangia are 251-300 su wide. Sporangia are 301-350 yu wide. Sporangia are 351-400 mu wide. Sporangia are > 400 uw wide. Columellae 008561: *043270: *043271: *043272: *043273: *043274: *043275: *043276: *043277: *043278: *043279: *043280: *043281: *043282: *043283: *043284: *043285: *043286: *043287: *043288: *043289: *043290: *043291: *043292: *043293: *043294: *043295: *043296: *043297: *043298: *043299: Columellae are formed. Columellae are spherical (length to breadth ratio is 1.0-1.05). Columellae are prolate spheroidal (length to breadth ratio is 1.06-1.15). Columellae are broadly ellipsoidal (length to breadth ratio is 1.16-1.30). Columellae are ellipsoidal (length to breadth ratio is 1.31-1.6). Columellae are conical. Columellae are applanate (flattened). 167 Columellae are pyriform (pear-shaped, attached at narrow end). Columellae are obpyriform (pear-shaped, attached at broad end). Columellae are cylindrical. Columellae are obovoid (egg-shaped, attached at narrow end). Columellae have collars. Columellae are pigmented. Columellae are gray. Columellae are brown. Columellae are 0.1-1.0 u long. Columellae are 1.1-2.0 uw long. Columellae are 2.1-3.0 u long. Columellae are 3.1-4.0 u long. Columellae are 4.1-5.0 uw long. Columellae are 5.1-6.0 u long. Columellae are 6.1-7.0 u long. Columellae are 7.1-8.0 u long. Columellae are 8.1-9.0 uw long. Columellae are 9.1-10 yu long. Columellae are 11-20 uw long. Columellae are 21-30 uw long. Columellae are 31-40 w long. Columellae are 41-50 wu long. Columellae are 51-60 yw long. Columellae are 61-70 yu long. 168 *043300: Columellae are 71-80 yu long. *043301: Columellae are 81-90 yu long. *043302: Columellae are 91-100 uw long. *043303: Columellae are 101-150 yu long. *043304: Columellae are 151-200 yu long. *043305: Columellae are > 200 yw long. *043306: Columellae are 0.1-1.0 u wide. *043307: Columellae are 1.1-2.0 u wide. *043308: Columellae are 2.1-3.0 uw wide. *043309: Columellae are 3.1-4.0 u wide. *043310: Columellae are 4.1-5.0 u wide. *043311: Columellae are 5.1-6.0 u wide. *043312: Columellae are 6.1-7.0 wide. *043313: Columellae are 7.1-8.0 u wide. *043314: Columellae are 8.1-9.0 u wide. *043315: Columellae are 9.1-10 uw wide. *043316: Columellae are 11-20 wu wide. *043317: Columellae are 21-30 u wide. *043318: Columellae are 31-40 u wide. *043319: Columellae are 41-50 u wide. *043320: Columellae are 51-60 yu wide. *043321: Columellae are 61-70 u wide. *043322: Columellae are 71-80 yu wide. *043323: Columellae are 81-90 uw wide. *043324: Columellae are 91-100 mu wide. *043325: Columellae are 101-150 yu wide. *043326: Columellae are 151-200 wu wide. *043327: Columellae are > 200 u wide. Sporangiospores 008173: Asexual spores (sporangiospores) are produced in sporangia (spore vesicles). 008592: Sporangiospores occur in masses. 008595: Sporangiospores are spherical (length to breadth ratio is 1.0-1.05). | *043328: Sporangiospores are prolate spheroidal (length to breadth ratio is 1.06-1.15). 169 Sporangiospores are broadly ellipsoidal (length to breadth ratio is 1.16-1.30). Sporangiospores are ellipsoidal (length to breadth ratio is 1.31-1.6). Sporangiospores are angular. Sporangiospores are fusiform. Sporangiospores are irregular in shape. Surfaces of sporangiospores are striated. Surfaces of sporangiospores are smooth. Surfaces of sporangiospores are spiny. Surfaces of sporangiospores are warty. *043329: 043047: *043330: *043331: 043053: *043332: 043054: 043056: 043057: 043061: 043068: 043067: 008602: 008603: 008604: *043333: *043334: *043335: *043336: *043337: *043338: *043339: *043340: 008597: 008598: 008599: 008600: 008601: *043341: *043342: *043343: *043344: *043345: *043346: *043347: *043348: *043349: Sporangiospores are pigmented. Sporangiospores are brown. Sporangiospores are black. Sporangiospores are 0.1-1.0 uw wide. Sporangiospores are 1.1-2.0 u wide. Sporangiospores are 2.1-3.0 uw wide. Sporangiospores are 3.1-4.0 u wide. Sporangiospores are 4.1-5.0 u wide. Sporangiospores are 5.1-6.0 “4 wide. Sporangiospores are 6.1-7.0 “ wide. Sporangiospores are 7.1-8.0 « wide. Sporangiospores are 8.1-9.0 wu wide. Sporangiospores are 9.1-10 uw wide. Sporangiospores are > 10 uw wide. Sporangiospores are 0.1-1.0 u long. Sporangiospores are 1.1-2.0 u long. Sporangiospores are 2.1-3.0 uw long. Sporangiospores are 3.1-4.0 u long. Sporangiospores are 4.1-5.0 x long. Sporangiospores are 5.1-6.0 uw long. Sporangiospores are 6.1-7.0 u long. Sporangiospores are 7.1-8.0 u long. Sporangiospores are 8.1-9.0 uw long. Sporangiospores are 9.1-10 yu long. Sporangiospores are 11-12 uw long. Sporangiospores are 13-14 yu long. Sporangiospores are 15-16 wu long. Sporangiospores are 17-18 uw long. 170 *043350: *043351: Sporangiospores are 19-20 yu long. Sporangiospores are > 20 wu long. Sexual Reproduction 008617: 008618: 008619: Sexual reproduction occurs. Strain is homothallic (both mating types on same mycelium). Strain is heterothallic (mating types on separate mycelia). Zygospores 008627: *043352: *043353: 008628: 008629: 008630: *043354: *043355: *043356: *043357: *043358: *043359: *043360: *043361: *043362: *043363: *043364: *043365: *043366: *043367: *043368: *043369: *043370: *043371: Zygospores are produced. Zygospores are spherical (length to breadth ratio is 1.0-1.05). Zygospores are prolate spheroidal (length to breadth ratio is 1.06-1.15). Zygospores are thick-walled. Surfaces of zygospores are rough. Surfaces of zygospores are spiny. Surfaces of zygospores are warty. Zygospores are pigmented. Zygospores are red. Zygospores are yellow. Zygospores are brown. Zygospores are black. Zygospores are ().1-1.0 « wide. Zygospores are 1.1-2.0 u wide. Zygospores are 2.1-3.0 uw wide. Zygospores are 3.1-4.0 u wide. Zygospores are 4.1-5.0 u wide. Zygospores are 5.1-6.0 u wide. Zygospores are 6.1-7.0 “ wide. Zygospores are 7.1-8.0 uw wide. Zygospores are 8.1-9.0 uw wide. Zygospores are 9.1-10 u wide. Zygospores are 11-15 yu wide. Zygospores are 16-20 wu wide. * (043372: * 043373: * (043374: *043375: *043376: *043377: *043378: *043379: *043380: *043381: *043382: *043383: Zygospores are 21-30 uw wide. Zygospores are 31-40 yu wide. Zygospores are 41-50 uw wide. Zygospores are 51-60 u wide. Zygospores are 61-70 yu wide. Zygospores are 71-80 u wide. Zygospores are 81-90 u wide. Zygospores are 91-100 yu wide. Zygospores are 101-150 wu wide. Zygospores are 151-200 u wide. Zygospores are 201-250 uw wide. Zygospores are > 250 mu wide. Azygospores *043384: Azygospores (parthenogenetic zygospores) are present. *043385: *043386: Azygospores are spherical (length to breadth ratio is Azygospores are terminal. 1.0-1.05). i7A *043387: Azygospores are prolate spheroidal (length to breadth ratio *043388: Surfaces of azygospores are rough. *043389: Surfaces of azygospores are spiny. *043390: Surfaces of azygospores are warty. *043391: *043392: *043393: *043394: *043395: *043396: *043397: *043398: *043399: *043400: *043401: *043402: *043403: *043404: *04340S: is 1.06-1.15). Azygospores are pigmented. Azygospores are brown. Azygospores are black. Azygospores are 0.1-1.0 “ wide. . Azygospores are 1.1-2.0 yu wide. Azygospores are 2.1-3.0 4 wide. Azygospores are 3.1-4.0 u wide. Azygospores are 4.1-5.0 4 wide. Azygospores are 5.1-6.0 wide. Azygospores are 6.1-7.0 4 wide. Azygospores are 7.1-8.0 4 wide. Azygospores are 8.1-9.0 u wide. Azygospores are 9.1-10 yu wide. Azygospores are 11-15 yu wide. Azygospores are 16-20 u wide. Lye *043406: Azygospores are 21-30 u wide. *043407: Azygospores are 31-40 u wide. *043408: Azygospores are 41-50 yu wide. *043409: Azygospores are 51-60 u wide. *043410: Azygospores are 61-70 yu wide. *043411: Azygospores are 71-80 su wide. *043412: Azygospores are 81-90 mu wide. *043413: Azygospores are 91-100 yu wide. *043414: Azygospores are 101-150 uw wide. *043415: Azygospores are 151-200 wu wide. *043416: Azygospores are 201-250 yu wide. *043417: Azygospores are > 250 u wide. Suspensors 008632: Two supporting cells (suspensors) are present. *043418: Suspensors (supporting cells) are equal in size. *043419: Both suspensors are spherical (globose). *043420: Only the larger suspensor is spherical (globose). Chlamydospores 008363: Chlamydospores are present. 008993: Chlamydospores occur singly. 008994: Chlamydospores are catenulate (in chains). 008996: Chlamydospores are spherical (length to breadth ratio is 1.0-1.05). *043421: Chlamydospores are prolate spheroidal (length to breadth ratio is 1.06-1.15). *043422: Chlamydospores are broadly ellipsoidal (length to breadth ratio is 1.16-1.30). | *043423: Chlamydospores are ellipsoidal (length to breadth ratio is 1.31-1.6). *043424: Chlamydospores are cylindrical. 008426: Chlamydospores are pigmented. *043425: Chlamydospores are refringent (refractive). *043426: Chlamydospore walls are smooth. *043427: Chlamydospores are 0.1-1.0 “ wide. *043428: *043429: *043430: *043431: *043432: *043433: *043434: *043435: *043436: *043437: *043438: *043439: *043440: *043441: *043442: *043443: *043444: *043445: *043446: *043447: *043448: *043449: *043450: *043451: *043452: *043453: *043454: *043455: *043456: *043457: Chlamydospores are 1.1-2.0 4 wide. Chlamydospores are 2.1-3.0 u wide. Chlamydospores are 3.1-4.0 u wide. Chlamydospores are 4.1-5.0 u wide. Chlamydospores are 5.1-6.0 u wide. Chlamydospores are 6.1-7.0 « wide. Chlamydospores are 7.1-8.0 u wide. Chlamydospores are 8.1-9.0 uw wide. Chlamydospores are 9.1-10 “ wide. Chlamydospores are 11-15 wu wide. Chlamydospores are 16-20 u wide. Chlamydospores are 21-25 yu wide. Chlamydospores are > 25 u wide. Chlamydospores are 0.1-1.0 uw long. Chlamydospores are 1.1-2.0 uw long. Chlamydospores are 2.1-3.0 u long. Chlamydospores are 3.1-4.0 u long. Chlamydospores are 4.1-5.0 u long. Chlamydospores are 5.1-6.0 u long. Chlamydospores are 6.1-7.0 uw long. Chlamydospores are 7.1-8.0 u long. Chlamydospores are 8.1-9.0 uw long. Chlamydospores are 9.1-10 u long. Chlamydospores are 11-15 yu long. Chlamydospores are 16-20 wu long. Chlamydospores are 21-25 yu long. Chlamydospores are 26-30 u long. Chlamydospores are 31-35 yu long. Chlamydospores are 36-40 yu long. Chlamydospores are > 40 uw long. Colony Pigmentation 173 020037: Nondiffusible pigment is produced. 020001: Colonies are pure (paper) white on solid medium. 020039: Nondiffusible brown pigment is produced. 020043: Nondiffusible golden (yellow) pigment is produced. 020002: Colonies are gray on solid medium. 174 Growth Temperatures 017032: 017012: 017013: 017066: 017037: 017014: 017033: 017052: 017034: 017074: 017043: 017017: 017018: 017019: 017020: Growth occurs at 5°C. Growth occurs at 10°C. Growth occurs at 15°C. Growth occurs at 17°C. Growth occurs at 20°C. Growth occurs at 25°C. Growth occurs at 30°C. Growth occurs at 33°C. Growth occurs at 35°C. Growth occurs at 36°C. Growth occurs at 40°C. Growth occurs at 45°C. Growth occurs at 50°C. Growth occurs at 55°C. Growth occurs at 60°C. Source of Isolation 002012: What was the specific source of isolation (e.g., kind of water, soil, etc., species and organ and tissue of plant, animal, etc.)? Acknowledgements This work was supported in part by National Science Foundation Grant DIR89-15137 to SCJ. The authors kindly thank Dr. Guozhong Ma for reviewing this paper. 175 REFERENCES Alexopoulos, C.J. and Mims, C.W. 1979. Introductory Mycology. John Wiley & Sons, New York, 632 pp. Dabinett, P.E. and Wellman, A.M. 1973. Numerical taxonomy of the genus Rhizopus. Can. J. Bot. 51:2053-2064. Ehrenberg, C.G. 1820. De Mycetogenesi. Bonnae. Ellis, J.J. 1985. Species and varieties in the Rhizopus arrhizus- Rhizopus oryzae group as indicated by their DNA complementarity. Mycologia 77:243-247. Ellis, J.J. 1986. Species and varieties in the Rhizopus microsporus group as indicated by their DNA complementarity. Mycologia 78:508-5 10. Fischer, A. 1892. Phycomycetes. In Rabenhorst’s Kryptogamen- flora, Die Pilze, Vol 4. Kummer, Leipzig, 505 pp. Hanzawa, J. 1915. Studien uber einige Rhizopus-Arten. Mykol. Zentralbl. 5:230-246. Hawksworth, D.L., Sutton, B.C., and Ainsworth, G.C. 1983. Ainsworth & Bisby’s dictionary of the fungi (including the lichens), 7th ed. Commonwealth Mycological Institute, Kew, Surrey, England, 445 pp. Hesseltine, C.W. and Ellis, J.J. 1961. Notes on Mucorales, especially Absidia. Mycologia 53:406-426. Inui, T., Takeda, Y., and lizuka, H. 1965. Taxonomical studies on genus Rhizopus. J. Gen. Appl. Microbiol. 11(suppl.):1-121. Jong, S.-C., Davis, E.E., McManus, C., and Krichevsky, M.I. 1991. Computer coding of strain features of the saprolegnian fungi. Mycotaxon 41:407-418. Jong, S.-C., Ho, H.H., McManus, C., and Krichevsky, M.I. 1989. Computer coding of strain features of the genus Phytophthora. Binary 1:187-193. Jong, S.-C., Ho, H.H., McManus, C., and Krichevsky, M.I. 1992. Computer coding of strain features of the genus Pythium. Mycotaxon 44:301-314. Jong, S.-C., Holloway, L., McManus, C., Krichevsky, M.I., and Rogosa, M. 1988. Coding of strain features for computer-aided identification of yeasts. Mycotaxon 31:207-219. 176 Lendner, A. 1908. Les Mucorinées de la Suisse. Matériaux pour la flora cryptogamique Suisse. K.-J Wyss, Berne, 180 pp. Liou, G.Y., Chen, C.C., Hsu, W.H., and Chien, C.Y. 1990. Atlas of the genus Rhizopus and its allies. Mycological monograph No. 3. Food Industry Research and Development Institute, Taiwan, 32 pp. Naumov, N.A. 1939. Clés des Mucorinées. Encyclopédia mycologique, Vol. 9. Lechevalier, Paris, pp. 1-137. Rogosa, M., Krichevsky, M.I., and Colwell, R.R. 1971. Method for coding data on microbial strains for computers (edition AB). Int. J. Syst. Bacteriol. 21:1A-184A. Rogosa, M., Krichevsky, M.I., and Colwell, R.R. 1986. Coding microbiological data for computers. Springer-Verlag, New York, 299 pp. Schipper, M.A.A. 1984. A revision of the genus Rhizopus. I. The Rhizopus stolonifer-group and Rhizopus oryzae. Stud. Mycol. 25:1-19. Schipper, M.A.A. and Stalpers, J.A. 1984. A revision of the genus Rhizopus. WU. The Rhizopus microsporus-group. Stud. Mycol. 25:20-34. Yang, G.F. and Jong, S.C. 1984 A new obligate azygosporic species. Mycotaxon 20:397-340. Zimmermann, O.E.R. 1871. Das genus Mucor. In Programm der Realschule I. Ordnung zu Chemnitz. Zycha, H. 1935. Pilze II. Mucorineae. In Kryptogamenflora der Mark Brandenburg, Vol. 6a. Gebruder Borntraeger, Leipzig, 264 pp. Zycha, H., Siepmann, R., and Linnemann, G. 1969. Mucorales. Eine Beschreibung aller Gattungen und Arten _ dieser Pilzgruppe. J. Cramer, Lehre, 355 pp. MY COTAXON Volume XLVI, pp. 177-182 April-June 1993 THREE NEW HYALOSCYPHACEOUS FUNGI FROM NORWAY AND GREENLAND SIGURD OLSEN * Skarveien 7, N-8550 Lédingen, Norway JOHN HAINES Biological Survey, rm 3132 CEC, NYS Museum Albany, NY 12230, U.S.A. SIGMUND SIVERTSEN UNIT, Botanical Department, N-7004 Trondeim, Norway SUMMARY: Three species of Hyaloscyphaceae from Norway and Greenland, Urceolella aasii sp. nov. on Urtica, Hyalopeziza archangelica sp. nov. on Angelica/Heracleum, and Hyalopeziza groenlandica sp. nov. on Angelica, are described and briefly discussed. Collections were made by the first author unless otherwise stated. KEY WORDS: Greenland; Hyaloscyphaceae; Norway Urceolella aasii Olsen & Haines, sp. nov. Pig) Apothecia diametro usque ad .5 mm, aurantio-ochracea. Pili usque ad 250 um longi, 6- 16 pm lati, non-septati, praecrassotunicati, 11 todo non colorati, tabida in 3% KOH. Sporae 6.5-9.5 um longae. 1.0-1.8 um crassi. Ad caulis emortuis Urticae. Holotypus: Norway, Nordland, Rana: Ytteren. On Urtica dioica. 22 V 1991, S. Olsen. In herb BG. Etymology: Named in honor of Norwegian mycologist Olav Aas. Apothecia up to .5 mm diam., orange to ochraceous, seated on very short stipe ca. 2/3 the diam. of the cup, disc orange-ochraceous, exposed and concave when moist, covered by hairs when dry. Excipulum thin-walled, buff-orange fextura angularis composed of cells ca. 12 4m in diam., outer layer flattened, almost forming a cortex. Subhymenium and inner cells of excipulum containing orange oil-globules in revived exiccata. Hairs up to 250 wm by 6-16 wm diam., cylindric or tapered to a blunt point at the apex, straight or occasionally slightly curved or forked, non-septate with a fine *Sigurd Olsen died November 23, 1992 178 lumen 1-2 um wide except at the base where it is wider, hyaline, smooth, arising perpendicularly from the margin or outer exciple, sometimes with a plate-like or rooting base, flexible, not easily breaking, slowly and almost completely dissolving in warm 3% KOH to leave only a ghost-like outline, unchanged in Melzer’s solution. Some hairs agglutinated apically by hyaline amorphous matter which stains in cotton blue. Hair walls and lumen unstained with cotton blue. Short cylindrical hairs with hemispherical tips and apically enlarged lumen also present. Asci 35-45 (57) x 4.5-6.0 um, 8-spored, cylindrical or enlarged in the mid portion, subtended by a crozier at least in early development, with a minute apical pore plug which turns bright blue in Melzer’s solution. Spores 6.5-9.5 1.0-1.8 (2.5) um, non-septate, cylindric-fusiform, slightly enlarged above, straight or very slightly curved, without conspicuous inclusions. Paraphyses evenly cylindrical, 1.3-1.7 um wide level with the ascus tips in the hymenium, containing yellowish oil globules, straight, unbranched except at the base. Host: Known only from moist previous years stems of Urtica dioica in Spring or summer. The type material also contains Crocicreas, Leptosphaeria, and a pycnidial fungus. Distributions: Known only by three collections from near or above the Arctic Circle in Norway. Collections examined: NORWAY; Trons., Harstad, Sdrvik. 27 VI 1989 on Urtica dioica in herb. BG; Nordland, Rana, Ytteren. 25 V 1990 on Urtica dioica in herbs. BG, NYS; See also type. This small, herbicolous discomycete fits into the Hyaloscyphaceous genus Urceolella as defined by Korf & Kohn (1980), with its long, thick-walled hairs that dissolve in KOH and remain unstained in Melzer’s solution. It is distinct within Urceolella with its orange to ochraceous pigmentation and long hairs up to 250 um arising perpendicularly from the substrate. It appears to be closely related to U. crispula, from which it is distinguished by its larger size, larger spores, and colored disc as compared to the watery white disc of U. crispula. Raschle (1977) describes U. crispula as white or light beige. Another, perhaps even more closely related, though isolated species, U. tuberculiformis, has the same type of hairs with T-like bases. This species, as described by Dennis (1963), differs in having no pigmentation, although Raschle (1977) describes it as yellowish white with a thick-walled yellow excipulum, and asci, ca. 70 x 10 wm, longer and broader spores (10-12 x 2.5-3.0 um), and growing on Aquilegia in the USA and Adenostyles in Italy. Figs. 1-5. 1. Urceolella aasii, Hairs, asci, paraphyses, and spores from Paratype, S6rvik. 2-4. Hyalopeziza archangelica. 2. Asci, paraphyses, hairs, and spores from holotype. 3. Spores and hairs from paratype, Trondheim. 4. Spores from paratype specimen, Skanland. 5. Hyalopeziza groenlandica Ascus, paraphyses, hairs, and spores from holotype. 180 Hyalopeziza archangelica Olsen & Sivertsen, sp. nov. Fig. 2-4 Apothecia cupulata, sessilia, ad 1 mm diametro, rose, arcte setulosa. Pili hyalini, 25-65 uum longis, basi ad 6 um crassi, solidis. Asci 70-110 um longis, 15-17 4m crassi, in Iodo non colorati. Paraphysis cylinribus. Sporae 10-15 (18.5) um longis, 4.0-6.5 (-7.0) pm crassi, 0-3-septatae. Ad caulis emortuis Angelica et Heracleum. Holotypus: Norway, Nordland, Rana, Fisktj6nna. On Angelica archangelica, 26 V 1990. Olsen & Sivertsen. In herb. BG. Etymology: Named for its substrate, Angelica archangelica. Apothecia sessile, cupulate, up to 1 mm across, mostly smaller, gregarious, whitish to pink. Rim and flanks set with scattered, short hairs. Excipulum of thick-walled, hyaline, angular, + isodiametric cells 6, 5-10 wm across. Hairs 25-70 ym. long, refractive, solid above with a lumen at the base, + straight, rather evenly tapering towards an acute apex. Not loosing glassiness in warm 10% KOH and not colored by Meltzer’s or Congo Red. Asci 70-110, 15-17 um thin-walled, some apical thickening when immature, 8-spored, arising from a crozier. Spores 10-15 (18.5) x 4-6 (7) um, hyaline, smooth, 0-2-, mostly 1-septate, often with prominent oil-globules. Paraphyses cylindrical, 2.0-2.8 jm thick, with rounded apex, sparingly septate, sometimes branched above the middle, not overtopping the asci, scarce. Host: On basal parts of previous years stems of Angelica and Heracleum. Distribution: Probably widespread over the northern hemisphere, but overlooked because of its small size and nondistinctive color. A specimen cited by Korf (1980) and presumed to be the same, is from Finmark, Norway, but it has not been examined by the authors. Collections examined: NORWAY, Nordland; Rana, Fisktjonna, 20 V 1991 on Angelica, S. Olsen, herbarium BG; ‘Tromso, Skdnland, near airport, 11 VI 1991 on Heracleum, S. Olsen, herbarium BG; Sér-Trondelag, Trondheim, Lillegardsbakken, III 1992, on Heracleum, S. Sivertsen, herbarium BG; GREENLAND, Kuanit, east of Godhavn, on Angelica, P. Milan Petersen, herbarium CUP as 5946; Also see type. This is the same fungus from Finmark and Greenland as tentatively refered to as "Chaetonaevia archangelica" by Korf (1980). Korf clearly did not intend to legitimize the name according to the code in that publication and since it has so many similarities to Hyalopeziza, it will be described here as a Hyalopeziza in subgenus Unguicularia as defined by Korf & Kohn (1980). Its hairs are unchanged by KOH, and the lumen is present only at the base. The excipulum is composed of + isodiametric angular cells, which, according to Huhtinen (1987) is acceptable in the genus. The species seems to be closely related to Hyalopeziza raripila (H6hn.) Huhtinen, but differs in its larger size, different color, longer hairs, asci, and spores which are septate at maturity. 181 Hyalopeziza groenlandica Olsen, sp. nov. Fig. 5 Apothecia gregaria, cupulata, ad 0.5 mm lata, sessilia, ochraceo-rosei. Pili hyalini, ad 30 um longis, in apicem solidis attenuatis. Asci 55-60 4m longis, octosponi, poro in Todi non colorati. Paraphyses cylindracea, apicem solidis attenuatis. Sporae 15-20 x 2.8-4.0 uum. Ad caulis emortuis Angelicae. Holotypus: Greenland, Kuanit, approx. 4 km. east of Godhavn. Leg. P. Milan Petersen 5 VIII 1970 in herbarium CUP (as CUP 59588). Etymology: For the type locality, Greenland. Apothecia yellowish-pink, sessile, cupulate, gregarious, up to 0.5 mm across. Margin with short, hyaline hairs. Excipulum of slightly thick-walled, elongate, hyaline cells. Hairs 25-35 um long, refractive, tapering, irregularly flexuose, hooked apically, straight or sharply bent basally, with lumen at the base, refractivity not lost in warm 10% KOH, without color change in Meltzer’s solution. Asci 55-60 um long, somewhat thick-walled, 8-spored, without croziers, pore not blued in Melzer’s solution even with KOH pretreatment. Spores 15-20 x 2.8-4.0 um, wide fusiform, slightly curved, aseptate, aguttulate. Paraphyses numerous, with irregular refractive, apical protuberances, unchanged in KOH or Melzer’s, slightly exceeding the asci. Host: Previous year’s stems of Angelica. Distribution: So far known only from the type collection. Korf (1980) tentatively assigned the name "Laetinaevia archangelica" to this collection, but he was clear in not formally describing it as a new species. The first author is inclined to assign this species to Hyalopeziza subgenus Unguiculella (sensu strictu) Korf & Kohn (1980) on the basis of its glassy hairs and paraphyses. This combination requires that a species name other than Korf’s be chosen as the epithet "archangelica" is occupied by the new species Hyalopeziza archangelica Olsen & Sivertsen described above. Korf stresses the paraphyses, which he described as "coiled" and, although the general appearance is very much the same as H. _archangelica Olsen & Sivertsen it differs in the glassy protuberances of the paraphyses, as well as its quite different spores. 182 ACKNOWLEDGEMENTS The authors wish to thank Dr. Linda Kohn for her prepublication review of this work. LITERATURE CITED Dennis, R. W. G. 1963. A redisposition of some Fungi ascribed to the Hyaloscyphaceae. Kew Bulletin 17: 319-379. Huhtinen, S. 1987 (1988). Five glassy-haired Hyaloscyphaceae. Karstenia 27: 8-14. Korf, R. P. 1980. Inoperculate discomycetes of the arctic and alpine zones of Finmark, Lapland, and Greenland. In G. Laursen & J. Ammirati. Arctic and Alpine Mycology I: 27-37. Korf, R. P. & Linda M. Kohn 1980. Revisionary studies in the Hyaloscyphaceae. I. On Genera with "glassy" hairs. Mycotaxon 10: 503-512. Raschle, P. 1977. Taxonomische Untersuchungen and Ascomyceten aus der Familie der Hyaloscyphaceae Nannfeldt. Sydowia 29: 170-236. MY COTAXON Volume XLVI, pp. 183-192 April-June 1993 TAXONOMIC HISTORY OF THE OOMYCETE GENUS THRAUSTOTHECA WILL H. BLACKWELL AND MARTHA J. POWELL Department of Botany, Miami University, Oxford, Ohio 45056 ABSTRACT The nomenclature of Thraustotheca(Saprolegniales) is reviewed. The rationale for the proper citation of authorship of TT. clavata as (de Bary ex Biisgen) Humphrey is explained. It is concluded that at present only two validly published or adequately described species of Thraustotheca_ should be recognized, T. clavata and T. primoachlya. The inclusion of the latter taxon in the genus is questioned based on current consideration of primary sporangial characters as most significant in generic delimitations. Many genera of Oomycetes need contemporary taxonomic and nomenclatural review. Sparrow’s 1960 review of Oomycete genera in his monumental work on Phycomycetes is the most recent extensive synoptic treatment of the family Saprolegniaceae. Dick (1973) provided a list of genera and species of the Saprolegniaceae in chronological order of their descriptions. New species of the family have been described in more recent years, yet within the last forty years the only genera of Saprolegniaceae which have been monographed are Achlya (Johnson, 1956), Aphanomyces (Scott, 1961) and Saprolegnia (Seymour, 1970). In addition 184 Langsam (1986, 1987) has carried-out revisionary investigations on the genus Brevilegnia . In the following account we review the current taxonomic state of one genus of the Saprolegniaceae, Thraustotheca Humphrey. We clarify nomenclatural errors and identify taxa which require revisionary study. History and Authorship of Thraustotheca clavata The original species of the genus Thraustotheca Humphrey (1893), 7. clavata , is based on the taxon Dictyuchus clavatus. Authorship of the epithet c/avatus is usually credited (Coker and Hyman, 1912; Coker, 1923; Coker and Matthews, 1937; Shanor, 1937; Wolf, 1944; Sparrow, 1960) solely to Anton de Bary (1888), although some also noted (Coker and Hyman, 1912; Weston, 1918; Coker and Matthews, 1937; Shanor, 1937) that Busgen (1882) first incidentally published the species (i.e., D. clavatus ). Busgen’s (1882) publication on sporangial and spore formation provided an extensive description of asexual development of D. clavatus , and as well an account of sexual structures of the organism, in effect a complete description. However, Busgen credited the species to de Bary, who had received it in 1880 as part of a mixed algal collection obtained by E. Stahl at Vendenheim, Germany. A preferred citation of authorship of Dictyuchus clavatus would thus be de Bary ex Biisgen (1882) or simply Busgen (cf. Article 46.3, CURSE Code of Botanical Nomenclature, Grueter et al., 1988). De Bary mentioned Dictyuchus clavatus in his 1884 text without description, referencing his student’s (i.e., M. Busgen’s) dissertation. De Bary’s 1888 actual description of the taxon provided additional morphological information, but this is not relevant to the point of its valid publication. In fact, de Bary’s 1888 publication was a posthumous paper edited by Solms-Laubach (Weston, 1918), and it is difficult to ascertain 185 de Bary’s intent as to the description of D. clavatus as a new species. Regardless, a proper citation of authorship under Thraustotheca would be T. clavatus (de Bary ex Busgen) Humphrey (1893), as listed in Farr et al. (1979). Concept of the Genus Thraustotheca Solms-Laubach (in de Bary, 1888) first noted the possible generic distinction of D. clavatus within the Saprolegniales; this was reiterated by Fischer (1892). Humphrey subsequently (1893) described the genus Thraustotheca based on this taxon, but without benefit of living material. Whereas Saccardo (1899) cited the date of publication of the generic name as 1892, Clements and Shear (1931) cited it as 1893. This discordance over date of publication is resolved by understanding that Humphrey read his manuscript on Saprolegniaceae of the United States before the American Philosophical Society in November of 1892, but the Proceedings were published in 1893, the actual year of effective publication of the name. Coker and Hyman (1912) clearly stated the morphological delimitation of Thraustotheca (i.e., T. clavata). Over the years the generic concept of Thraustotheca , however, has evolved (Coker and Couch, 1923, 1924; Coker and Braxton, 1926; Coker, 1927; Coker and Matthews, 1937; Wolf, 1944; Sparrow, 1960) , and we can summarize it now as: those Saprolegniaceae producing a thraustothecoid sporangium (=generally club-shaped or subcylindric with an inoperculate, quickly disintegrating or rupturing sporangial wall), which contains no dictyuchoid network but which usually contains numerous primary cystospores (i.e., primary motile stage suppressed); oogonia with multiple oospores; and hyphae with relatively broad diameters which form a spreading hyphal mass. 186 These characters, in combination and in predominance, distinguish Thraustotheca from other genera of the Saprolegniaceae which are often considered morphologically similar: e.g. Achlya, Brevilegnia, Calyptralegnia, Dictyuchus, and Geolegnia. Setting generic limits within the Saprolegniaceae is sometimes problematic because of morphological plasticity. Some genera exhibit more than one type of sporangial discharge, yet it is generally accepted that the mechanism of sporangial discharge is a primary character distinguishing genera. For example, one species of Thraustotheca recognized (T. primoachlya ) does not exclusively produce sporangia with thraustothecoid type of spore discharge. Decisions on generic placements have traditionally been based on which sporangial type predominates (Coker, 1923). Instead, more recently, the type of spore discharge found in first-formed sporangia (Langsam, 1986) has been used for generic assignment. Accordingly, Langsam (1986) transferred Brevilegnia bispora to Achlya because of achlyoid spore discharge of primary sporangia and minisporangia. These viewpoints (i.e, relative importance of predominant versus initial sprorangial type) caution that clearer delimitation of taxa requires developmental studies of isolates grown under varying environmental conditions, as well as numerical comparison of genera using all putatively distinguishing generic characteristics. Nomenclatural Summary Several taxa (in addition to 7. clavata) have been described within Thraustotheca , two of these (T. achlyoides and TJ. unisperma ) eventually constituting bases for recognition of new genera (respectively Calyptralegnia and Brevilegnia) of Oomycetes. Following is a chronological nomenclatural summary of names published as taxa of Thraustotheca. Names in boldface are presently construed as valid taxa of the genus. 187 Thraustotheca Humphrey, 1893 Thraustotheca clavata (de Bary ex Biisgen) Humphrey, 1893. The basionym is Dictyuchus clavatus de Bary ex Biisgen, 1882. Citation as D. clavatus Busgen is equally permissible (Article 46.3, ICBN). Thraustotheca caucasica Voronov, 1922. This name has been overlooked by virtually all authors. It was, however, noted in 1969 in Index of Fungi, a Supplement to Petrak’s Lists. Although validly published, T. caucasica must be considered a doubtful or exluded taxon. Though seemingly similar to T. primoachlya , e.g., possessing tuberculate oogonia, there is insufficient descriptive information (including the lack of an illustration, as well as any information on antheridia) to make a positive determination of identity. Should it be possible to determine that it is the same as T. primoachlya , the epithet caucasica would assume priority. Thraustotheca achlyoides Coker and Couch, 1923. = Calyptralegnia achlyoides (Coker and Couch) Coker, 1927. Thraustotheca primoachlya Coker and Couch, 1924. Although we are recognizing J. primoachlya herein, further assessment of this taxon as a species of Thraustotheca is needed. In contrast to T. clavata, this species has androgynous antheridial branches and primary sporangia of the achlyoid type (producing the usually predominating thraustothecoid type sporangia later). 188 Thraustotheca unisperma Coker and Braxton, 1926. = Brevilegnia unisperma (Coker and Braxton) Coker and Couch in Coker, 1927. This is tantamount to transfer of Thraustotheca unisperma var. unisperma, an autonym created when var. /itoralis (see below) was described (Coker and Braxton, 1926). Thraustotheca unisperma var litoralis Coker and Braxton, 1926. = Brevilegnia unisperma var. litoralis (Coker and Braxton) Coker and Couch in Coker, 1927. In Coker’s work, the transfer to Brevilegnia is erroneously credited to Coker and Braxton. Because of similarities of sporangial structure and dehiscence, Salvin (1942) seemingly proposed that Brevilegnia be only a subgenus of Thraustotheca , and hence that all taxa described in or transferred to Brevilegnia should be returned toZhraustotheca. Based on total morphology (including one-spored oogonia), we favor retention of Brevilegnia as a separate genus, and do not favor such transfer of taxa back to Thraustotheca. We note that Langsam (1987), though excluding certain species, nonetheless retained the genus Brevilegnia . Thraustotheca irregularis Coker and Ward in Ward, 1939. This isa nomen invalidum, lacking a Latin diagnosis. Should it become desirable to recognize this taxon, it would be necessary to provide such descriptive information in Latin. Dick (1973) questioned the status of T. irregularis. Johnson (1956) considered the taxon possibly similar to Achlya dubia Coker (1923), in effect questioning its acceptance as a species. Two taxa, thus, are presently recognized as valid species of Thraustotheca, T. clavata and T. primoachlya. This is in fact the composition of the genus arrived at de facto by Coker (1927) when T. achlyoides and T. unisperma were transferred out of the genus. This circumscription 189 recognizes that T. primoachlya does not exclusively produce thraustothecoid sporangia, and hence at this time we are following Coker’s (1923) conservative concept of predominance of sporangial discharge for generic assignment. Thraustotheca clavata shows only thraustothecoid sporangia, whereas 7. primoachlya demonstrates a progression from achlyoid to thraustothecoid sporangial structures. Since Achlya dubia Coker , like T. primoachlya , can produce both achlyoid and thraustothecoid types of sporangia, both taxa should be _ investigated further. In fact, based on diclinous antheridia and primary sporangia of the thraustothecoid type, Achlya dubia may be more similar to T. clavata than is T. primoachlya.. Revisionary studies on species of Achlya and Thraustotheca with multiple types of sporangial discharge will have to be conducted to decide the validity of including species which demonstrate achlyoid sporangial discharge, even if only for a limited duration of the asexual cycle, within Thraustotheca. Central to the issue is whether primary sporangial type should take precedence over predominance of sporangial type. The dilemma is complex because temperature may influence the relative ratio of each type produced (Alabi, 1972). The current study is foundational for future investigations. Key to Recognized Species of Thraustotheca 1. Sporangia all thraustothecoid, club-shaped; oogonia smooth; antheridia diclinous.......... T. clavata 2. Early sporangia achlyoid, later ones thraustothecoid, often more elongate; oogonia with blunt projections; antheridia androgynou................ T. primoachlya 190 ACKNOWLEDGEMENTS We are indebted to the following persons for their helpful and critical review of this manuscript: Dr. Deborah M. Langsam, Department of Biology, The University of North Carolina at Charlotte; Dr. Michael A. Vincent, Department of Botany, Miami University; and Dr. David E. Padgett, Department of Biological Sciences, The University of North Carolina at Wilmington. This work was supported by a grant (BSR 91-07451) from the National Science Foundation. LITERATURE CITED ALABI, R.O. 1972. Thraustothecoid sporangium formation in three Saprolegniaceae from Nigeria. Trans. Brit. _. Mycol. Soc. 58: 519-521. BUSGEN, M._ 1882. Die Entwicklung der Phycomycetensporangien. Jahrb. Wiss. Bot. 13: 253-285. CLEMENTS, F.E. and C.L. SHEAR. 1931. The Genera of Fungi. The H.W. Wilson Co., New York. iv + 496 p. and 58 pits. COKER, W.C. 1923. The Saprolegniaceae, with Notes on Other Water Molds. Univ. of North Carolina Press, Chapel Hill. 201p. and 63 pits. COKER, W.C. 1927. Other water molds from soil. J. Elisha Mitchell Sci. Soc. 42: 207-226. COKER, W.C. and H.H. BRAXTON. 1926. New water molds from soil. J. Elisha Mitchell Sci. Soc. 42: 139-149. COKER, W.C. and J.N. COUCH. 1923. A new species of Thraustotheca. J. Elisha Mitchell Sci. Soc. 39: 112-114. COKER, W.C. and J.N. COUCH. 1924. Revision of the genus Jhraustotheca , with a description of a new species. J. Elisha Mitchell Sci. Soc. 40: 197-201. COKER, W.C. and O.W. HYMAN. 1912. Thraustotheca clavata. Mycologia 4: 87-90. COKER, W.C. and V.D. MATTHEWS. 1937. Saprolegniales. North America Flora 2: 15-67. 191 DE BARY, A. 1884. Vergleichende Morphologie und Biologie der Pilze, Mycetozoen und Bacterien. Wilhelm Engelmann, Leipzig. xvi + 558 p. DE BARY, A. 1888. Species der Saprolegnieen. Bot. Zeit. 46: 597-610, 613-621, 629-636, 645-653. (Paper edited posthumously by H.G. Solms-Laubach. ) DICK, M.W. 1973. Saprolegniales. IN: The Fungi, An Advanced Treatise. Ainsworth, G.C., F.K. Sparrow, and A.S. Sussman, eds. Academic Press, New York and London. pgs. 113-144. FARR, E.R., J.A. LEUSSINK and F.A. STAFLEU (eds.). 1979. Index Nominum Genericorum (Plantarum), Vol. 3. Reg. Veg. 102: 1277-1896. FISCHER, A. 1892. Saprolegninae. IN: Rabenhorst’s Kryptogamenflora 1(4): 310-383. GREUTER, W., H.M. BURDET, W.G. CHALONER, V. DEMOULIN, R. GROLLE, D.L. HAWKSWORTH, D.H. NICOLSON, P.C. SILVA, F.A. STAFLEU, E.G. VOSS and J. McNEILL. 1988. International Code of Botanical Nomenclature (ICBN) . Adopted by the Fourteenth International Botanical Congress, Berlin, July-August, 1987. Regnum Vegetabile 118: xiv + 328 p. HUMPHREY, J.E. 1893. The Saprolegniaceae of the United States, with notes on other species. Trans. Amer. Phil. Soc. (N.S.) 17: 63-148. INDEX OF FUNGI. 1969. A Supplement to Petrak’s Lists. 1920-1939. Commonwealth Mycological Institute, Kew, Surrey. p. 153. (citation source of T. caucasica ). JOHNSON, T.W. 1956. The genus Achlya: Morphology and Taxonomy. Univ. Michigan Press, Ann Arbor. xv + 180 p. LANGSAM, D.M. 1986. Achlya bispora: A taxonomic reassessment of Brevilegnia bispora . Mycologia 78: 600- 604. LANGSAM, D.M. 1987. Notes on the genus Brevilegnia: Two exluded taxa. Mycologia 79: 323-324. SACCARDO, P.A. 1899. Sylloge Fungorum 14: 36. 192 SALVIN, S.B. 1942. Variations of specific and varietal character induced in an isolate of Brevilegnia. Mycologia 34: 38-51. SCOTT, W.W. 1961. A monograph of the genus Aphanomyces. Virginia Agri. Exp. Sta. Tech. Bull. 151: 1-95. SEYMOUR, R.L. 1970. The genus Saprolegnia. Nova Hedwigia 19: iv -124. SHANOR, L. 1937. Observations on the development and cytology of the sexual organs of Thraustotheca clavata (de Bary) Humph. J. Elisha Mitchell Sci. Soc. 53: 119-136. SPARROW, F.K. 1960. Aquatic Phycomycetes, 2nd. rev. ed. Univ. Michigan Press, Ann Arbor. xxv + 1187 p. VORONOV, I. 1922. Thraustotheca caucasica. Mater. Mikol. Fitopat. Ros. 4(1): 57-58. WARD, M.W. 1939. Observations on a new species of Thraustotheca. J. Elisha Mitchell Sci. Soc. 55: 346- 352. WESTON, W.H. 1918. The development of Thraustotheca, a peculiar water-mould. Ann. Bot. 32: 155-173. WOLF, F.T. 1944. The Aquatic Oomycetes of Wisconsin. Part 1. Univ. Wisconsin Press, Madison. 64 p. MYCOTAXON Volume XLVI, pp. 193-199 April-June 1993 BIOSTATISTICAL CHARACTERIZATION OF THE GENUS OIDIUM MARCO T. IALONGO Istituto Sperimentale per la Patologia Vegetale, Via C.G. Bertero 22, Rome, Italy A method that takes advantage of biostatistical analysis is proposed for differentiating between species of the genus Oidium (anamorph of Erysiphoideae), since it is often difficult to distinguish one from the other using the traditional parameters. In addition to the usual parameters, new quantitative morphological parameters are introduced: area, circumference, maximum diameter, shape factor of the circumference, shape factor of the area of the appresoria; true length of conidial foot-cell; area, circumference, shape factor of circumference, shape factor of the area of the conidia. Measurements were made of the anamorphs of various species of Erysiphaceae selected from those that have indistinguishable forms according to the traditional measurement methods. The series of values obtained for the single parameters were compared two by two (the values of each sample with the corresponding values of all the others), calculating Student’s t for each comparison. The average linkage cluster analysis was then applied, and the relative dendrogram constructed. The latter demonstrates how the anamorphs most similar to one another are found on the plants systematically most like one another, and so on. Apart from demonstrating the possibility to differentiate between anamorphs that would otherwise be indistinguishable, the strict adherence of the conidial form of these obligate pathogens to the relative matrix is also shown. This method is useful both during epidemiological surveys, thus permitting suitable chemical intervention, as well as during systematic investigations on the pathogens under study. The standardization of the method, proposed for the first time in the field of powdery mildews, begins with the considered anamorphs being distributed in a sequence of six-figure codes (given the name "morphotype") expressing the following six groups of characteristics: type of appressoria, conidial shape, placement of conidia, absence or presence of fibrosin bodies, conidial outline, shape of conidiophore foot-cell. The biostatistical analysis described above will successively be applied to the anamorphs of each single morphotype. Introduction and Aims As is known, in the subfamily Erysiphoideae of the family Erysiphaceae, the teleomorphs of the various species can be fairly straightforwardly differentiated. The same cannot be said for the anamorphs (belonging to the genus Oidium Link), which, in the light of what is currently known, are often morphologically indistinguishable from each other. Furthermore, the habit of using the name of the teleomorph to indicate the anamorphs as well has more than ever made people refrain from carrying out a close study of the latter. Commenting on the lack of literature on the subject, Braun has recently published a few contributions (1980, 1982, 1982a). 194 In addition, since several powdery mildews are found, in nature, at the anamorphic state and infrequently, or at least, not always, develop into the perfect form, it has been considered worthwhile to make an accurate investigation of this conidial form. However, a deeper study of the individual anamorphs is not just an attempt to answer taxonomic queries; the possibility of differentiating one form from another also leads to a more thorough understanding of the epidemiology of the individual species and thus knowing the likelihood, or not, of the mildews on spontaneous plants infecting agricultural crops or spreading among them. The advantages to the planning of control methods are obvious. The aim of the present work is to reach such results, proposing a methodology based on biostatistical analysis. The analysis is carried out using both classical and new quantitative morphological parameters. The research on new parameters is even more necessary for the anamorphs whose descriptions (when based only on the parameters usually adopted) do not permit exact attribution to one species rather than another. Materials and Methods Five samples of "powdery mildews" were selected in the conidial state, according to the following criteria. As the first criterion, five anamorphs were chosen that, on the basis of the usual characteristics considered in the description of an Oidium, presented an identical "profile", 1.e., - type of appressoria: multi-lobed (type C of Boesewinkel, 1977); - conidial shape: cylindrical; - conidial placement: carried singly by the conidiophore (Pseudoidium-type anamorph: cfr. Blumer, 1967); - fibrosin bodies: inconspicuous; - conidial outline: curvilinear; - shape of the foot-cells of the conidiophore: cylindrical, flexuous at the base. Thus, according to the above characteristics, no differentiation exists between the five samples. As already mentioned, such a criterion was used to select five conidial forms of Erysiphaceae, which are cosmopolitan on the following host plants: 1&2) Erysiphe convolvuli DC. on two hosts of the same species: Calystegia sylvatica (Kit.) Griseb. - Fam. Convolvulaceae, but collected in areas far from one another (this pair is witness to the effectiveness of the proposed method); 3) The same E. convolvuli collected on a plant belonging to the family of 1 & 2), but of another genus - Convolvulus arvensis L.; 195 4) Another powdery mildew of a different genus than the preceding one - Microsphaera trifoli (Grev. ) U. Braun on a host, Trifolium pratense L., belonging again to a different family (Leguminosae); 5) Another powdery mildew of the same genus as above but belonging to another species - Microsphaera euonymi-japonici Viennot-Bourgin, on a host of a family that is very different from the two preceding ones - Euonymus japonicus L. fil. (fam. Celastraceae). The identification of the first four conidial forms was confirmed also with the respective teleomorphs. All five samples were collected on the same day and (obviously with the exception of one of the two Calystegia) in the same area, from plants found on the outskirts of Rome that had not undergone any chemical treatments. The measurements of the individual parameters of the anamorphs under study were Statistically processed using a Kontron MOP. Videoplan image analyzer connected via a telecamera to a Zeiss photomicroscope. The quantitative characteristics considered for each sample are relative to the following parameters: 1) area (projection surface) of the appressoria; 2) circumference (projection outline) of the appressoria; 3) maximum diameter (of the projection surface) of the appressoria; 4) shape factor of the circumference of the appressoria, based on the ratio area/circumference (used to define regular or irregular structures: for ellipses and irregular structures its value remains less than 1, while it reaches unity in the case of a circular structure); Dy) shape factor of the appressorium area, based on the minimum and maximum area/diameter ratio (as above: in the case of circular structures its value is 1, for irregular structures it remains less than unity); 6) real length of basal cell of the conidiophore; 7) area (projection surface) of conidia; 8) circumference (projection outline) of conidia; 9) shape factor of the conidial circumference (see above); 10) shape factor of the conidial area (see above); 11) conidial length; 12) maximum width of conidia; 13) ratio length/maximum width of conidia. From a research of the literature, the type of processing reported above has never been previously applied to Erysiphaceae, in general. 196 A broad outline for statistical processing has, at most, been inferred by Zar (1974) and Nordio (1977). The series of values obtained for the individual parameters (totalling 13 series for each sample) have been compared two by two (the series from each sample with the corresponding series of all the others), calculating Student’s t for each comparison. In order to visualize the ratio between the above series, the average linkage cluster analysis was applied to the symmetrical matrix of t, to allow the construction of the relative dendrogram. Standardization of the Method As already explained, in order to process the data collected, the preliminary condition was imposed that the anamorphs to be compared belong to the same "profile", based on six characteristics (obviously it is not necessary to differentiate between two or more anamorphs if they already belong to different profiles). To standardize this sequence of descriptive characteristics, for which the term "morphotype", as used by Ainsworth, 1971, is proposed, it is considered convenient to have a coding. The morphotype will be characterized by numbers chosen in the six figures, always in the same succession; one then proceeds with the following coding (Fig. according to Boesewinkel (loc. cit) and Braun (1987)). I-FIGURE Type of appressoria Indistinct: Boesewinkel, fig. 3E, Braun, Fig. 1a. Nipple-shaped: Boesewinkel, fig. 3A, Braun, Fig. 1b-g. Slightly or moderately lobed: Boesewinkel, Fig. 3D, Braun, fig. 1h. Multi-lobed: Boesewinkel, fig. 3C, Braun, Fig. 11. II-FIGURE Conidial shape Sashes As is known, conidia of the genus Oidium can assume very different shapes, which basically lead back to three: 1. _Ellisoid or ovoid: Braun, Figs. 3A and 3B. 2. Cylindric (when the two largest sides are parallel for a long tract: Braun, Fig. 3C). 3. Botuliform or doliform (with nonparallel sides and with the two ends prominent: Braun, Fig. 3D). III-FIGURE Placement of the conidia — Formed singly. 2. Produced in chains. IV-FIGURE Absence or presence of fibrosin bodies 1. Present - well-developed fibrosin bodies. 2. Absent - inconspicuous fibrosin bodies. 197 V-FIGURE Conidial outline 1. Curvilinear. 2. Angular, as for the case of Sawadaea. VI-FIGURE Shape of conidiophore foot-cell This is also very variable but lead back to seven basic morphologies: Gradually decreasing in diameter from base to top: Boesewinkel, Fig. SE. Cylindrical (straight, cylindrical foot-cell: Boesewinkel, Fig. 5A). Gradually decreasing in diameter from top to base: Boesewinkel, Fig. 5D. Swollen at the base. Moniliform (swollen foot-cell with a conspicuously enlarged basal area: Boesewinkel, Fig. 5B; typical of anamorphs of Blumeria graminis (DC.) Speer). 6. Flexuous. 7. Conidiophore curved in the basal half: Braun, Fig. 65c. A cat ad dae As an example of what is listed above, the morphotype under study is: 4 2 1 2 1 6, and so on. Thus, in order to recognize and differentiate these anamorphs, one first determines the respective morphotypes, grouping together those that are identical. Then, the anamorphs of each morphotype are biostatistically analyzed as described above, with the aim of further differentiating one from the other. Results The dendrogram constructed with the processed data is given in Fig. 1. It shows how the measurements that are most similar to one another belong to the two Enrysiphe convolvuli on the two samples of Calystegia. Then, with increasing order of t, it can be seen that this pair is close to the same E. convolvuli collected on the other convolvulacea, but well differentiated from it. The measurements on the powdery mildews of the other two _ botanical families are considerably dissimilar from the preceding measurements. Finally, by calculating, for each of the three families under study, the overall average of their Student’s t, we can note how a closer similarity exists between the leguminosa and the three convolvulaceae than between the latter and the celastracea: this is in agreement with the similarity between the botanical families as reported by Hutchinson (1969). Using the methodology applied here, we can obtain the differentiation of the anamorphs under study, which would be impossible to do by observation only. - Moreover, the biostatistical analysis used on the individual samples leads to a deeper insight into the morphological characterization of the various anamorphs: this information constitutes a valuable databank that can also be used for systematic analyses of the genus Oidium. 198 Euonymus japonicus Convolvulus arvensis Trifolium pratense S = 3s > aS} Ry & S —| Calystegia sylvatica 2.0 3.0 4.0 Se 6.0 7.0 8.0 Fig. 1 Cluster analysis dendrogram for single linkage applied to the symmetrical matrix of Student’s t. 199 Discussion and Conclusions The methodology proposed in this work has enabled to distinguish the positions of reciprocal similarity between the five anamorphs examined. The Statistical analysis has outlined both a strict adherence of the conidial shape of the pathogen to its host, as well as its precise relationship to the relative teleomorph. The results confirm the well-known determining influence of the host plants, a true and proper matrix, on at least the anamorphic morphology of the fungus; thus, it can be utilized to investigate the natural distribution of such obligate pathogens. In the meanwhile the method provides valuable information to enable a deeper insight into the systematics of Erisifaceae. It can be concluded that, at least in the case of anamorphs, the method presented appears to justify the authors who have proposed a classification of these pathogens that strictly adheres to the taxonomic position in the botanic system of the respective host plants. The author wishes to thank Prof. A. Rambelli for kindly reviewing the manuscript. References Ainsworth, G.C. (1971). Aisnworth & Bisby’s Dictionary of the Fungi. Commonwealth Mycological Institute, Kew, Surrey, 663 pp. Blumer, S. (1967). Echte Mehltaupilse (Erysiphaceae). Gustave Fischer Verlga, Jena, 436, pp. Boesewinkel, H.J. (1977). Identification of Enrysiphaceae by _ conidial characteristics. Revue Mycol., 41: 493-507. Braun, U. (1980). Morphological studies in the genus Oidium. Flora, 170: 77-90. Braun, U. (1982). Morphological studies in the genus Oidium (II). Zbl. Mikrobiol., 137: 138-152. Braun, U. (1982a). Morphological studies in the genus Oidium (III). Zbl. Mikrobiol., 137: 314-324. Braun, U. (1987). A monograph of the Erysiphales (powdery mildews). Nova Hedwigia, beiheft 89. J. Cramer, Berlin-Stuttgart, 1-700. Hutchinson, J. (1969). Evolution and phylogeny of flowering plants. Academic Press, London and New York, 717 pp. Nordio, V. (1977). Festuca alpestris R. et S. nelle prealpi bellunesi. Giorn. Bot. Lialoeh1129123-128, Zar, J.H. (1974). Biostatistical analysis. Prentice-Hall Inc., Englewood Cliffs, N.J. USA, 620 pp. — — SS ee i = = .. —— yi Ah st — f ; i : a ai. As at ay Bit} Avnet Ay i Wid 4 i a é ; Oe, y i Pi Ne m ne 7 t q Berta 1 Het " es \ vn ps ise » is ’ h bd % i im ie hy bf 1 N “ih + shaped P meet Wh i ei A a Mm i Ae eye Lt oh ihn Pay Ay (het F pi MR LBD i 7 Bee Phi, LFA, 1 Baie: ihe Heat ‘iui hege aaah Site | ee in TUNA La a PUI WA aes | ne Me uh oes a uk wy ie : a EL liee ae ae Shey hy uroyt are eae wife nyt . : ; hy " wis rhe til: Mik a ay, hs lhe ) ii a casio i | | ea ea Aa : sista A} i ' yi ’ ey + been: d aT Le al ‘fe ee i Are ites Rae ae Abie a tal ub \ t 45 Pg let Pies ’ i i ‘ i ) by re 4 1. " 7 Denes teas, Peg 8) Se : volt Se Pye Wed A has iniiy tit) oak AY a) 7 ae oe re (i'n ct rah si i pi : Rit ‘ ut he ‘ ae j uy { a | iy th; Toa hb 1, Sip: i aur , r = SAt j . 7 : i 14% ey 7 al "in inh y > nt a Py wae a Ai) rat tat i ‘ \ i Mig aa q i4 ie Ae | Lo I } i i) Oh 7 Ceres sit wet a AF oat t . i i Lan a L A oa Ny a a AAs ne apse vai i si a nit rs ROR ssl ‘ ae r bee f } te “ “ae eo p A, ty Be eh hg Ltt a Na ebay rg) abe iy) i, SR Lech law On a) 2 NET Wao tint 280 ey ; * i i { a av vi te | ne ¥, “t \y ae a! ay i Ps Ni fA bet Pe Pa i: j ' Nae Tt Rab ee Canosa ? 4 waht u j f i} ; P bat So ‘4 mi ‘ hs 1 Bo) i) aN i “sf MH A Hint A Mh laa’ Dey Bs Pa | ayes wali eee ‘ r) wy Wy, | ed . , min ante Mi, “ph vi Bes aia , Li se basi § h, ; Lhe Oe Pe ay ms Sey ny ag Pa Hi) ’ ih hye: anol as Vinee wa) aa Pte cat $a ty tlt ha me aa ai i A deeattntlgn neh ial bn d te ae. F 1 Lan uh H Ayat y hy lf LAT 4 ice 1 | od ath? i hp ear a : fo) Y ata \ a a i en i 1h eae ‘ apd yi pa ansaileny Ye luna th tee y ee Avene | WO et eal ey ne He, oud ity ee herent Lil Hy ‘et “Aull, 4 hae py, . Ht es, ia iy eet We a vey vr py ay eer | Vik geal : ane ¥ i pees a ia a Ai : tly ‘th wee Reh Pek a up a 3 cae aay 4 te “iy Eh sneetihty sate ti ise ‘ ee pth 12 by Rae ’ , MeL oe MYCOTAXON Volume XLVII, pp. 201-217 April-June 1993 HYPOGEOUS MYCORRHIZAL FUNGI OF SPAIN Isabel F. Alvarez’, Javier Parladé', James M. Trappe’, and Michael A. Castellano® "Departamento de Patologia Vegetal. Institut de Recerca i Tecnologia Agroalimentaries (IRTA). Centre de Cabrils. 08348 Cabrils, Barcelona, Spain. 7Department of Forest Science and Botany-Plant Pathology. Oregon State University, Corvallis, Oregon 97331. *United States Department of Agriculture, Forest Service, Pacific Northwest Research Station. Forestry Sciences Laboratory, Corvallis, Oregon 97331. ABSTRACT Although scientific study of hypogeous fungi in Spain began early in this century, our collections over five years (specially in the northern area of the Iberian Peninsula) produced 18 new species records, including two new species and one new genus. This paper includes first reports in Spain for E/aphomyces muricatus, Glomus macrocarpum, Hymenogaster sp., Hymenogaster albus, H. muticus, H. niveus, H. tener, Hysterangium clathroides, H. gardneri, Loculotuber gennadi, Rhizopogon subareolatus, R. ventricisporus, Tuber gennadii, T. levissimum, T. maculatum, T. malencgonii and T. multimaculatum. Information on previously reported species is expanded to other geographical areas. The updated relation of Spanish hypogeous fungi and its ecological significance in the forests of northern Spain are discussed. KEY WORDS: Ascomycotina, Basidiomycotina, truffle, ectomycorrhiza, Elaphomyces, Glomus, Hymenogaster, Hysterangium, Loculotuber, Rhizopogon, Tuber. 202 INTRODUCTION The study of hypogeous fungi in Spain was initiated by Lazaro Ibiza in 1908. The first published compilation of data available on hypogeous fungi in Spain reported 34 species in 12 genera (Calonge et al. 1977). Subsequent additions increased the number of reported fungi considerably (Calonge 1982, Calonge et al. 1985b, Calonge 1990). Most hypogeous fungi are desirable ectomycorrhizal symbionts of tree seedlings because they increase water and nutrient uptake (Bowen 1973, Luoma et al. 1991). Additional interest for their use in reforestation practices stems from their capability to reduce soil drought stress in tree seedlings (Parke et al. 1983). To use hypogeous fungi for increasing plantation success, they must be collected in the field, identified, isolated or kept for spore inoculum, and successfully infect seedlings in the nursery (Trappe 1977, Castellano et al. 1985, Castellano and Molina 1989). Many hypogeous ectomycorrhizal fungi show plant- fungus specificity (Molina and Trappe 1982). Available information on hypogeous fungi in humid northern Spain has been scarce. This area extends from Galicia to Catalonia, a total of ca. 14 million ha. Presently, 4.4 million ha are covered with conifer, broadleaf and mixed forests. An additional 3.3 million ha will be reforested in the future (Anonymous 1989). The objective of this study was to collect and identify hypogeous mycorrhizal fungi occurring with the major forest species of northern Spain for possible later use in either conventional reforestation, or truffle production programs. MATERIALS AND METHODS Fungi were collected at least twice a year from fall through spring of 1985- 1989. The Cantabric zone (from Galicia to the west part of the Pyrenees range) was sampled from sea level to 1200 m elevation in a range of forest associations. At the lower elevations (0 - ca 400 m) the stands consisted of either pure Pinus pinaster Ait., Pinus radiata D. Don or a mixture of the two pines with Castanea sativa Mill. At the middle elevations (300 - 700 m) similar mixed forests were examined as also were plantations of Eucalyptus globulus Lab. on the coastal slopes. Inland, in the Cantabric range, the tree species were Acer pseudoplatanus L., Corylus avellana L., Fraxinus excelsior L., Ilex aquifolium L., Populus spp., and Quercus petraea (Matts.) Liebl. At the higher elevation (1000 - 1200 m) in Asturias (Picos de Europa) the forests were composed of Fagus sy/vaticaL. and Quercus pyrenaica Willd. In Catalonia (Montseny Range) the mixed forests were located at 1000-1200 m elevation and composed of Abjes alba Mill, Fagus sylvatica L., Quercus ilex L., Quercus pubescens Willd. and plantations of Picea abies (L.) Karst. and Pseudotsuga menziesii (Mirb.) Franco. Other areas were occasionally searched for hypogeous fungi. Among them 203 were the Beceite range in southern Catalonia (1200 m elevation in a native Pinus sylvestris L. forest) and plantations of introduced pines at the lower elevations of the Montseny Range. Hypogeous fungi were sought by raking the soil and litter. Care was taken to replace the litter afterwards. Field data were taken on fresh sporocarp features, overstory species composition, associated ectomycorrhizal hosts, and soil characteristics. A few specimens were found with a trained dog. Some collections provided by commercial truffle hunters lacked field data. Because of their mycological interest, available collections of hypogeous fungi from other areas of Spain recently described have been included in this review (Alvarez et al. 1992). Hypogeous fungi from the Canary Islands were compiled previously by Fogel (1980) and are not included here. Microscopic observations were made by sectioning sporocarp tissue with a razor blade and mounting the material in water, KOH, cotton blue or Melzer’s reagent. Sporocarps were dried at 30°C for herbarium deposit. Isolations were made from fresh sporocarps on MMN agar slants (Molina and Palmer 1982). Herbarium specimens were deposited in the Instituto Antonio José Cavanilles, Real Jardin Botanico (MA), Madrid, Spain and in the Oregon State University Herbarium (OSC), Corvallis, Oregon. Other herbaria cited are the National Fungus Collections, Washington D.C. (BPI), The Royal Botanic Gardens, Kew (K), and the Istituto ed Orto Botanico, Torino (TO). RESULTS A total of 36 species of hypogeous fungi were collected and identified. Eighteen are new records for Spain, including two new species. The following list attempts to update the known species of hypogeous fungi that occur in Spain. The list includes data on collection site characteristics and associated mycorrhizal hosts when available, as well as the positive results of isolation attempts. Newly reported species and new localities (provinces) for previously reported species are written in bold characters. Collection numbers, place of deposit, and previous references are given in parentheses. An asterisk after the locality indicates that it is an Autonomous Community (composed of one or more provinces) and the exact locality is not known. The symbol (+) after the species name indicates that it was isolated successfully. The identification of the species reported by other authors has not been verified except when indicated. Balsamia vulgaris Vitt. BARCELONA, March 1988; 30-40 yr.- old C. avellana plantation on dry, stony pH 7.8 soil (MA-28295, OSC), (Calonge et al. 1985b). CORDOBA, (Moreno et al. 1991). Associated mycorrhizal hosts: Cistus albidus L. and C. avellana. 204 Choiromyces magnusii (Matt.) Paol. EXTREMADURA%, (Calonge et al. 1985a). Associated mycorrhizal hosts: unknown. Chofiromyces venosus (Fries) Th. Fries (= C. meandriformis Vitt., Tuber sinuosum Laz.) ALAVA, (Calonge et al. 1977). BADAJOZ, (Lazaro Ibiza 1908, Moreno et al. 1991). CACERES, (Moreno et al. 1991). Associated mycorrhizal hosts: Cistus /ladanifer L., Fagus sp., and Quercus sp. NOTE: The type collection of Tuber sinuosum Laz. is lost (Calonge et al. 1985a). Lazaro’s illustrations and descriptions, however, are readily compatible with C. venosus. Delastria rosea Tul. BARCELONA, (Calonge et al. 1985b). Associated mycorrhizal hosts: mixed forest, specific host unknown. Elaphomyces anthracinus Vitt. CORDOBA, (Moreno et al. 1991). Associated mycorrhizal hosts: Q. i/ex. Elaphomyces granulatus Fries ASTURIAS, September 1986; in a Q. petraea and Populus sp. forest at 760 m. elevation, soil pH:4 (MA-28299, OSC). GIRONA, (Unamuno 1941). PAIS VASCO%*, (Calonge et al. 1977). PONTEVEDRA, November 1987; under P. pinaster on a controlled-burned, flat area. Soil pH: 4.7 (MA-28297, MA-28298, OSC). SEGOVIA, (Calonge et al. 1977). SORIA, November 1987; under P. sy/vestris at 1200 m. elevation. Soil pH: 5.2 (MA-28296, OSC). Associated mycorrhizal hosts: F. sylvatica, P. sylvestris, and Quercus sp. Elaphomyces muricatus Fries SANTANDER, May 1986; under Q. pyrenaica at 1200 m. elevation (MA-28300, OSC). Associated mycorrhizal hosts: Q. pyrenaica. Elaphomyces personii Vitt. GIRONA, (Codina and Font-Quer 1931). Associated mycorrhizal hosts: C. sativa. Elaphomyces trappei Galan & Moreno BADAJOZ, (Moreno et al. 1991). Associated mycorrhizal hosts: C. /adanifer. Gautieria mexicana (Fisch.) Zeller & Dodge ALAVA, (Calonge et al. 1985b). Associated mycorrhizal hosts: P. radiata. Gautieria morchelliformis Vitt. BARCELONA, (Aymerich and Llimona 1986). CORDOBA, (Moreno et al. 1991). CUENCA, (Calonge et al. 1985a). LERIDA, (Calonge et al. 1977). Associated mycorrhizal hosts: Abies sp., Pinus nigra Arnold, and Q. ilex. Genea klotzschii Berk. & Broome BARCELONA, February, March 1988; collected in a 30-40 year-old C. avel//ana plantation on stony, pH 7.8 soil (MA-28301, MA-28302, OSC), (Calonge et al. 1985b). VALLADOLID, (Calonge et al. 1985b). Associated mycorrhizal hosts: C. ave//ana and Pinus pinea L. 205 Genea verrucosa Vitt. CORDOBA, (Moreno et al. 1991). TOLEDO, (Arroyo et al. 1989). Associated mycorrhizal hosts: C. albidus and Q. ilex. Geopora arenicola (Lév.) Kers (= Sepultaria arenicola (Lév.) Cooke, S. arenosa (Fuck.) Cooke)). GIRONA, (Heim et al. 1934). GRANADA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986). MURCIA, (Honrubia et al. 1983). VALLADOLID, (Moreno et al. 1991). Associated mycorrhizal hosts: Pinus spp., Pinus halepensis Mill. and P. pinaster. Geopora cervina (Vel.) Schum. GRANADA, (Moreno et al. 1986). Associated mycorrhizal hosts: Quercus faginea Lamk. and Q. ilex. Geopora cooperi Harkn. ALAVA, (Mendaza and Diaz 1987). GRANADA, (Ortega et al. 1981). GUADALAJARA, (Moreno et al. 1986). MADRID, (Moreno et al. 1991). MURCIA, (Honrubia et al. 1990). Associated mycorrhizal hosts: Pinus spp. and Q. ilex. Geopora foliacea (Schaeff.) Anmad (= Sepultaria foliacea (Schaeff.) Boud.) ALBACETE, (Honrubia et al. 1983). BARCELONA, (Heim et al. 1934). GRANADA, (Ortega et al. 1981). GUADALAJARA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986). Associated mycorrhizal hosts: P. halepensis and Populus alba L. Geopora nicaensis (Boud.) Torre ALMERIA, (Calonge and Oria de Rueda 1988). BARCELONA, (Losa Quintana 1974, Moreno et al. 1986). GRANADA, (Moreno et al. 1986). GUADALAJARA, (Moreno et al. 1991). MADRID, (Moreno et al. 1986). Associated mycorrhizal hosts: conifers and Q. i/ex. Geopora sumneriana (Cook.) Torre GRANADA, (Moreno et al. 1986). MADRID, (Moreno et al. 1986). Associated mycorrhizal hosts: Cedrus sp. Geopora tenuis (Fuck.) Gal.& Ort. GRANADA, (Galan and Ortega 1982). Associated mycorrhizal hosts: unknown. Glomus macrocarpum Tul. & Tul. ASTURIAS, September 1986; in a mixed forest of A. pseudoplatanus, F. excelsior, and Q. petraea at 700 m. elevation. Soil pH: 4 (MA-28303, OSC). Associated mycorrhizal hosts: unknown. Glomus microcarpum Tul. & Tul. (= Endogone microcarpa (Tul. & Tul.) Tul.& Tul. BARCELONA, (Heim et al. 1934). Associated mycorrhizal hosts: unknown. Hydnangium aurantiacum Heim & Mal., (= H. carotaecolor Berk.) BARCELONA, (Codina and Font-Quer 1931, Heim et al. 1934). Associated mycorrhizal hosts: P. halepensis and Q. ilex. 206 Hydnangium carneum Wallr. 4 BADAJOZ, (Moreno et al. 1991). LA CORUNA, (Jeppson 1987). Associated mycorrhizal hosts: Eucalyptus spp. Hydnangium pila Pat. GIRONA, (Heim et al. 1934). Associated mycorrhizal hosts: Quercus suber L. NOTE: The generic disposition of this species is disputable by modern concepts. Svrcek’s designation of it as Octaviania pila (Pat.) Svrc. is nomenclaturally unacceptable by International Code of Botanical Nomenclature (Svrcek 1958). Hydnotrya tulasnei (Berk.) Berk. & Broome BARCELONA, (Calonge et al. 1985a). VALLADOLID, (Calonge 1982). VIZCAYA, (Mendaza and Diaz 1987). Associated mycorrhizal hosts: C. avellana, F. sylvatica, Pinus spp. and Quercus spp. Hymenogaster sp. No. 1 : GIRONA, November, December 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil (MA-28314, MA-28315, MA-28316, MA-28317, OSC). PONTEVEDRA, November 1987; in a 15-yr.-old plantation of P. menziesii on flat ground with no understory (MA-28313, OSC). Associated mycorrhizal hosts: P. menziesii. Hymenogaster sp. No. 2 GIRONA, October 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil (MA-28318, OSC). Associated mycorrhizal hosts: P. menziesii. Hymenogaster aibus Berk. & Br. ASTURIAS, October 1986; under E. globulus at 500 m. elevation (MA-28304, OSC). Associated mycorrhizal hosts: E. globulus. Hymenogaster bulliardi Vitt. TOLEDO, (Calonge et al. 1985b). Associated mycorrhizal hosts: unknown. Hymenogaster calosporus Tul. & Tul. BARCELONA, (Heim et al. 1934). Associated mycorrhizal hosts: C. avellana, Q. ilex and Sorbus aucuparia L. Hymenogaster cerebellum Cavara CACERES, (Moreno et al. 1991). Associated mycorrhizal hosts: Q. pyrenaica. Hymenogaster citrinus Vitt. ASTURIAS, May 1986; under Quercus robur L. at 160 m elevation (MA-28306, OSC). BARCELONA, (Heim et al. 1934). CORDOBA, (Moreno et al. 1991). GIRONA, October 1987; under F. sy/vatica at 1100 m elevation (MA-28305, OSC). Associated mycorrhizal hosts: F. sylvatica, Q. ilex and Q. robur. Hymenogaster muticus Berk. & Br. GIRONA, December 1987; under P. sy/vestris (MA-28307, OSC). Associated mycorrhizal hosts: P. sy/vestris. 207 Hymenogaster niveus Vitt. SANTANDER, May 1986; under F. sy/vatica at 1200 m elevation (MA-28308, OSC). Associated mycorrhizal hosts: F. sylvatica. Hymenogaster olivaceus Vitt. BARCELONA, (Heim et al. 1934). Associated mycorrhizal hosts: C. avellana and Q. ilex. Hymenogaster populetorum Tul. & Tul. BARCELONA, (Calonge et al. 1985b). Associated mycorrhizal hosts: unknown. Hymenogaster remyi Zeller & Dodge in Dodge & Zeller GUADALAJARA, (Calonge 1982). Associated mycorrhizal hosts: Pinus sp. Hymenogaster tener Berk. & Br. ASTURIAS, May 1986; under mixed forest of E. giobulus and P. radiata at475m elevation, October 1986; under F. sy/vatica at 600 m. elevation (MA-28309, MA-28310, OSC). Associated mycorrhizal hosts: F. sylvatica. Hymenogaster vulgaris Tul.& Tul. in Berk. & Br. BARCELONA, (Heim et al. 1934, Calonge et al. 1985a). GIRONA, November, December 1989; under P. menziesii at 1100 m. elevation in pH 5.5 soil (MA-28311, MA-28312, OSC). Associated mycorrhizal hosts: Pinus sp. and P. menziesii. Hysterangium clathroides Vitt. ASTURIAS, May 1986; in a 15-years-old P. radiata plantation, at 270 m. elevation. (MA- 28319, OSC). Associated mycorrhizal hosts: P. radiata. Hysterangium gardneri Fischer ASTURIAS, May 1986; in a mixed stand of C. sativa and E. globulus at 575 m. elevation (MA-28320, OSC). Associated mycorrhizal hosts: E. globulus. Hysterangium rickenii Soehn. GRANADA, (Moreno et al. 1986). Associated mycorrhizal hosts: Q. faginea and Q. ilex. Hysterangium separabile Zeller CUENCA, (Calonge et al. 1985a). MADRID, (Calonge et al. 1977). Associated mycorrhizal hosts: P. nigra and P. sylvestris. Hysterangium stoloniferum Tul. var. rubescens (Quél.) Zeller & Dodge. CACERES, (Moreno et al. 1991). CORDOBA, (Moreno et al. 1991). Associated mycorrhizal hosts: C. sativa and Q. ilex. Hysterangium thwaitesii Berk. & Broome ALAVA, (Calonge et al. 1977). Associated mycorrhizal hosts: unknown. 208 Labyrinthomyces donkii Mal. ALICANTE, (Honrubia 1984). CADIZ, (Calonge 1982, 1983). MURCIA, (Honrubia 1984). Associated mycorrhizal hosts: E. g/obulus. NOTE: This genus has been recently revised by Trappe et al. (1992). Leucogaster floccosus Hesse MURCIA, (Honrubia and Llimona 1981). Associated mycorrhizal hosts: unknown. Loculotuber gennadi Trappe, Parladé & Alvarez CACERES, May 1987; (MA-26771, OSC). (Alvarez et al. 1992). Associated mycorrhizal hosts: unknown. Martellia mediterranea Moreno, Galan & Montecchi CACERES, (Moreno et al. 1991). Associated mycorrhizal hosts: C. /adanifer and Q. ilex. Melanogaster ambiguus (Vitt.) Tul. (+). BARCELONA, (Calonge 1982). GIRONA, November 1989 and June 1990 under P. menziesii at 1100 m. elevation in pH 5 soil (MA-28321, MA-28322, MA-28323, MA-28324, MA-28325, OSC), (Martin 1988). MADRID, (Moreno et al. 1986). Associated mycorrhizal hosts: F. sy/vatica, P. menziesii, Q. ilex, and Q. suber. Melanogaster broomeianus Berk. in Tul. & Tul ALBACETE, (Calonge and Demoulin 1975). BARCELONA, (Calonge et al. 1985b, Llistosella and Aguasca 1986). GIRONA, (Maire 1937, Singer 1947). MADRID, (Calonge et al. 1985a). Associated mycorrhizal hosts: P. sy/vestris and P. pinea. Melanogaster variegatus (Vitt.) Tul. & Tul. ALBACETE, (Calonge 1973). CACERES, (Moreno et al. 1991). CORDOBA, (Moreno et al. 1991). GUIPUZCOA, (Lazaro Ibiza 1912). MURCIA, (Honrubia et al. 1990). VALENCIA, (Lazaro Ibiza 1912). Associated mycorrhizal hosts: Pinus spp. and Q. suber. Octaviania asterosperma Vitt. CACERES, (Moreno et al. 1991). Picoa juniperi Vitt. MADRID, (Calonge et al. 1985a). ZARAGOZA, (Calonge 1982). Associated mycorrhizal hosts: unknown. Rhizopogon /uteolus Fries & Nordh. (+) ALAVA, December 1989; under P. sy/vestris (MA-28326, OSC). ASTURIAS, October 1986: under P. radiata at 100 m. elevation (MA-28327). BARCELONA, (Maire et al 1933, Heim et al. 1934, Llimona 1984, Martin 1988). GIRONA, (Maire et al. 1933, Llimona 1983, Martin 1988). LERIDA, (Maire 1937). MADRID, (Calonge and Zugaza 1973, Calonge and Demoulin 1975). PONTEVEDRA, November 1985, 1986, and 1987; under P. pinaster in compacted, sandy pH 4.7-6 soils and sometimes abundant in controlled-burned areas (MA-28328 MA- 28329 MA-28330, MA-28331, MA-28332, MA-28333, MA-28334, MA-28335, MA-28336, OSC). TARRAGONA, (Maire 1937). Associated mycorrhizal hosts: P. nigra, P. pinaster, P. pinea, P. radiata and P. sylvestris. 209 Rhizopogon roseolus (Corda ex Sturm) Th. Fries (= R. aestivus (Tul. & Tul.) Tul. & Tul., R. rubescens Tul & Tul.) (+) ALBACETE, (Honrubia et al. 1982). ALICANTE, (Honrubia et al. 1982). ALMERIA, (Honrubia et al. 1982). ASTURIAS, December 1985, May and October 1986; under P. radiata plantations at 100-350 m. elevation in pH 5.1 soil (MA-28337 and MA-28338, OSC). BALEARES, (Malengon and Bertault 1972). BARCELONA, (Codina and Font-Quer 1931, Maire et al. 1933, Heim et al. 1934, Maublanc 1936, Singer 1947, Calonge and Demoulin 1975, Tabares and Rocabruna 1987, Martin 1988). GIRONA, (Maire et al. 1933, Maire 1937, Font- Quer 1954, Calonge and Demoulin 1975, Llimona 1983, Martin 1988). JAEN, (Honrubia et al. 1982). LEON, November 1989; under P. sy/vestris in sandy soil (MA-28342, MA-28343, OSC). LERIDA, (Calonge and Demoulin 1975, Llimona 1983, Martin 1988). MADRID, (Calonge 1973, Calonge and Zugaza 1973, Calonge and Demoulin 1975). MALAGA, (Calonge and Demoulin 1975). MURCIA, (Honrubia et al. 1982). PONTEVEDRA, November 1986; under P. pinaster (MA-28339, OSC). TARRAGONA, September 1987, August 1989; under P. sylvestris at 1100 m. elevation in pH 7.4 soil (MA-28340, MA-28341, OSC). VALLADOLID, (Calonge and Demoulin 1975). Associated mycorrhizal hosts: P. halepensis, Pinus mugo Turra., P. nigra, P. pinaster, P. pinea, P. radiata and P. sylvestris. Rhizopogon subareolatus Smith (+) GIRONA, November, December 1987 and 1988, abundant under P. menziesii plantations or under Abies sp. with P. menziesii present on slopes at 1100 m. altitude in pH 4.7-6 soil (MA-28344, MA-28345, MA-28346, MA-28347, MA-28348, MA-28349, MA-28350, and MA- 28351, OSC). Associated mycorrhizal hosts: P. menziesii. NOTE: This species has not previously been reported in continental Europe. It was reported in the Madeira islands where it was probably introduced with P. menziesii (Fogel 1980). Rhizopogon ventricisporus Smith (+) TARRAGONA, September 1987; under P. sy/vestris at 1100 m. elevation in pH 7.4 soil (MA- 28352, OSC). Associated mycorrhizal hosts: P. sylvestris. Rhizopogon vulgaris (Vitt.) M. Lange (= R. provincialis Tul. & Tul.) (+) ALBACETE, (Honrubia et al. 1982). ASTURIAS, May 1986 and October 1986; under P. radiata at 100-350 m elevation in pH 5.1 soil (MA-28353, MA-28354, MA-28355 and MA- 28356, OSC). BARCELONA, (Codina and Font-Quer 1931, Martin 1988). GIRONA, (Heim et al. 1934, Malencon and Bertault 1971). MADRID, (Calonge and Demoulin 1975). MALAGA, (Calonge and Demoulin 1975). PONTEVEDRA, November 1985, 1986 and 1987; under P. pinaster on coast in pH 4.7 soil in controlled-burned areas and under P. menziesii in 15 year-old plantations (MA-28357, MA-28358, MA-28359, MA-28360 and MA-28361, OSC). SORIA, November 1987; under P. sy/vestris at 1200 m elevation in pH 5.2 soil (MA-28362 and MA-28363, OSC). Associated mycorrhizal hosts: P. halepensis, P. nigra, P. pinaster, P. radiata, P. sylvestris and P. menziesii. Terfezia arenaria (Mor.) Trappe (= T. /eonis Tul.) BADAJOZ, (Calonge et al. 1977, Calonge et al. 1985a). CACERES, May 1987 (MA-28364 and MA-28365, OSC), (Moreno et al. 1986). HUELVA, (Calonge et al. 1985a). TOLEDO, (Calonge et al. 1985b). Associated mycorrhizal hosts: C. Jadanifer, Cistus monspeliensisL., Cistus salvifolius L. and Helianthemum guttatum Mill. 210 Terfezia berberiodora (Lesp. ex Tul. & Tul.) Tul. & Tul. LA CORUNA, (Castro and Freire 1982). Associated mycorrhizal hosts: unknown. Terfezia claveryi Chat. BURGOS, (Calonge et al. 1985b). GRANADA, (Calonge et al. 1985b, Moreno et al. 1986). MADRID, (Calonge et al. 1985b). Associated mycorrhizal hosts: Helianthemum sSsalicifolium Pers. Terfezia hispanica Laz. ALBACETE, (Lazaro Ibiza 1908). BADAJOZ, (Lazaro Ibiza 1908). CACERES, (Lazaro Ibiza 1908). CIUDAD REAL, (Lazaro Ibiza 1908). CORDOBA, (Lazaro Ibiza 1908). GRANADA, (Lazaro Ibiza 1908). GUADALAJARA, (Lazaro Ibiza 1908). MADRID, (Lazaro Ibiza 1908). SALAMANCA, (Lazaro Ibiza 1908). TOLEDO, (Lazaro Ibiza 1908). ZARAGOZA, (Lazaro Ibiza 1908). Associated mycorrhizal hosts: Helianthemum paniculatum Durm., H. salicifolium, Helianthemum vulgaris Gaertn., and Tuberaria variabilis Wk. NOTE: T. hispanica may be an aberrant collection of 7. o/biensis; the type collection is lost (Calonge et al. 1985a). Because of its distinctive vinaceous color as illustrated by Lazaro Ibiza (1908), however, we prefer to keep it separate from 7. o/biensis pending new collections. Terfezia olbiensis (Tul. & Tul.) Tul. & Tul. (= T. feptoderma (Tul. & Tul.) Tul. & Tul., 7. cadevalli Font-Quer, Tuber lutescens Laz., T. pallidumLaz.) ALAVA, (Calonge et al. 1977). ALBACETE, (Lazaro Ibiza 1908). AVILA, (Lazaro Ibiza 1908). BADAJOZ, (Calonge et a!. 1977, Calonge et al. 1985b). BARCELONA, (Codina and Font-Quer 1931). BURGOS, (Calonge et al. 1985b). CACERES, May 1987 (MA-28366 and MA-28367, OSC), (Moreno et al. 1986). CIUDAD REAL, (Lazaro Ibiza 1908). MADRID, (Lazaro Ibiza 1908, Calonge 1982, Calonge et al. 1985b). SALAMANCA, (Lazaro !biza 1908). SEGOVIA, (Calonge et al. 1985b). TOLEDO, (Calonge et al. 1985b). VALLADOLID, (Lazaro Ibiza 1908). ZAMORA, (Lazaro ibiza 1908). Associated mycorrhizal hosts: H. guttatum, H. paniculatum, Q. ilex, and T. variabilis. NOTE : We have found many sporocarps intermediate in morphology between those reported for 7. olbiensis and T. leptoderma in the numerous collections we have examined. Most differences appear to be a matter of developmental stage, as indicated by Fogel (1980). Hence, we regard 7. /eptoderma as a synonym of 7. olbiensis (we choose T. o/biensis as the epithet to retain, because its type collection contains more mature material than does that of 7. /Jeptoderma). Although the type collections of Terfezia cadevalli Font-Quer, Tuber lutescens Laz. and T. pallidum Laz. are lost (Calonge et al., 1985a), the descriptions of each present no reason to regard them other than as somewhat inmature to mature stages of 7. olbiensis. Trappea darkeri (Zeller) Castellano GUADALAJARA, (Calonge 1982). Associated mycorrhizal hosts: Pinus sp. NOTE: This collection was initially reported as Hysterangium membranaceum Vitt. (Castellano 1990). Tuber aestivum Vitt. BARCELONA, March 1987 (MA-28368, OSC), (Unamuno 1941, Calonge et al. 1985a). CORDOBA, (Moreno et al. 1991). GIRONA, (Codina and Font-Quer 1931, Unamuno 1941). LERIDA, March 1987; under Pinus sp. (MA-28369, OSC). TERUEL, March 1987; under Pinus sp. (MA-28370, OSC). Associated mycorrhizal hosts: C. avellana, Pinus sp. and Q. ilex. 211 Tuber asa Tul. & Tul. BADAJOZ, (Moreno et al. 1991). CACERES, May 1987, (MA-28371 and MA-28372, OSC), (Moreno et al. 1991). Associated mycorrhizal hosts: C. ladanifer. Tuber borchii Vitt. MURCIA, (Honrubia and Llimona 1981). Associated mycorrhizal hosts: P. halepensis. Tuber brumale Vitt. BARCELONA, March 1988; under C. avellana in 30-40 year-old plantations in pH 7.8 soil (MA-28373, OSC), (Codina and Font-Quer 1931), and three collections at TO. CASTELLON, (Calonge et al. 1977). GIRONA, (Lazaro Ibiza 1912, Codina and Font-Quer 1931, Heim et al. 1934, Unamuno 1941). GUADALAJARA, (Calonge et al. 1977). MADRID, one collection at TO. TARRAGONA, March 1987 (MA-28374, OSC). Associated mycorrhizal hosts: C. ave/l/ana and Quercus sp. Tuber excavatum Vitt. BARCELONA, (Calonge et al. 1985a, b). CORDOBA, (Moreno et al. 1991). GUADALAJARA, December 1987, under Populus sp. at 1100 m. altitude (MA-28375, OSC). Associated mycorrhizal hosts: C. avellana, Pinus sp., Populus sp. and Q. ilex. Tuber foetidum Vitt. BALEARES, (Calonge 1982, 1983). BARCELONA, (Calonge et al. 1985b). Associated mycorrhizal hosts: Pinus sp. Tuber gennadii (Chat.) Pat. CACERES, May 1987 (MA-28376, OSC). Associated mycorrhizal hosts: unknown. Tuber levissimum Gilkey GIRONA, November 1989, December 1989, and June 1990; under P. menziesii in pH 5.5 soil (MA-28377, MA-28378, MA-28379, MA-28380, OSC). Associated mycorrhizal hosts: P. menziesii. Tuber maculatum Vitt. GIRONA, October, December 1989; under P. menziesii in pH 5.5 soil (MA-28381, MA-28382, OSC). Associated mycorrhizal hosts: P. menziesii. Tuber malenconi Don., Riouss. & Chev. BARCELONA, March 1987 (MA-28385, OSC). LERIDA, March 1987 (MA-28384, OSC). TERUEL, March 1987 (MA-28383, OSC). Associated mycorrhizal hosts: unknown. Tuber maresa Font-Quer GIRONA, (Codina and Font-Quer 1931). Associated mycorrhizal hosts: unknown. NOTE: 7. maresa seems likely to be an immature collection of 7. mesentericum, but we have not located the holotype for study. Tuber melanosporum Vitt. BARCELONA, (Unamuno 1941), one collection at BP! (Lloyd 22118) pius five collections at TO. GIRONA, (Lazaro Ibiza 1912, Codina and Font-Quer 1931, Heim et al. 1934). 212 GUADALAJARA, December 1987, under Q. i/ex at 1200 m. altitude (MA-28390, OSC), (Calonge et al. 1985b, Moreno et al. 1986). HUESCA, December 1987; under Q. ilex (MA-28386, MA-28387, MA-28388, MA-28391, OSC). TARRAGONA, February - March 1987; under C. sativa (MA-28392 and MA-28393, OSC). TERUEL, March, 1987 (MA-28389, OSC). Associated mycorrhizal hosts: C. sativa, C. avellana, and Q. ilex. Tuber mesentericum Vitt. BARCELONA, (Unamuno 1941). TARRAGONA, March 1987; under C. sativa (MA-28394, OSC). TERUEL, (Calonge 1983). Associated mycorrhizal hosts: C. sativa. Tuber multimaculatum Parladé, Trappe & Alvarez TARRAGONA, March 1987; (MA-26769, MA-26770, OSC). (Alvarez et al. 1992). Associated mycorrhizal hosts: unknown. Tuber nitidum Vitt. CORDOBA, (Moreno et al. 1991). Associated mycorrhizal hosts: Q. i/ex. Tuber oligospermum (Tul. & Tul.) Trappe (= Terfezia oligosperma Tul. & Tul.) LA CORUNA, (Castro and Freire 1982). Associated mycorrhizal hosts: unknown. Tuber panniferum Tul. & Tul CORDOBA, (Moreno et al. 1991). Associated mycorrhizal hosts: Q. i/ex. Tuber puberulum Berk. & Broome BARCELONA, (Heim et al. 1934). Associated mycorrhizal hosts: Q. lex. Tuber rufum Pico:Fries (= Tuber ferrugineum Vitt.) BARCELONA, March 1988; in a 30-40 yr.-old C. ave//ana plantation in a stony pH 7.8 soil (MA-28396, OSC), (Calonge et al. 1985a, b; Moreno et al. 1991). CACERES, (Moreno et al. 1991). CASTELLON, (Calonge et al. 1977). CUENCA, (Calonge et al. 1977). GIRONA, November, December 1989; under P. menziesii in Ph 5.5 soil (MA-28395, MA-28397, OSC). MALAGA, one collection at K. Associated mycorrhizal hosts: C. sativa, C. avellana, Pinus sp., Q. faginea, and Q. ilex. DISCUSSION Throughout five years of field sampling in forests of northern Spain, we observed that hypogeous fungi are scarce and difficult to find. Only species of the genus Rhizopogon were obtained with relative ease every fall and spring, once the field sites where they occur had been identified. The finds of two new taxa of hypogeous Pezizales (Tuberales) and new records of others indicate that systematic research is needed in many areas of Spain to complete the list of hypogeous mycoflora. Animal digging (holes, channels or scraped soil) was observed in mild 215 coastal areas of Asturias and Galicia and in mountains of Catalonia. Animal activity of this sort was not observed at other places where hypogeous fungi were collected. The interrelationship between mycophagous small mammals and hypogeous fungi has been proven in North America (Fogel and Trappe 1978, Maser et al. 1978, Ure and Maser 1982) and in Australia (Malajczuk et al. 1987). Also, wild boars and foxes readily penetrate commercial plantations of Tuber melanosporum protected with electric fences in search of sporocarps. The relationship between animal vectors and hypogeous fungi has not been studied in Spain, although Durrieu et al. (1984) report substantial utilization by small mammals in the French Pyrenees. The scarcity of sporocarps observed in many Spanish forests may be because of the low number of species, or low populations of animal vectors thus reducing spore dissemination of fungi already present in the forests. Many genera of hypogeous fungi, both Ascomycetes and Basidiomycetes, have never been found in Spain. This may indicate lack of long term studies on the hypogeous mycoflora of Spanish forests. Hunt and Trappe (1987) indicate that documenting all species of hypogeous ectomycorrhizal fungi in a forest stand requires long-term collecting over several years. The production of sporocarps varies from year to year (Fogel 1976) and hypogeous fungal species are scarce when compared to epigeous species in stands of conifers (Luoma et al. 1991). The relatively low diversity of tree species (particularly conifers) in Spanish forests compared with that in places such as the American Northwest, where hypogeous fungi abound, could also explain in part the low species diversity of hypogeous — fungi in Spain. Northern Spain has residual virgin forests, but most extensive forests are the result of human reforestation, often with species native to other continents such as Pinus radiata, P. lambertiana, Pseudotsuga menziesii, and Eucalyptus spp. Research programs to introduce exotic forest species into Europe are likely to continue, given the poverty of indigenous genetic material available (CEE 1984). Hence, parallel research on the hypogeous mycorrhizal symbionts of forest tree species in their area of origin and on the native hypogeous fungi in the introduction area is desirable to assess the need of introducing a particular fungal symbiont for successful establishment of the trees. In some Spanish plantations of P. menziesii, the host-specific hypogeous mycorrhizal fungi Rhizopogon subareolatus has been collected consistently and was probably introduced with its host seedlings long ago. Isolations from sporocarps were obtained with relative ease only from the Rhizopogon species. Some species such as R. roseo/us, were collected in both acid (pH 5) and slightly basic (pH 7.4) soils. This genetic diversity is an important consideration in selecting fungal candidates for use in nursery inoculations (Trappe 1977). Knowledge of the adaptability and host range of the fungal symbiont as well as the receptivity and infectivity of the planting site are important in evaluating the success of introduced fungi in improving forestry practices. 214 ACKNOWLEDGEMENTS Financial support for this research was provided by U.S.- Spain Joint Committee for Scientific and Technological Cooperation grant No. CCA-84 11/013. We thank Drs. R. Fogel and C. Walker for their constructive comments on the manuscript. LITERATURE CITED Alvarez, |.F., J. Parladé and J.M. Trappe. 1992. Locu/otuber gennadii gen. & comb. nov. and Tuber multimaculatum sp. nov. (Tuberaceae). Mycologia 84:926-929. Anonymous. 1989. Anuario de Estadistica Agraria. Minist. Agric. Pesca Aliment. Madrid. 678 pp. Arroyo, |., F.D. Calonge, G. Carrascosa and E. Sau. 1989. Nuovi funghi ipogei di Spagna Ill. Hydnocystis clausa (Tul.) Cerutie Genea verrucosa Vitt. Micol. Ital. 1:3-7. Aymerich, J. and X. Llimona. 1986. Gautieria morchellaeformis Vitt. (Gastrosporiales, Gasteromicetes) al Bergueda (Catalunya). Folia Bot. Misc. 5:145. Bowen, G.D. 1973. Mineral nutrition of ectomycorrhizae. In Ectomycorhizae. Their ecology and physiology. Edited by G.C. Marks and T.T. Kozlowski. Academic Press, New York. pp. 151-205. Calonge, F.D. 1973. Estudios sobre hongos. IV. Aportacion al catalogo de las provincias de Madrid y Sevilla. An. Inst. Bot. Cavanilles 30: 19-32. Calonge, F.D. 1982. Adiciones al catalogo de hongos hipogeos de Espana. Garcia de Orta Ser. Estud. Agron. 9:143-146. Calonge, F.D. 1983. Algunos hongos raros 0 nuevos para Espana. Lazaroa 5:283-289. Calonge, F.D. 1990. Check-list of the Spanish Gasteromycetes (Fungi, Basidiomycotina). Crypt. Bot. 2:33-55. Calonge, F.D. and V. Demoulin. 1975. Les Gasteromycetes d’Espagne. Bull. Soc. Mycol. Fr. 91:247-292. Calonge, F.D. and J.A. Oria de Rueda. 1988. Aportacion a la micoflora de la provincia de Almeria. Bol. Soc. Micol. Cast. 12:93-106. Calonge, F.D. and A. Zugaza. 1973. Catalogo de los hongos presentados en la | exposicion de Madrid, del 1 al 5 de Noviembre de 1972. An. Inst. Bot. Cavanilles 30:33-55. Calonge, F.D., A. Rocabruna and M. Tabarés. 1985a. Nuevos datos sobre los hongos hipogeos de Espana. Bol. Soc. Micol. Cast. 9:45-54. Calonge, F.D., A. Rocabruna, M. Tabarés and N.B. Rodriguez. 1985b. Nuevos datos sobre los hongos hipogeos de Espana. Il. Géneros Balsamia, Delastriay Genea, novedades para el catalogo espanol. Butll. Soc. Cat. Micol. 9:57-64. Calonge, F.D., M. de la Torre and M. Lawrynowicz. 1977. Contribucién al estudio de los hongos hipogeos de Espana. An. Inst. Bot. Cavanilles 34:15-31. 215 Castellano, M.A. 1990. The new genus Trappea_ (Basidiomycotina, Hysterangiaceae), a segregate from Hysterangium. Mycotaxon 38: 1-9. Castellano, M.A. and R. Molina. 1989. Mycorrhizae. In: The container tree nursery manual, Vol. 5, Handbk. 674. Edited by T.D. Landis, R.W. Tinus, S.E. McDonald and J.P. Barrett. U.S.D.A. For. Serv., Washington, D.C. pp. 101-167. Castellano, M.A., J.M. Trappe and R. Molina. 1985. Inoculation of container- grown Douglas-fir with basidiospores of Rhizopogon vinicolor and R. colossus: effects of fertility and spore application rate. Can. J. For. Res. 15:10-13. Castro, M.L. and L. Freire. 1982. Terfezia berberiodora y T. oligosperma encontradas en Galicia, nuevas para Espana. Collect. Bot. (Barc.) 13:44 1- 444. CEE. 1984. L’Europe verte. La Couverture verte de L’Europe. Nos fdrets d’aujourd’hui et de demain. Brussels. 24 pp. Codina, J. and P. Font-Quer. 1931. Introduccié a l’estudi dels macromicets de Catalunya. Cavanillesia 3: 100-189. Durrieu, G., M. Genard and F. Lescourret. 1984. Les micromammiféres et la symbiose mycorrhizienne dans une forét de montagne. Bul. Ecol. 15:253- 263. Fogel, R. 1976. Ecological studies of hypogeous fungi. Il. Sporocarp phenology in a western Oregon Douglas-fir stand. Can. J. Bot. 54:1152-1162. Fogel, R. 1980. Additions to the hypogeous mycoflora of the Canary Islands and | Madeira. Contr. Univ. Mich. Herb. 14:75-82. Fogel, R. and J.M. Trappe. 1978. Fungus consumption (mycophagy) by small mammals. Northwest Sci. 52:1-31. Font-Quer, P. 1954. Plantas de Greixa (Pirineo catalan). Collect. Bot. (Barc.) 4:173-177. Galan, R. and A. Ortega. 1982. Geopora tenuis (Fuckel) Galan & Ortega comb. nov., nuevo pezizal para la micoflora espanola. Bol. Soc. Micol. Cast. 7:49-51. Heim, R., P. Font-Quer and J. Codina. 1934. Fungi Iberici. Observations sur la flore mycologique catalane. Treb. Mus. Cienc. Nat. Barc, Ser. Bot. 15, 146 pp. Honrubia, M. 1984. Labyrinthomyces donkii Malengon, en el S.E. de Espana. Int. J. Mycol. Lichenol. 1:345-349. Honrubia, M. and X. Llimona. 1981. Aportacion al conocimiento de los hongos del S.E. de Espafia. IV. Tres citas nuevas para la microflora espafola. Pustularia insignis, Tuber borchii, Leucogaster cf. floccosus. An. Univ. Murcia Cienc. 37:81-90. Honrubia, M., P. Berthet and X. Llimona. 1983. Contribution a la connaissance des champignons du sud-est de I’Espagne. VII. Pezizales (Ascomycetes). Bull. Soc. Linn. Lyon 52:46-62. Honrubia, M., F.D. Calonge, V. Demoulin, G. Moreno and X. Llimona. 1982. Aportacion al conocimiento de los hongos del S.E. de Espana VI: Esclerodermatales, Licoperdales, Nidulariales, Falales, Himenogasterales, Podaxales (Gasteromicetes, Basidiomicetes). An. Univ. Murcia Cienc. 38: 101-132. 216 Honrubia, M., A. Cano and C. Molina-Nifirola. 1990. Nota sobre hongos hipogeos del semiarido murciano. Resumenes V Reunién Conjunta de Micologia. Rev. Iberoamericana Micol. 7:95. Hunt, G.A. and J.M. Trappe. 1987. Seasonal hypogeous sporocarp production in a western Oregon Douglas-fir stand. Can. J. Bot. 65:438-445. Jeppson, M. 1987. Notes on some Spanish Gasteromycetes. Bol. Soc. Micol. Madr. 11:267-282. Lazaro Ibiza, B. 1908. Nuevos Tuberaceos de Espana. Rev. R. Acad. Cienc. Exactas Fis. Nat. Madr. 6:801-826. Lazaro Ibiza, B. 1912. Notas micoldgicas. Coleccién de datos referentes a los hongos de Espafia. 3° serie. Mem. R. Soc. Esp. Hist. Nat. 7:287-341. Llimona, X. 1983. Sobre fongs de primavera a Catalunya. Butll. Soc. Cat. Micol. 7:33-45. Llimona, X. 1984. Llista d’espécies de l’exposicio de fongs organitzada a Barcelona l’octubre de 1977. Butll. Soc. Cat. Micol. 8:39-46. , Llistosella, J. and M. Aguasca. 1986. El 1° mini foray de la British Mycological Society a Catalunya. Butll. Soc. Cat. Micol. 10:19-33. Losa Quintana, J.M. 1974. Contribuci6n al conocimiento de los Ascomycetes con apotecios de Catalufia. Collect. Bot. (Barc.) 9:45-60. Luoma, D.L., R.E. Frenkel, and J.M. Trappe. 1991. Fruiting of hypogeous fungi in Oregon Douglas-fir forests: seasonal and habitat variation. Mycologia 83: 335-353. Maire, R. 1937. Fungi Catalaunici. Series Altera. Contribution a l'étude de la Flore Mycologique de la Catalogne. Publ. Inst. Bot. Barc. 3, 128 pp. Maire, R., J. Codina and P. Font-Quer. 1933. Fungi Catalaunici. Serie Botanica. Contributions a l’étude de la Flore Mycologique de la Catalogne. Publ. Mus. Cienc. Nat. Barc. 15: 1-120. Malajczuk, N., J.M. Trappe and R. Molina. 1987. Interrelationships among some ectomycorrhizal trees, hypogeous fungi and small mammals. Western Australian and northwestern American parallels. Aust. J. Ecol. 12:53-55. Malengon, G. and R. Bertault. 1971. Champignons de la Peninsule Ibérique. I. Explorations entre le Midi valencien et le Montseny. Acta Phytotaxon. Barc. 8: 1-68. Malencon, G. and R. Bertault. 1972. Champignons de la Peninsule ibérique. IV. Les lles Baléares. Acta Phytotaxon. Barc. 11:1-64. Martin, M.P. 1988. Aportacidn al conocimiento de las higroforaceas y los gasteromicetos de Catalufia. Edic. Esp. Soc. Cat. Micol. 2. 508 pp. Maser, C., J.M. Trappe and R.A. Nussbaum. 1978. Fungal-small mammal interrelationships with emphasis on Oregon coniferous forests. Ecology 59:799-809. Maublanc, M.A. 1936. Rapport sur la session générale de la Societe Mycologique de France, tennue a Barcelone de 19 a 27 octubre 1935. Bull. Soc. Mycol. Fr. 52:17-32. Mendaza, R. and G. Diaz. 1987. Las setas. Ed. Iberduero, Vizcaya. 932 pp. Molina, R. and J.G. Palmer. 1982. Isolation, maintenance, and pure culture manipulation of ectomycorrhizal fungi. In: Methods and principles of mycorrhizal research. Edited by N.C. Schenck. Amer. Phytopathol. Soc., St. Paul, MN. pp. 115-129. 247, Molina, R. and J.M. Trappe. 1982. Patterns of ectomycorrhizal host specificity and potential among Pacific Northwest conifers and fungi. For. Sci. 28:423- 458. Moreno, G., R. Galan and A. Ortega. 1986. Hypogeous fungi from continental Spain. |. Cryptogam. Mycol. 7:201-229. Moreno, G., R. Galan and A. Montecchi. 1991. Hypogeous fungi from peninsular Spain. Mycotaxon 42:201-238. Ortega, A., R. Galan and M. de la Torre. 1981. AportaciOn al estudio de los hongos de Andalucia. IV. El Orden Pezizales en la provincia de Granada. Bol. Soc. Micol. Cast. 6:39-50. Parke, J.L., R.G. Linderman and C.H. Black. 1983. The role of ectomycorrhizas in drought tolerance of Douglas-fir seedlings. New Phytol. 95:83-95. Singer, R. 1947. Champignons de la Catalogne. Espéces observées en 1934. Collect. Bot. (Barc.) 1:199-246 Svrcek, M. 1958. Ill. Rad Hymenogastrales - Hlizotvaré. /n: A. Pilat, Flora CSR, Gasteromycetes. Prace Cesk. Akad. Véd, Prague. pp. 121-208. Tabares, M. and A. Rocabruna. 1987. Aportaci6n al conocimiento de los hongos de la sierra de Collcerola (Catalunya). Butll. Soc. Cat. Micol. 11:83-98. Trappe, J.M. 1977. Selection of fungi for ectomycorrhizal inoculation in nurseries. Annu. Rev. Phytopathol. 15:203-222. Trappe, J.M., M.A. Castellano and N. Malajczuk. 1992. Labyrinthomyces, Dingleya and Reddellomyces gen. nov. (Ascomycotina). Aust. Syst. Bot. 5:597-611. Unamuno, L.M.. 1941. Enumeraci6n y distribuciOn geografica de los Ascomycetes de la Peninsula Ibérica y de las Islas Baleares. Mem. R. Acad. Cienc. Exactas Fis. Nat. Madr. Ser. Cienc. Nat. 8: 1-403. Ure, D.C. and C. Maser. 1982. Mycophagy of red-backed voles in Oregon and Washington. Can. J. Zool. 60:3307-3315. ee ut a hy My er BY hy ait ; Ny 7 WG v va \. : > at es ‘ ak } mee a a yt y | e. A ] ite = ‘ \ asta A 6 1 Rea A! Weibel iF ¥ Taahe af eek Sih an M4 | et Ct ae ; we ya has 2 ss a vente, ee ty eee 10 10. Ascospores subspherical with rounded ends, dark brown to almost black, (10-) 11-12 (-13) 227-89) ema ee iets clef, 6 keh LAR gees octane. ele, eit Biscogniauxia cf. nummularia NOTE: Ascospores of Mexican material of B. nummularia frequently have pinched ends. 10. Ascospores ellipsoid-inequilateral to navicular ............. 11 11. Ascospores with narrow to narrowly-rounded ends, dark brown to almost black, (13-)13.5-17(-19) x 8-8.5 um. Stromata on Quercus spp. wood in temperate forests}. ar. ee Biscogniauxia nummularia var. exutans 11. Ascospores smaller. Stromata on wood in subtropical forests or in tropical FALUNMOTESIS (eee ere ean ne IV Wicenp dare occ eaters Pee car ek ae eee ne 12 12,» 5 -AsCospores: (8.54)9-12)x9.5:6(-6.5)) [Mens = eae ee te ne SCO CAEP CTR EME ey RE Biscogniauxia nummularia var. merrillii 12s), Ascospores.10212.5(-13.5)sx.6.5-7(-8), Me se. ee ee I A ee Ste Biscogniauxia nummularia var. "pseudopachyloma" Biscogniauxia atropunctata (Schwein.) Pouzar, Cesk4 Mykol. 33:216. 1979. The teleomorph is much as described by Miller (1961). Ascus apical ring bluing in Melzer’s iodine reagent, cuneate, 3-3.5 wm high x 7-8 “wm broad. Ascospores (30.5-)32-34(-37) x (12-)13-15 ym, with straight germ slit full-length on convex side. The anamorph was described by Martin (1969) (as Numulariola atropunctata). SPECIMEN EXAMINED: Nuevo Leén state, Iturbide municipality, Ejido La Purisima, Alt. 2000 m, no date, Marmolejo 185, on wood of Quercus sp. in an oak forest (ITCV;JDR). 233 NOTES: This pathogen of Quercus spp. principally in the southern USA (Miller, 1961), was found in México on its expected host genus. Hypoxylon cf. comedens Ces., Atti Accad. Sci. Fis. 8:19. 1879. Teleomorph is as described by Miller (1961) except that the thickness of the stroma is greater, i.e., 1.8-4 mm vs 1-1.5 mm, and perithecia are broader, i.e., 1-2.0 mm vs 1-1.2 mm. SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de Chajul, VII.87, San Martin 376, on wood in evergreen tropical rain forest (ITCV;JDR). NOTES: This fungus is similar to Biscogniauxia in the absence of a basal entostroma which at maturity is partially lost (as in Ustulina Tul. & C. Tul.), the applanate and erumpent stroma, the ostiolar pits, and the light brown, smooth ascospores. It seems most prudent, however, to await the availability of fresh material so that further observations and cultural data can be brought to bear on the taxonomic status of this fungus. Martin (1969) described material from Mexico (as Numulariola). He reported cultures bearing conidiophores of the Nodulispor- tum type (as Acrostaphylus) and of the Xylocladium type (as Basidiobotrys). This indicates that his fungus might, in fact, be a species of Camillea. We have not seen Martin’s material, however. Type material of H. comedens is from Indonesia and its smooth ascospores that possibly bear a germ slit are unlike Camillea (Laessge et al., 1989). Biscogniauxia divergens (Theiss.) A. J. S. Whalley & T. Laessge var. macrospora (J. H. Miller) A. J. S. Whalley & T. Laessge, Myc. Res. 94:239. 1990. The teleomorph is much as described by Miller (1961). Ascospore germ slit Straight, running the entire colored part on convex side. SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules Biosphere Reserve, 21.VIII.87, San Martin 400 & 400B, on dicot wood in evergreen tropical rain forest (ITCV;JDR). NOTES: This material is overmature, but stromatal characters, and shape and size of ascospores fit well Miller’s (1961) description of H. divergens var. macrospora. The type material of this variety was originally collected somewhere in México in 1895 by C. L. Smith. This material was identified by Berkeley as Hypoxylon tinctor (unpublished data) and later recognized as a new variety of Hypoxylon divergens by Miller (1961). Biscogniauxia fuscella (Rehm) San Martin & J. D. Rogers, comb. nov. Basionym: Nummularia fuscella Rehm, Ann. Mycologici 2:176. 1904. Hypoxylon fuscellum (Rehm) J. H. Miller, Mycologia 25:325. 1933. Stromata applanate, erumpent, 0.7-25 cm long x 0.7-2.5 cm broad x 0.5-0.7 234 mm high; externally whitish-gray, internally black. Texture hard but fragile. Surface smooth to undulate due to irregularities of the substrate. Perithecia ovoid, 0.4-0.7 mm broad. Ostioles impressed to slightly papillate. Asci cylindrical, eight- spored, with uniseriate spore arrangement, very short-stipitate, 100-118 2m total length x 9.5-12 wm broad, the spore-bearing part 92-106 zm, with apical ring bluing in Melzer’s iodine reagent, slightly cuneate, 1.5-2 zm high x 4-4.5 wm broad. Ascospores brown to dark brown, ellipsoid-inequilateral to navicular, with narrowly rounded ends, or some with one end narrowly rounded and the other truncate, 13.5-16(-16.5) x 7-8 um, with straight spore-length germ slit whose margins protrude in the manner of clam shell sutures, often with a cellular (?), hyaline appendage at one end ca. 1.5 wm long. Colonies covering Petri plate in 3 wk, sectored, at first white, velvety, somewhat zonate, then with white, woolly growth overlaying the surface, with well- defined margins. Odor strongly yeasty. Reverse at first yellow, then green. Conidium-bearing regions over the entire surface of the colony. Conidiophores erect, branched, with well-defined main axis, 3-4.5 4m broad, brownish toward bases, warted. Conidiogenous cells terminal and lateral, di- or triverticillately branched, cylindrical, 9-15 x 3-4 wm, hyaline, bearing poroid conidial secession scars, with apices somewhat swollen due to conidial production. Conidia cream in mass, individually hyaline, subglobose to ellipsoid, (4.5-)5-6 x 3-4 4m, with flattened bases indicating former points of attachment to conidiogenous cell. Conidia germinating in water. SPECIMEN EXAMINED: Tamaulipas state, Victoria municipality, Km 2 Victoria City-Matamoros City road, 5.1X.88, San Martin 1011 (CULTURED), on wood of Celtis laevigata Willd. ("palo blanco") in semidesertic scrub with Pithecellobium flexicaule (Benth.) Coult. (TCV;JDR). ADDITIONAL SPECIMENS EXAMINED: Nummularia fuscella, USA, Texas, Long 481, Cat. # 11202, (BPI;HOLOTYPE); Hypoxylon atropurpureum Fr. = Hypoxylon cinereo-lilacinum J. H. Miller, USA, Iowa, Decorah, Sept. 1882, Cat. # 1180 (BPI;ISOTYPE). NOTES: The Mexican fungus is similar to the type of Nummulana fuscella Rehm. The applanate stroma, the cuneate apical ring, the color of the ascospores, and the - conidial state assignable to Nodulisponum Preuss clearly link N. fuscella to species of Biscogniauxia. Miller (1961) had placed N. fuscella in his subsection Papillata of section Papillata Hypoxylon due to the more or less papillate ostioles. The color of the ectostroma, the shape of the ostioles, and the size of the ascospores of the Figs. 1-8. Biscogniauxia sp. (José Castillo 1100B), Camillea magnifica, and Camillea mexicana. 1,2. Biscogniauxia sp. (J. Castillo 1100B). 1. Stromata, x 3.8. 2. Stromatal surface, x 16. 3-5. C. magnifica. 3. Stroma, x 6. 4. Stromatal surface, x 16. 5. Stroma (arrow) accompanied by other stromata of a Biscogniauxia, x 1. 6-8. C. mexicana. 6. Portion of stroma, x 1. 7. Stromatal surface, x 16. 8. Stromatal surface, x 24. 235 236 Mexican collection are shared by Biscogniauxia cinereo-lilacina (J. H. Miller) Pouzar (= H. cinereo-lilacinum J. H. Miller), but the latter lacks the strong amyloid reaction of the ascal apical ring, lacks appendaged ascospores, and apparently is restricted to Tilia spp. in cooler localities (Miller, 1961). Biscogniauxia grenadensis (J. H. Miller) A. J. S. Whalley & Laessge, Myc. Res. 94:239. 1990. Teleomorph is much as described by Miller (1961). Ascospores with straight, short germ slit in dark body of spore. SPECIMENS EXAMINED: Chiapas state, La Trinitaria municipality, Montebello Lagoons, 25.V.88, San Martin 840 & 848, on dicot wood in an oak forest mixed with tropical species (ITCV;JDR); San Cristobal de las Casas municipality, El Huitepec Hill, 4.V1.88, San Martin 727, on dicot wood in oak forest (TCV;JDR). NOTES: Collection San Martin 727 is young, but some mature ascospores were found and these were typical of B. grenadensis. Biscogniauxia mediterranea (De Not.) Kuntze, Rev. Gen. Plant. 2, p. 398. 1891. The teleomorph is as described by Miller (1961). The anamorph was described by Martin (1969) (as Numulariola mediterranea), and Jong & Rogers (1972). SPECIMEN EXAMINED: Tamaulipas state, GOmez Farias town, 14.1X.88, San Martin 1130, on wood in median subdeciduous tropical rain forest (ITCV;JDR). ADDITIONAL SPECIMEN EXAMINED: Biscogniauxia mediterranea, Portugal, 1969, N-275 (JDR). NOTES: Mexican material is typical of the species. It was collected in a tropical rain forest. Figs. 9-20. Biscogniauxia sp. (José Castillo 1100B), Biscogniauxia sp. (San Martin 725), Camillea fossulata. 9-11. Biscogniauxia sp. (J. Castillo 1100B). 9. Ascospore, x 1500. 10. Ascospore with germ slit, x 1300. 11. Ascus apical ring, x 1400. Figs. 12-14. Biscogniauxia sp. (SM 725). 12. Ascus apical ring, x 2300. 13. Ascospore, x 2400. 14. Ascospore with short, straight germ slit on concave side, x 2300. 15-20. C. fossulata. 15. Ascospore, x 2000. 16. Portions of two asci with ascospores, x 1900. 17. Ascus apical ring, x 2250. 18. Ascospore with ornamentation, x 20000. 19. Ascospore with ornamentation, x 2500. 20. Ascospore showing ornamentation, x 11000. Figs. 9-14 by BF. Figs. 15-17 by DIC. Figs. 18-20 by SEM. Figs. 9, 10, 13- 16 from water mounts. Figs. 11, 12, 17 from mounts in Melzer’s reagent. 237 238 Hypoxylon microplacum (Berk. & M. A. Curtis) J. H. Miller, Monogr. of the world species of Hypoxylon, p. 129. 1961. The teleomorph is as described by Miller (1961). The anamorph is as described by Jong & Rogers (1972). SPECIMENS EXAMINED: Tamaulipas state, Casas municipality, El Lajeadero Ranch, 8.X1.86, Jesis Garcia 5824, on wood in submontane scrub (ITCV;JDR); Gé6mez Farias town, II.21.88, Chac6n-Jiménez 240B, on wood in median subdeciduous tropical rain forest ITCV;JDR). NOTES: This species was reported by Pérez-Silva (1983) from the state of Oaxaca, México. Martin (1969) considered it to be a Numulariola, but did not observe the scolecosporous conidia reported by Jong & Rogers (1972) and Glawe & Rogers (1986). We are tempted to move it to Biscogniauxia, but she a that it will ultimately be placed in another genus. Biscogniauxia nummulana (Bull.:Fr.) Kuntze var. exutans (Cooke) Van der Gucht, Mycotaxon 45:266. 1992. Hypoxylon nummularium Bull.:Fr. var. excutans (Cooke) J. H. Miller, Monogr. of the world species of Hypoxyion, p. 124. 1961. Teleomorph is much as described by Miller (1961) and Ju (1990). Asci cylindrical, eight-spored, short-stipitate, 119-150 42m total length x 10-11(-11.5) “zm broad, the spore-bearing part 100-120 wm, with apical ring bluing in Melzer’s iodine reagent, cuneate, 3(-3.5) 4m high x 4 “4m broad. Ascospores dark brown to almost black, ellipsoid-inequilateral to navicular with narrow to narrowly rounded ends, (13-)13.5-17(-19) x 8-8.5 4am, with germ slit straight spore-length on convex side. The anamorph was described by Martin (1969) (as Numulariola exutans). SPECIMENS EXAMINED: Nuevo Leon state, Iturbide municipality, Ejido La Purisima, Alt. 2000 m, no date, Marmolejo 199, on wood of Quercus affinis Mart. & Gal. in an oak forest (ITCV;JDR). Tamaulipas state, Hidalgo municipality, Fjido Conrado Castillo, VII.87, San Martin 453, on Quercus sp. wood in oak-pine forest ITCV;JDR). NOTES: This is what Miller (1961) called Hypoxylon nummularium var. exutans. We reluctantly accept it as a variety of Biscogniauxia nummularia because we are uncertain if it is to be considered as a variety or as B. exutans as Martin (1969) believed (as Numulaniola). We require cultural and other data to reach a firm decision. Mexican material differs from Taiwan material (Ju, 1990) in having smaller apical rings, ie., 3(-3.5) zm high x 4 wm broad vs 5-6 4m high x 5 zm broad. The taxonomic significance of apical ring dimensions in Biscogniauxia is unknown. Biscogniauxia nummulana (Bull.:Fr.) Kuntze var. merrillii (Bres.) Van der Gucht, Mycotaxon 45:276. 1992. Hypoxylon nummularium Bull.:Fr. var. merrillii (Bres.) J. H. Miller, Monogr. of 239 the world species of Hypoxylon, p. 126. 1961. The teleomorph is as described by Miller (1961). Colony morphology and anamorphic structures were described by Ju (1990) and Martin (1969) (as Numulariola mermillit). SPECIMENS EXAMINED: Campeche state, Escarcega municipality, Ing. Eduardo Sangri Serrano Forestry Experimental Station, 9.X1.88, San Martin 1180, on wood in median subevergreen tropical rain forest (ITCV;JDR). Quintana Roo state, Oth6n P. Blanco municipality, Tamalcab Island, 11.X1.88, San Martin 1191, on wood in low subdeciduous tropical rain forest (ITCV;JDR); Ejido La Union, 8.XI1.86, San Martin 66, on wood in median subdeciduous tropical rain forest (ITCV;JDR). Tamaulipas state, Ocampo municipality, El Tigre Hill, 8. VIII.88, San Martin 990, on wood in median subdeciduous tropical rain forest (ITCV;JDR); 10. VIII.88, Chac6én 480, on wood in median subdeciduous tropical rain forest (ITCV;JDR); GOmez Farias town 16.VII.87, San Martin 225; 14.X.88, San Martin 1122, on wood in median subdeciduous tropical rain forest (ITCV;JDR). NOTES: This taxon poses the same problems as the previous one. We are undecided as to whether it should be considered as a variety of Biscogniauxia nummulana or aS a separate species (see NOTES on the previous taxon). According to Ju (1990), this taxon and the following one cannot be distinguished by anamorphic characters, the only apparent difference being the slightly smaller ascospores of var. mer7illii. There is a lack of knowledge about host preferences of both taxa. Biscogniauxia cf. nummularia (Bull.:Fr.) Kuntze var. nummularia, Rev. Gen. Plant 2:398.. 1891. The teleomorphis as described by Miller (1961) and Petrini & Miiller (1986) except that Mexican material has some spores with pinched ends. SPECIMEN EXAMINED: Chiapas state, La Trinitaria municipality, Montebello Lagoons, 24.V.88, San Martin 802, on wood in oak-pine forest (ITCV;JDR). NOTES: Biscogniauxia nummulania is primarily a European species. Despite the presence of some spores with pinched ends, Mexican material is very similar to it. On the other hand, the ascospores of this material have almost the same shape of those of H. maculum (Schwein.) J. H. Miller, but are smaller (see Miller, 1961 for description of H. maculum). Welden & Guzman (1978) previously reported this species, as Hypoxylon nummularium, from the state of Veracruz. Biscogniauxia nummularia (Bull.:Fr.) Kuntze var. "pseudopachyloma" Hypoxylon nummularium Bull.:Fr. var. pseudopachyloma (Speg.) J. H. Miller, Monogr. of the world species of Hypoxylon p. 125. 1961. The teleomorph is as described by Miller (1961). The anamorph was described by Ju (1990). 240 SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules Biosphere Reserve, 27.V.88, San Martin 780, on wood in evergreen tropical rain forest (ITCV;JDR). Nuevo Leon state, Santiago municipality, El Cercado, 23.X.88, San Martin 1142, on wood in submontane scrub with Quercus spp. ITCV;JDR). Tamaulipas state, GOmez Farias town, 20.VII.88, San Martin 976, on wood of Citrus sinensis (L.) Osbeck. in an abandoned orchard. NOTES: Material listed above seems to represent what Miller (1961) considered to be variety pseudopachyloma of B. nummularia (as Hypoxylon). We are presently unable to decide whether pseudopachyloma should be considered a variety of B. nummularium or a separate species as did Martin (1969) (as Numulanrola pseudopachyloma). (See also remarks on other putative varieties of B. nummulana herein.) Biscogniauxia sp. (José Castillo 1100B). Figs. 1-2, 9-11. Stromata applanate, erumpent, covering extensive areas of the host, more than 50 cm long x 0.4-0.6 mm thick; externally at first bluish-gray then blackish- white, internally black. Texture hard. Surface uneven due to perithecial ostioles and irregularities of the substrate. Perithecia ovoid to more or less pyriform, 0.2- 0.4 “m high x 0.1-0.3 mm broad. Ostioles prominently papillate surrounded by a blackish area. Asci cylindrical, eight-spored, with uniseriate spore-arrangement, very short-stipitate, 140-156 4m total length x 10-12 4m broad, the spore-bearing part 132-143 zm, with apical ring bluing in Melzer’s iodine reagent, rectangular, 1.5-3 zm high x 3-4.5 4m broad. With hyaline paraphyses 8-9 4m broad at bases. Ascospores dark brown, ellipsoid-inequilateral to navicular with narrow to rounded ends, 18.5-20(-21) x (7-)7.5-9 4m, with straight germ slit spore-length on convex side. SPECIMENS EXAMINED: Tamaulipas state, Casas municipality, E] Lajeadero Ranch, 8.X.86, José Castillo 1100B, on wood of living Quercus sp. in an oak forest (ITCV;JDR); Km 45 Victoria City-Soto La Marina road, San Martin 1090, on wood of Quercus sp. in an oak forest (ITCV;JDR). NOTES: In its host preference and the whitish color of the surface in mature specimens this fungus resembles Biscogniauxia atropunctata, but differs in the presence of blackish areas surrounding perithecial ostioles, and smaller asci, apical ring and ascospores (see descriptions of H. atropunctatum in Miller, 1961, and -Biscogniauxia atropunctata herein). It might be a variant of B. atropunctata or even a new species. We prefer to postpone this decision until cultures are established and examined. Biscogniauxia sp. (San Martin 725). Figs. 12-14. Stromata applanate, erumpent, 0.9-2.5 cm long x 0.7-1 cm broad; externally and internally black. Surface smooth with papillae. Texture carbonous. Perithecia more or less tubular, 0.8-0.9 mm high x 0.4-0.5 mm broad. Ostioles moderately to prominently papillate-conical. Asci cylindrical, eight-spored, with partly biseriate 241 spore-arrangement, short-stipitate, 117-135 fim total length x 6-7.5 4m broad, the spore-bearing part 95-106 44m, with apical ring bluing faintly in Melzer’s iodine reagent, short cylindrical to slightly cuneate, 1.5-2 4m high x 2.5-3 4m broad. Ascospores brown, ellipsoid-inequilateral to navicular, with narrow to acute ends, 13-16(-17) x (4.5-)5-5.5(-6) Am, with germ slit straight, much less than spore-length on the flattened side. SPECIMEN EXAMINED: Chiapas state, San Cristobal de las Casas municipality, E] Huitepec Hill, 4. VI.88, San Martin 725, on wood in oak forest ITCV;JDR). NOTES: Stromatal characters of this species are reminiscent of Hypoxylon nummularium vat. theissenit (Sydow) J. H. Miller, but the ascospores have narrow to acute ends and the germ slit is shorter than that of the ascospores as depicted by Miller (1961). Camillea Fr. Camillea Fr. includes taxa with stromata that are applanate, cylindrical, or broadly conic-truncate. They are usually dark brown or black and highly carbonaceous. The stroma is bipartite, the outermost layer shed to expose the ostiolar openings. Ostioles take various forms and configurations and are highly diagnostic for many species. A single ostiolar canal services one or several perithecia, depending upon the species. Asci feature apical rings that blue in Melzer’s iodine reagent, are approximately as broad as high, and are rhomboid or diamond-shaped. Ascospores are almost hyaline, pale yellow, or light brown and lack a germ slit. By light microscopy the ascospores appear smooth to granular or slightly ornamented. By scanning electron microscopy they are seen to be ornamented with warts, spines, pits, reticulations, or ribs combined with an intricate substructure. Anamorphs where known are accommodated in Xylocladium Sydow (= Basidiobotrys Hohnel). Many of the species now recognized as Camillea were, until recently, included in Nummularia or Hypoxylon. A recent contribution toward a monograph of Camillea reviews most current information on the genus (Laessge et al., 1989). KEY TO SPECIES OF CAMILLEA FROM MEXICO ii Ascospores light brown to brown, verrucose or echinulate .......... Z iL Ascospores subhyaline to light brown, punctate to reticulate or more complexly ornamented (ornamentation seen clearly only by SEM) .... 3 fap Stromata short-cylindrical or pulvinate, protruding 3 mm or more above the host surface level. Ostioles papillate-conical to hemispheric. Ascospores (9-)10-11(-12) x 4-5 um ............. Bre eee. rE, «oh REI pons C. cf. verruculospora Ze Stromata nearly applanate, or short cylindrical, not protruding more than 3 mm above the host surface level (in our material). Ostioles obscure to slightly papillate. Ascospores 12-16 x 5-6 242 in 11. Level 28 eerie tat ae ones eae ac ee SEE OPC eae LL eo eS Se 4 Stromata applanate, not reaching 3 mm above the host surface level .. 6 4. Ostioles in radiating depressions. Ascospores (12-)13-15 x 3.5- ACD EAI Gringc 22h Lora DINE, jo cl's. £48 ogee, MARIS Sale hh Feria Fees C. stellata 4, Ostioles not in radiating depressions... ee ee 5 Stromata with mucronate-labiate apex. Ascospores cylindrical, 8-9 x 4-5 PEM es ate Cra anate Capa ae atOeE ocala! cick - seaiaeaiity Gg Levies ta C. mucronata Stromata with the upper part discoid-convex. Ascospores navicular to somewhat crescentic, (13.5-)14-16 x 4(-4.5) um ........... C. labellum 6. Ascospores ellipsoid with one end beveled ................. 7 6. Ascospores ellipsoid-ineq uilateral, navicular, or crescentic, with both endsmoreorlessisimilar ee Se ee ee eae 10 Ostioles in minute pits. Ascospores 8-10(-11.5) x (3.5-)4-4.5(-5) um. Stromata chiefly on Quercus wood ................64. C. punctulata OStIOIES NOUN MING: DIES PN ee eras esos ote cock ec suaesl ates ah eee ae 8 8. Ostioles papillate, not located in depressions. Ascospores (7) 15-10. S(T Xx 4-8. IN oem ce cele cine l=. ences sata os C. hainesii 8. Ostioles;locatediin depressions 2) 70) 01%), .s,ersrer ea rata ere vie 9 Depressions tending to remain separate, generally 1 ostiole per depression. Ascospores (7.5-)8-9(-10) x 4-4.5(-5) um .........-22006. C. fossulata Depressions tending to coalesce, generally more than one ostiole per depression. Ascospores 9-11 x 4-5 um .......... C. mexicana sp. nov. 1055 -ASCOSpores 2417 AM OL JONPET oo eres cate ie ohetsseoe Nel tare tee 11 10s." Ascospores less than 21° fim Jong 00) t.rs ok. aay degra bersteee 13 Ostioles prominently papillate, on the upper part of hemispheric projections surrounded by depressions. Ascospores (20-)22-29(-30) x 8-9 wm ...... bite wrhay ee? Ske saat ky fl Mara ae gr LCE re a ee at ee ae Cee C. cf. gigaspora Ostioles not on the top of hemispheric projections; stromatal surface between ostioles planGi coche icles tales nie ye pita oe ee eles oe ant 12 12. Ascospores (19-)21-24 x (4.5-)5-6 fam. Ascus apical ring somewhat rectangular, discrete, flaring upward, 3-4 wm high ..... AOA rch as MUN Gh UO RR ne kc ey AL as C. guzmanii sp. nov. 12. Ascospores (22-)23-33 x (7-)8-9 4m. Ascus apical ring cup- shaped, massive, (12-)12.5-14(-14.5) 4m high x 5-6 “4m broad..... Ss EA A, Madman AE TP 8c MOA oa aa ARS SU IE C. magnifica sp. nov. 13. Ostioles obscure to conspicuously papillate; wood beneath stromata usually stained orange. Ascospores 13-20(-21) x 6-9 wm........... C. tinctor 13. Ostioles obscure to punctate; wood beneath stromata never orange-stained. Ascospores (13-)15-19 x (6-)7-7.5(-8) Mm «6.2... eee eee. C. cyclisca Camillea cyclisca (Mont.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:128. 1989. Teleomorphic features and a Xylocladium anamorph were discussed in Laess¢ge et al. (1989). SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de Chajul, VII.87, San Martin 371, on wood in evergreen tropical rain forest (ITCV;JDR). Tamaulipas state, Gomez Farias town, VII.87, San Martin 380, on wood in median subdeciduous tropical rain forest (ITCV;JDR); Ocampo municipality, 16.V.88, San Martin 672, on wood in low subdeciduous tropical rain forest (ITCV;JDR). NOTES: This taxon seems to be related to C. finctor. The latter stains wood beneath the stromata orange; C. cyclisca does not. The differences between these taxa are subtle. Camillea fossulata (Mont.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:131. 1989. Figs. 15-20. Teleomorph is as described in Laess¢e et al. (1989) except that ascospores are hyaline instead of dilute yellow, and ascus apical rings are diamond-shaped _ instead of urn-shaped. Ascospore surface under SEM is punctate-reticulate. SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, "Montes Azules" Biosphere Reserve, 28.V.88, San Martin 811, on wood in evergreen tropical rain forest (ITCV;JDR); Ejido Boca de Chajul, 29.V.88, San Martin 861, on wood in evergreen tropical rain forest (ITCV;JDR). NOTES: It appears that examination of more material with asci and ascospores in good condition, as well as cultural data, are needed in order to achieve a better concept of this taxon. It seems to be collected fairly regularly in the tropics, but seldom with ascospores and asci in good condition. Camillea cf. gigaspora (Massee) Laessge, J. D. Rogers & Whalley, Myc. Res. 33132511989. 244 Teleomorphic features are described in Miller (1961) under Hypoxylon hemisphaericum J. H. Miller; ascospore ornamentation is as described by Rogers (1977) for H. hemisphaericum. SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Pico de Oro, 31.V.88, San Martin 884, on wood in median subdeciduous tropical rain forest (ITCV;JDR). i NOTES: Few ascospores were found in this material. Nevertheless, their shape and size, and the characteristic disposition of the perithecial ostioles permitted a certain degree of confidence about the specific identity. Camillea guzmanii sp. nov. Figs. 28-31 Stroma fragmentarium, applanatum, erumpens, 4-5 mm longum x 3-3.5 mm latum x 0.7-1 mm crassum, marginibus sterilibus distinctis; extus atrum; intus nigellum. Textura dura. Perithecia ampulliformia, 0.4-0.6 mm alta x 0.3-0.4 mm lata. Ostiola indistincta vel parum papillata. Asci octospori, cylindrici, stipitati, 188-203 4m longitudine x 7-8 4m crassi, partibus sporiferis 135-145 4m, annulo apicali in liquore iodata Melzeri immerso cyanescente, rotundato ad basim et sursum aliquantum infundibuliformi, 3-4 zm alto x 3-4 “wm lato. Paraphyses persistentes, septatae, 5.5-9.5 fam latae. Ascosporae luteolae, unicellulares, ellipsoideo-inequilaterales vel plus minusve lunatae, extremis angustis vel acutis, sub lente laeves, per SEM observatae punctato-reticulatae, (19-)21-24 x (4.5-)5-6 fim. Rima germinationis nulla. Stroma fragmentary, applanate, erumpent, 4-5 mm long x 3-3.5 mm broad x 0.7-1 mm high, with distinct sterile margins; exterior black; interior dark. Texture hard. Perithecia flask-shaped, 0.4-0.6 mm high x 0.3-0.4 mm broad. Ostioles obscure to slightly papillate. Asci eight-spored, cylindrical, stipitate, 188-203 4m total length x 7-8 44m broad, the spore-bearing part 135-145 fim, with apical ring bluing in Melzer’s iodine reagent, rounded at the base and flared upward, 3-4 “4m high x 3-4 fém broad. Paraphyses persistent, septate, 5.5-9.5 fm broad. Ascospores pale yellow, unicellular, ellipsoid-inequilateral to somewhat crescentic, with narrow to acute ends, smooth by light microscopy, punctate-reticulate by SEM, Figs. 21-31. Camillea labellum and C. guzmanii. 21-27. C. labellum. 21. Ascospore, x 2425. 22. Ascospore in upper part of ascus, x 5000. 23. Ascus apical ring, x 2900. 24. Ascospore showing ornamentation, x 5000. 25. Detail of ascospore ornamentation, x 15000. 26. Anamorph. Ampulla with conidiogenous cells and conidial primordium (arrow),-x 1650. 27. Conidiogenous cells bearing conidia (arrow), x 1650. 28-31. C. guzmanit. 28. Ascospore, x 1400. 29. Ascus apical ring, x 2300. 30. Ascospore showing ornamentation, x 4000. 31. Detail of ascospore ornamentation, x 15000. Figs. 21, 22, 26-28 by DIC. Figs. 23 and 29 by BF. Figs. 24, 25, 30, 31 by SEM. Figs. 21, 22, 26-28 from water mounts. Figs. 23 and 29 from mounts in Melzer’s reagent. 245 LQE < 246 (19-)21-24 x (4.5-)5-6 fm. Lacking a germination slit. SPECIMEN EXAMINED: Veracruz state, Catemaco municipality, Catemaco- Montepio road, 4.X1.88, San Martin 1399, on wood in evergreen tropical rain forest (ITCV,HOLOTYPE). NOTES: Camillea guzmanii differs from Camillea magnifica described herein, in having obscure to slightly papillate ostioles instead of punctate ones, in having a much smaller ascus apical ring, ie. 3-4 2m high x 3-4 “wm broad vs (12-)12.5- 14(-14.5) um high x 5-6 4m broad, and in having shorter ascospores, i.e. (19-)21-24 x (4.5-)5-6 fam vs (22-)23-33 x (7-)8-9 Am. Both species have a similar poroid- reticulate ascospore ornamentation seen by SEM, but pores in C. guzmanii are more elongated. The Mexican fungus shares with Camillea hyalospora (Pat.) J. D. Rogers, Laessge, & Lodge its slightly papillate ostioles, but the latter has larger ascospores, i.e. 26-31 x (5-)7.5-9 zm. Ascospore ornamentation of C. hyalospora as seen by SEM is poroid-reticulate but has more elongated pores than those of C. guzmanii [see Rogers, 1975b for ascospore ornamentation of N. hyalospora (as Hypoxylon glycyrrhiza Berk. & M. A. Curtis), and Rogers et al., 1991 for description of C. hyalospora}. Camillea guzmanii is named after the eminent Mexican mycologist, our friend Gast6n Guzman. Camillea hainesii (J. D. Rogers & Dumont) Laess¢e, J. D. Rogers & Whalley, Myc. Res. 93:134. 1989. Teleomorph was described by Rogers & Dumont (1979) as Hypoxylon hainesii based on fragmented material. Stromata applanate, orbicular, erumpent, with sterile margins, 0.9-2.5 cm long x 5-7 mm broad x 0.3-0.8 mm high. Ascospores rectangular to ellipsoid, with one end somewhat acute, the other rounded, (7-)7.5- 10.5(-11) x 4-4.5 wm. SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, "Montes Azules" Biosphere Reserve, 30.V.88, San Martin 823 and 872, on wood in evergreen tropical rain forest (ITCV;JDR). Oaxaca state, Temazcal town, 8.X.88, San Martin 1093, on wood in median subdeciduous tropical rain forest (ITCV;JDR). NOTES: Compared with type material of C. hainesti (see Rogers & Dumont, 1979), the Mexican collections differ in having ascospores more variable in shape and in being slightly larger both in length and width. Camillea labellum Mont., Ann. Sci. nat. Bot. 4:113. 1855; Sensu Dennis, Kew Bull. 2:319. 1957. Figs. 21-27. Teleomorph described in Dennis (1957) and Laessge et al. (1989). Ascospores are smooth under LM, but are seen by SEM to be composed of closely packed rod-shaped elements. Colonies covering 9 cm diam Petri dish in 11 days, with mycelium at first 247 white with yellowish tones, then turning pale-yellow, floccose, becoming felty in old colonies; margin indistinct, with peripheral hyphae woolly to somewhat appressed. Odor strongly yeasty which by continuous transferring gradually disappears. Reverse at first yellowish then greenish, finally dark-green. Conidia produced in 15 days. Hyphae septate, hyaline, of two types: one torulose ca. 4 4m broad, the other even, 6 4m broad. Conidiophores upright, with determinate growth, arising singly and laterally from hyphae, each composed of a mononematous, smooth stipe more than 450 4m long, terminated by a swollen ampulla. Ampullae subglobose to clavate, 10-12 x 16-18 um, bearing conidiogenous cells, with poroid scars at the point of conidiogenous cell attachment. Conidiogenous cells hyaline, continuous, oblong-elliptical, inflated at the apex, numerous, often closely compacted on the surface of the ampullae, 5-9 4m high x 2-4 44m broad, with conspicuous secession scars. Conidia grayish in mass, hyaline individually, borne on the upper part of conidiogenous cells or directly from apparently unspecialized hyphae, ellipsoid to somewhat allantoid with one end rounded, the other truncate to somewhat attenuated, bearing basal scar indicating former point of attachment to conidiogenous locus, 6-11(-13) x (2-)2.5-3 zm, with internal oil droplets. SPECIMENS EXAMINED: Campeche state, Escarcega municipality, "Ing. Eduardo Sangri Serrano" Forestry Experimental Station, 9.X1.88, San Martin 1181C, on wood in median subevergreen tropical rain forest (ITCV;JDR). Chiapas state, Ocosingo municipality, Ejido Boca de Chajul, 29.V.88, San Martin 839, on wood in evergreen tropical rain forest (ITCV;JDR); Chancal4, Rio Chancala, Alt 100 m., 12.23.85, Chacon-Zapata 3346, on wood in evergreen tropical rain forest (XAL). Quintana Roo state, Oth6n P. Blanco municipality, Ejido La Union, IX.87, San Martin 313, on wood in median subdeciduous tropical rain forest (ITCV;JDR); San Felipe Bacalar, 10.X1.88, San Martin 1219 and 1233 (CULTURED), on wood in an acahual (ITCV;JDR). Tamaulipas state, Ocampo municipality, 16.V.88, San Martin 663, on wood in low subdeciduous tropical rain forest (ITCV;JDR). Veracruz state, Catemaco municipality, "Los Tuxtlas" UNAM Biological Station, 4.X1.88, San Martin 1264, on wood in evergreen tropical rain forest ITCV;JDR). NOTES: Mexican material matches the concept of Dennis (1957) for C. labellum. The Xylocladium anamorph supports the hypothesis that the remaining members of the genus Camillea for which the anamorphs have not yet been obtained, will likewise have a Xylocladium state. Camillea magnifica sp. nov. Figs. 3-5, 32-35 Stromata erumpentia, applanata, circularia vel irregularia, restricta, 0.2-3.4 cm longa x 0.15-1.4 cm lata x 0.7-0.8 mm alta, marginibus sterilibus aliquantum elevatis; extus brunneola vel nigella; intus nigella. Textura dura. Perithecia ampulliformia, 0.7 mm alta x 0.3-0.4 mm lata. Ostiola indistincta vel subtiliter punctata vel parum papillata. Asci octospori, cylindrici, breviter stipitati, 185-210 Hm longitudine tota x 9-10 zm crassi, partibus sporiferis 157-177 4m, annulo apicali in liquore iodata Melzeri immerso cyanescente, cupulato, magno, (12-)12.5- 14(-14.5) um alto x 5-6 zm lato. Paraphyses persistentes, septatae, longior quam asci, 4-6 4m latae. Ascosporae luteolae, unicellulares, ellipsoideo-inequilaterales 248 vel plus minusve lunatae, extremis angustis vel acutis, sub lente subtiliter aspris, per SEM observatae punctato-reticulatae, (22-)23-33 x (7-)8-9 Am. Rima germinationis nulla. Stromata erumpent, applanate, circular or irregular, restricted, 0.2-3.4 cm long x 0.15-1.4 cm broad x 0.7-0.8 mm high, with somewhat raised sterile margins; exterior brownish to black; interior blackish. Texture hard. Perithecia flask- shaped, 0.7 mm high x 0.3-0.4 mm broad. Ostioles obscure to finely punctate to slightly raised. Asci eight-spored, cylindrical, short-stipitate, 185-210 4m total length x 9-10 4m broad, the spore-bearing part 157-177 vm, with apical ring bluing in Melzer’s iodine reagent, cup-shaped, massive, (12-)12.5-14(-14.5) 4m high x 5-6 fm broad. Paraphyses persistent, septate, longer than asci, 4-6 (4m broad. Ascospores yellowish, unicellular, ellipsoid-inequilateral to somewhat crescentic, with narrow to acute ends, finely roughened by LM, punctate-reticulate by SEM, (22-)23-33 x (7-)8-9 Am. Lacking germination slit. SPECIMEN EXAMINED: Tamaulipas state, GOmez Farias town, no date, San Martin 258, on wood in median subdeciduous tropical rain forest (ITCV, HOLOTYPE; JDR;ISOTYPE). NOTES: Camillea magnifica has ascospores in the size range of C. gigaspora (Massee) Laess¢ge, J. D. Rogers & Whalley but its ostioles are punctate instead of papillate on raised hemispherical elevations, and its apical ring is much larger, i.e., (12-)12.5-14(-14.5) 4m high vs 4.5-6.5 4m high. The ascospore surface as seen by SEM is poroid-reticulate in both species [see ascospore ornamentation of C. gigaspora (as Hypoxylon hemisphaericum J. H. Miller) in Rogers, 1977]. See NOTES on Camillea guzmanii herein for differences between C. guzmanii and C. magnifica). The name magnifica refers to the large apical ascus ring. Camillea mexicana sp. nov. Figs. 6-8, 36-39 Stromata erumpentia, applanata, ca 1 m longa x 7-10 cm lata x 0.4-0.5 mm alta, marginibus sterilibus; extus brunneola vel nigella; intus nigella. Textura dura. Perithecia oblonga, 0.4-0.5 mm diam. Ostiola subliter papillata, 2-4 disposita in depressionibus non profundis et saepe coalescentibus. Asci octospori, cylindrici, breviter stipitati, 104-119 “zm longitudine tota x 6-6.5 1m crassi, partibus sporiferis 68-78 4m, annulo apicali in liquore iodata Melzeri immerso cyanescente, valde Figs. 32-39. Camillea magnifica and C. mexicana. 32-25. C. magnifica. 32. Ascospore, x 1240. 33. Ascus apical ring, x 1750. 34. Ascospore showing ornamentation, x 3500. 35. Detail of ascospore ornamentation, x 11000. 36-39. C. mexicana. 36. Ascospore, x 23000. 37. Ascus apical ring, x 1625. 38. Ascospore showing ornamentation,x 8000. 39. Detail of ascospore ornamentation, -x 15000. Figs. 32 and 36 by DIC. Figs. 33 and 37 by BF. Figs. 34, 35, 38, 39 by SEM. Figs. 32 and 36 from water mounts. Figs. 33 and 37 from mounts in Melzer’s reagent. 250 rhombeo, 3 um alto x 4 fm lato. Paraphyses persistentes, septatae, 7-10 4m latae. Ascosporae hyalinae vel luteolae, unicellulares, plus minusve oblongae, extremo saepe devexo, sub lente laeves, per SEM observatae plus minusve punctato- reticulatae, 9-11 x 4-5 4m. Rima germinationis nulla. Stromata erumpent, applanate, ca. 1 m long x 7-10 cm broad x 0.4-0.5 mm high, with sterile margins; exterior brownish to blackish; interior blackish. Perithecia oblong, 0.4-0.5 mm diam. Ostioles finely papillate, in groups of 2-4 in shallow depressions which frequently coalesce. Asci 8-spored, cylindrical, short- stipitate, 104-119 zm total length x 6-6.5 44m broad, the spore-bearing part 68-78 jum, with apical ring bluing in Melzer’s iodine reagent, strongly rhomboid, 3 “4m high x 4 4m broad. Paraphyses persistent, septate, 7-10 im broad. Ascospores hyaline to pale yellow, unicellular, more or less rectangular, often sloping at one end, smooth by light microscopy, more or less punctate-reticulate by SEM, 9-11 x 4-5 um. Lacking germination slit. SPECIMEN EXAMINED: Chipas State, Ocosingo municipality, "Montes Azules" Biosphere Reserve, 29.V.88, San Martin 856, on wood in evergreen tropical rain forest ITCV,HOLOTYPE;JDR,ISOTYPE). NOTES: Camillea mexicana differs from C. fossulata (Mont.) Laessge, J. D. Rogers & Whalley in having a stromatal surface that is blackish or brownish-black instead shiny-black, in its faint, more indefinite surface depressions which tend to coalesce rather than remaining separate, in having several ostioles per depression, in having slightly larger ascospores, i.e. 9-11 x 4-5 zm vs (7.5-)8-9(-10) x 4-4.5(-5) fim, and in having unique interlocked, puzzle-like ascospore ornamentation as seen by SEM rather than poroid-reticulate [see description of C. fossulata in Miller, 1961, (as Hypoxylon fossulatum Mont.), and herein]. Camillea mexicana approximates Camillea flosculosa (Starb.) Laessge, J. D. Rogers & Whalley in the disposition of the ostioles and ascospore morphology seen by LM, but differs in having smaller ascospores, i.e. 9-11 x 4-5 fam vs 11-15.5 x 5- 6.5 44m, and in having the ascospore ornamentation referred to above instead of poroid-reticulate [see Rogers, 1977, for description of ascospore ornamentation in Hypoxylon flosculosum (Starb.) J. H. Miller]. The Mexican fungus shares with Camillea punctulata (Berk. & Ravenel) Laessge, J. D. Rogers & Whalley the shape of the ascospores and to a lesser degree the ascospore ornamentation, but the latter species has pitted ostioles instead of slightly papillate ones in faint depressions. This highly distinctive fungus seems to have a transitional type of ascospore ornamentation from poroid reticulate to ribbed anastomosing, or vice versa. Camillea mucronata Mont., Ann. Sci. Nat. Bot. 4:112. 1855. Teleomorph is as described by Medel & Chac6n (1988), and Laess¢e et al. (1989). SPECIMEN EXAMINED: Chac6n-Zapata 3316, 22.XII.85, Michol-Ha falls, Palenque municipality, Chiapas state, on wood in evergreen tropical rain forest (XAL). 251 NOTES: Medel & Chacén (1988) reported the species from the Mexican state of Chiapas. It is interesting to note that, with one exception, material of C. mucronata so far reported was collected near rivers, suggesting that the species is confined to some tropical, riparian host(s) (see Laessge et al. 1989 for distribution of C. mucronata). Camillea obularia (Fr.) Laessge, J. D. Rogers & Lodge, Mycologia 83:224. 1991. Camillea broomeiana (Berk. & M. A. Curtis) Laessge, Rogers & Whalley, Myc. Res. 93:149. 1989. Teleomorph is as described in Laess¢ge et al. (1989) under C. broomeiana; the anamorph was described in Rogers (1975a) [as Nummulania broomeiana (Berk. & M. A. Curtis) J. H. Miller]. SPECIMENS EXAMINED: Oaxaca state, Km 65 Loma Bonita-Sayula off-side road, 21.V.88, San Martin 719, on wood of Delonix regia (Bojer ex Hook) Raf., ("flambollan"), (ITCV;JDR). Tamaulipas state, Gomez Farias town, 17.1.85, Garcia J. 4599, on wood of Mangifera indica L. ("mango"), in an orchard (ITCV;JDR). NOTES: Mexican material fits well the species concept of C. obularia. It is interesting to note that out of 3 collections made by F. C. Deighton in Sierra Leone, one was on Delonix regia and the other on Mangifera indica (see Laessge, et al., 1989). Mexican material was found on those hosts (see SPECIMENS EXAMINED above). Camillea punctulata (Berk. & Ravenel) Laess¢ge, J. D. Rogers & Whalley, Myc. Res. 93:143. 1989. The teleomorph was as described by Miller (1961), and the anamorph was described by Jong & Rogers (1972) [as Hypoxylon punctulatum (Berk. & Ravenel) Cooke in both publications]. SPECIMENS EXAMINED: Chiapas state, La Trinitaria municipality, Montebello Lagoons, 50 km from the Guatemalan border, 24.V.88, San Martin 789, on wood in pine-oak forest ITCV;JDR). Nuevo Le6n state, Santiago municipality, El Cercado, VIII.1987, San Martin 395, on wood of Quercus ryzophylla Weath. in submontane scrub with Quercus spp. (ITCV;JDR). Tamaulipas state, Gomez Farias municipality, Malacate, 31.11.88, Baldazo 219, on wood of Quercus aff. germana Cham. & Schlecht. in cloud forest (ITCV;JDR). NOTES: This species seems to be confined almost exclusively to Quercus spp. (Barnett, 1957; Jong & Rogers, 1972). Its ascospore shape and ornamentation by SEM approximates that of Camillea mexicana described above, but the ascospores of the latter are larger and its ostioles are not in pits [see description of ascospore surface of H. punctulatum in Rogers (1977), and description of C. mexicana herein]. Medel et al. (1989) reported this species [as Hypoxylon punctulatum (Berk. & Ravenel) Cooke] from the Mexican state of Veracruz growing on wood in a 252 cloud forest. Camillea stellata Laessge, J. D. Rogers & Whalley, Myc. Res. 93:144. 1989. Teleomorph is as described in Laess¢e et al. (1989). SPECIMENS EXAMINED: Chiapas state, Palenque municipality, Palenque Ruins, [X.1987, San Martin 296, on wood in evergreen tropical rain forest (ITCV;JDR). Quintana Roo state, Oth6n P. Blanco municipality, Ejido La Union, 8.X11.86, San Martin 37, on wood in median subdeciduous tropical rain forest (ITCV;JDR). NOTES: Mexican material seems to be typical of the species even though it has few ascospores in good condition. Camillea tinctor (Berk.) Laessge, J. D. Rogers & Whalley, Myc. Res. 93:145. 1989. Teleomorph and anamorph are more or less as described by Laess¢e et al. (1989). SPECIMENS EXAMINED: Chiapas state, Ocosingo municipality, Montes Azules Biosphere Reserve, 27.V.88, San Martin 830, on wood in evergreen tropical rain forest (TCV;JDR); Palenque municipality, Palenque Ruins, 2.VI.88, San Martin 876, on wood in evergreen tropical rain forest (ITCV;JDR). Nuevo Le6n state, Santiago municipality, El Cercado, 26.V.88, Moreno 221 and 225, on wood in submontane scrub with Quercus spp. (ITCV;JDR); 23.X.88, San Martin 1146, (same data as Moreno 221 and 225 above); 29.1.89, Moreno 267 (CULTURED), on wood of Morus nigra L. ("mora"), in submontane scrub with Quercus spp., (ITCV;JDR). Tamaulipas state, G6mez Farias municipality, "El Nacimiento’, 29.V.88, San Martin 620, on wood in median subdeciduous tropical rain forest (ITCV;JDR); Hidalgo municipality, Ejido Los Mimbres, "Los Alamos", 7.VII.87, San Martin 200, on wood of Platanus sp., on a river bank (ITCV;JDR). Veracruz state, Jalapa Enriquez municipality, "Francisco Javier Clavijero" Botanical Garden, VIII.87, San Martin 620 A, on wood in cloud forest ITCV;JDR); San Andrés Tuxtla municipality, Eyipantla falls, 5.X1.88, San Martin 1339 (CULTURED), on wood of a riparian tree (ITCV;JDR). NOTES: Mexican material does not always impart an orange stain to the wood beneath stromata; perithecial size is variable, i.e., in Moreno 225 they measure 0.2- 0.3 mm broad and in San Martin 620 "A" they measure 0.6-0.7 mm; ascospore size is variable among collections, i.e., in Moreno 267 the range is 13-17 fam while in San Martin 1339 it is 18-20(-21) zm; ascospores of Moreno 267 readily germinated in water and produced germ tubes at one or both ends of the propagules, even when stromata lacked asci and thus appeared old. Anamorphic features are variable as well, i.e., San Martin 1339 produced conidia highly variable in size, 6-18 x (1.5-)2-3(-3.5) 4m, and Moreno 267 produced ampullae not as broad as in "typical" isolates, and Nodulisporium-type conidio- phores were more common than in other isolates. It appears necessary to obtain additional fresh collections of this species and study the teleomorphic and 253 anamorphic features in order to assess its limits. Camillea cf. verruculospora J. D. Rogers, Laessge & Lodge, Mycologia 83:224. 1991; The teleomorph is as described by Rogers et al. (1991), except that ascospores of Mexican material are smaller, i.e., (9-)10-11(-12) x 4-5 zm vs (11-)12- 13.5 x 4.5-5(-6.5) wm. SPECIMEN EXAMINED: Chiapas state, Ocosingo municipality, Ejido Boca de Chajul, 29.V.88, San Martin 837, on wood in an acahual (ITCV;JDR). NOTES: The Mexican collection is probably not fully mature. This might account for the spores being smaller than usually encountered. ACKNOWLEDGMENTS PPNS No. 0121, Department of Plant Pathology, Project 1767, Washington State University, College of Agriculture and Home Economics. This study was supported by National Science Foundation Grant BSR-9017920 to JDR. We thank the Consejo del Sistema Nacional de Educaci6n Tecnologica for furnishing support to SM during 1986-88 for the project Ecologia y Distribuci6n de Hongos Destructores de Madera en el Noreste de Mexico and to Consejo Nacional de Ciencia y Tecnologia de Mexico (CONACYT) for a grant to SM during his graduate studies. We thank Donald P. Rogers, Auburn, WA, for correcting the Latin descriptions. We thank the following associates at Washington State University: Y.-M. Ju for his aid in numerous taxonomic matters; Michael J. Adams for aid with SEM and photography; Jane Lawford for reconstructing the manuscript; Lori M. Carris for reading the manuscript. We thank Brenda E. Callan, Forestry Canada, Victoria, B.C., Canada for reading the manuscript. We thank the curators of cited herbaria for the loan of specimens. SM had extensive aid and encouragement in the portion of the studies conducted in Mexico. He thanks the following persons: Professor José Castillo Tovar for the first lectures on mycology and encouragement; Dr. Gaston Guzman for his friendship and encouragement; Biol. Lucrecia Garcia Alanis, Biol. Arnulfo Moreno, Dr. Efrén Cazares Gonzalez, Biol. Santiago Chac6n Jiménez, Biol. Bertha Baldazo, Biol. Pablo Lavin, Biol. Catalina Flores, M. en C. Gonzalo Guevara, Biol. Jess Garcia, and Biol. Maria Concepci6n Herrera for aid in collecting and the latter also for her aid in identifying plant material. SM likewise thanks Biol. Santiago Chac6n-Jiménez and Biol. Arnulfo Moreno for sharing their interest in the adventure of collecting; Dr. Alberto Rodriguez Fernandez and Biol. Javier Chavelas Polito for providing collecting facilities in Quintana Roo and Biol. Francisco Quinto Adrian for his friendship and aid in Quintana Roo; Biol. Antonio Espinoza Banda for collecting facilities in Campeche and Biol. Jorge Ayala Guajardo for his friendly aid while collecting in the beautiful and interesting state of Chiapas. 254 LITERATURE CITED Barnett, H. L. 1957. Hypoxylon punctulatum and its conidial stage on dead oak trees and in culture. Mycologia 49:588-595. Callan, B. E. and J. D. Rogers. 1990. Teleomorph-anamorph connections and correlations in some Xylaria species. Mycotaxon 36:343-369. Dennis, R. W. G. 1957. Further notes on tropical American Xylariaceae. Kew. Bull. 1957. 297-332. Glawe, D. A. and J. D. Rogers. 1986. Conidial states of some species of Diatrypaceae and Xylariaceae. Can. J. Bot. 64:1493-1498. Holmgren, R. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum. Part I: The herbaria of the world. Ed. 8. New York Botanical Garden. 693 p. INEGI. 1983. Sintesis Geografica y Nomenclators de los estados de Coahuila, Morelos, Nuevo Leé6n y Tamaulipas. Secretaria de Programacién y Presupuesto. México. INEGI. 1988. Carta Topografica de México 1:250 000. Secretaria de ProgramaciOn y Presupuesto. México. Jong, S. C., and J. D. Rogers. 1972. Illustrations and descriptions of conidial states of some Hypoxylon species. Wash. State Agric. Exp. Sta. Bull. 71. 51 p. Ju, Y.-M. 1990. Studies of Xylariaceae from Taiwan. M. Sc. Thesis. Washington State University, Pullman, WA. 171 p. Koeppen, W. 1948. Climatologia. Fondo de Cultura Econémica. México, D.F. 478 p. Laessge, T., J. D. Rogers, and A. J. S. Whalley. 1989. Camillea, Jongiella and light-spored species of Hypoxylon. Myc. Res. 93:121-155. Martin, P. 1969. Studies in the Xylariaceae: VI. Daldinia, Numulanola and their allies. J. S. African Bot. 35:267-320. Medel, R. and S. Chacon. 1988. Ascomycetes lignicolas de México, II. Algunos Pyrenomycetes y Discomycetes. Micologia Neotropical Aplicada 1:87-96. Medel, R., S. Chac6n, and G. Guzman. 1989. Especies conocidas y nuevos registros de Hypoxylon (Sphaeriales, Xylariaceae) en México. Rev. Mex. Mic. 5:149-168. Miller, J. H. 1961. A monograph of the world species of Hypoxylon. Univ. Georgia Press, Athens. 158 p. Pérez-Silva, E. 1983. Distribucién de algunas especies del género Hypoxylon (Pyrenomycetes) en México. An. Inst. Biol. Univ. Nac. Aut. Méx. 54:1-22. Petrini, L. E. and E. Miller. 1986. Haupt- und nebenfruchtformen europaischer Hypoxylon-Arten (Xylariaceae, Sphaeriales) und verwandter pilze. Mycologia Helv. 1:501-627. Pouzar, Z. 1979. Notes on taxonomy and nomenclature of Nummulana (Pyrenomycetes). Ceska Mykologie. 33:207-219. Rogers, J. D. 1975a. Nummularia broomeiana: conidial state and taxonomic aspects. Am. J. Bot. 62:761-764. Rogers, J.D. 1975b. The ascospore of Hypoxylon glycyrrhiza. Mycologia 67:657- 662. 255 Rogers, J. D. 1977. Surface features of the light-colored ascospores of some applanate Hypoxylon species. Can. J. Bot. 55:2394-2398. Rogers, J. D. and K. P. Dumont. 1979. Los hongos de Colombia VI. Two new applanate species of Hypoxylon. Mycologia 71:807-810. Rogers, J. D., T. Laessge and J. Lodge. 1991. Camillea: New combinations and a new species. Mycologia 83:224-227. Rzedowski, J. 1978. La vegetacion de México. Ed. LIMUSA. México. 431 p. San Martin, F. and J. D. Rogers. 1989. A preliminary account of Xylaria of México. Mycotaxon 34:283-373. Welden, A. L. and G. Guzman. 1978. Lista preliminar de los hongos, liquenes y myxomycetos de las regiones de Uxpanapam, Coatzacoalcos, Los Tuxtlas, Papaloapan y Xalapa (Parte de los estados de Veracruz y Oaxaca). Bol. Soc. Mex. Mic. 12:59-102. APPENDIX Location of Collecting Sites For the exact location of the sites where the field work was conducted, the Atlas de México and the Geographical Syntheses and Nomenclators of the pertinent states, edited by the Instituto Nacional de Geografia y Estadistica (INEGI, 1983, 1988) were consulted. The main locations are listed below. A. Campeche state: 1) Escarcega municipality, Ing. Eduardo Sangri Serrano Forestry Experimental Station, 18° 40’ Lat; 90° 45’ Long; Topographic Map E15-6. B. — Chiapas state: 1) Ejido Boca de Chajul, 16° 07’ Lat; 99° 52’ Long; Topographic Map 5 12;D 153; 2) Ejido Pico de Oro, 16° 20’ Lat; 90° 45’ Long; Topographic Map F152;D153. 3) SEI wHuitepeci Hill.) 16; 645°) Lats.92° 4.43" sLongah27509m s Alt Topographic Map E1511. 4) El Triunfo Hill, 15° 40’ Lat; 112° 48” Long; 2450 m Alt; Topographic Map D15-2. 5) | Malpaso Dam, 16° 48’ Lat; 93° 20’ Long; Topographic Map E-15-11. 6) Montebello Lagoons, 16° 08’ Lat; 91° 42’ Long; Topographic Map Bi512;D 153: 7) Palenque Ruins, 17° 30’ Lat; 92° 01’ Long; Topographic Map E-15- } 256 Coahuila state: 1) | Arteaga Municipality, San Antonio de las Alazanas, 25° 16’ Lat; 10° 34’ Long; Topographic Map G14-C35. Federal District: 1) Km 43 Mexico-Cuernavaca Road, 19° 02’ Lat; 99° 12’ Long; Topographic Map E-14-2. Jalisco state: 1) | Cacoma Mountains, 19° 58’ Lat; 104° 32’ Long; Topographic Map E-13-2-5. 2) Guadalajara City, 20° 39’ Lat; 103° 18’ Long; 1550 m Alt; Topographic Map F-13-12. 3) La Manzanilla, 20° 00’ Lat; 103° 09’ Long; 2050 m Alt; Topographic Map F13-12. 4) Manantl4n Biosphere Reserve, 19° 37’ Lat; 104° 13’ Long; Topographic Map E-13-2-5. 5) | Mazamitla, 19° 55’ Lat; 103° 01’ Long; Topographic Map E-13-3. 6) Tecalitl4n, 19° 28’ Lat; 103° 18’ Long; 1140 m Alt, Topographic Map E13-3. Morelos state: 1) La Pera Curve, 25° 14’ Lat; 99° 14’ Long; Topographic Map E14- AS9. Nuevo Le6n state: 1) El Chipinque, 25° 55’ Lat; 100° 22’ Long; Topographic Map G-14- C15. 2) El Cercado, 25° 24’ Lat; 100° 08’ Long; Topographic Map G14-C36. 3) El Salto, La Ciénega, 25° 22’ Lat; 100° 14’ Long; Topographic Map G14-C36. 4) Guadalupe City, 25° 40’ Lat; 100° 15’ Long; Topographic Map G14- C26. 5) La Encantada, 23° 49” Lat; 99° 41’ Long; Topographic Map F14- Al7. 6) San Roque, 25° 36’ Lat; 100° 09’ Long; Topographic Map G14-C26. Oaxaca: 1) | Temazcal, 18° 15’ Lat; 96° 21’ Long; Topographic Map E14-6. 2) Teotitlan del Camino, 18° 08’ Lat; 96° 53’ Long; Topographic Map E14-6. 2544 Quintana Roo state: 1) 2) 3) 4) 5) Ejido La Uni6n, 17° 55? Lat; 887 52° Long; Topographic Map E16-4- is Km 12 Cafetal-Mahahual Road, 18° 47 Lat; 87° 45’ Long; Topographic Map E16-2-5. La Noria Ranch, 19° 40’ Lat; 88° 44’ Long; Topographic Map E16-1. San Felipe Bacalar, 18° 40’ Lat; 88° 25’ Long; Topographic Map E16-4-7. Tamalcab Island, 18° 35’ Lat; 88° 13’ Long; Topographic Map E16- 4-7. San Luis Potosi state: 1) Tamasopo Bath Resort, 21° 50’ Lat; 99° 25’ Long; Topographic Map F14-8. Tabasco state: 1) 2) Colegio Superior de Agricultura Tropical, 17° 59’ Lat; 93° 35’ Long; Topographic Map E15-8. La Venta Archaeological Zone, 18° 05’ Lat; 94° 03’ Long; Topographic Map E15-1-4. Tamaulipas state: 11) Alta Cima, 23° 03’ Lat; 99° 12’ Long; Topographic Map F14-A49. Barra del Tordo, 23° 03’ Lat; 97° 45’ Long; Topographic Map F14- B44. Ejido Conrado Castillo, 23° 57’ Lat; 99° 28’ Long; Topographic Map F14-A18. Ejido Julilo, 23° 08’ Lat; 99° 12’ Long; Topographic Map F14-A49. EI Bernal Hill, 22° 45’ Lat; 98° 35’ Long; Topographic Map F14- B52. El Cielo Ranch, 23° 05’ Lat; 99° 11’ Long; Topographic Map F14- A49. El Madrojfio, 23° 36’ Lat, 99° 13’ Long; Topographic Map F14-A29. El Tigre Hill, 24° 37° Lat; 98° 37° Long; Topographic Map G14-D62. Gomez Farias town, 23° 02’ Lat; 99° 09’ Long: Topographic Map F14-A49. Km 2 Victoria City-Matamoros City Road, 22° 02’ Lat; 99° 08’ Long; Topographic Map F14-A19. La Libertad, 23° 47’ Lat; 99° 11’ Long; Topographic Map F14-A19. Veracruz State: 1) Axocoapan, 19° 12’ Lat; 97° 00’ Long; Topographic Map E14-3. 258 2) 3) 4) Eyipantla Falls, 18° 25’ Lat; 95° 13’ Long; Topographic Map E15-1- 4. Jalapa, 19° 32’ Lat; 96° 55’ Long; 1440 m Alt; Topographic Map E14-3. Los Tuxtlas UNAM Biological Station, 18° 33’ Lat, 95° 04’ Long; Topographic Map E15-1-4. MY COTAXON Volume XLVII, pp. 259-274 April-June 1993 NEW FUNGI FROM YUCCA: PLANISTROMELLA YUCCIFOLIORUM, GEN. ET SP. NOV., ITS ANAMORPH, KELLERMANIA YUCCIFOLIORUM, SP. NOV., AND PLANISTROMELLA UNISEPTATA, SP. NOV., THE TELEOMORPH OF KELLERMANIA YUCCIGENA ANNETTE W. RAMALEY 7 Animas Place Durango, Colorado 81301 Planistromella, a new genus closely related to Planistroma, is described. The anamorph of Planistromella yuccifoliorum is Kellermania yuccifoliorum, a new species. The anamorph of Planistromella uniseptata is Kellermania yuccigena. Key words: Loculoascomycete, Coelomycete, Planistromella, Kellermania. INTRODUCTION Stagonospora gigantea Heald & Wolf (3) was described from Agave americana Linnaeus, and has been reported from several species of Yucca in the United States (2). The name has been a catch-all designation for coelomycetes from Yucca which have unappendaged, large, hyaline, septate conidia born singly on conidiogenous cells lining a thick-walled pycnidial chamber, because there were no other satisfactory described genera for their disposition. The type of S. gigantea was examined (10), and Piptarthron uniloculare Ramaley was estab- lished to accommodate some of the fungi formerly called S. gigantea. Further study and collections indicate that another anamorph species can be reliably separated from the fungi placed in S. gigantea for lack of an alternative taxon. This new anamorph, Kellermania yuccifoliorum Ramaley, has finger-shaped, 2- septate, somewhat pointed conidia that often have a small apical appendage. The teleomorphs of K. yuccifoltorum and K. yuccigena Ellis & Everhart comprise a new ascomycete genus, Planistromella, which has septate ascospores but otherwise resembles Planistroma, the teleomorph for the Pzptarthron species in which a teleomorph has been observed (9, 10, and unpublished). MATERIAL and METHODS Lactophenol-cotton blue was utilized when necessary in the study of appendage formation during conidiogenesis. 260 Cultures were obtained by suspending freshly extruded conidia in a drop of sterile tap water and spreading the drop over the surface of commercial potato dextrose agar (PDA) in a petri plate. Germinating conidia were isolated onto plates of fresh PDA and grown at room temperature in incident light. Asci were washed 3 times in sterile tap water and placed on petri plates of PDA. Growth was isolated onto petri plates of fresh PDA and grown at room temperature in incident light. Planistromella yuccifoliorum Ramaley and Kellermania yucctfoliorum Ramaley were discovered among specimens in a loan from the University of California at Berkeley (UC). Fresh K. yucctfoliorum was collected subsequently from the desert of the southwestern United States at eleven sites from Washington County, Utah, southward to San Bernardino and Riverside Counties, California, and Cocconino County, Arizona, on Yucca baccata Torrey, Y. brevifolia Engelmann, and Y. thornberi McKelvey. | Measurements were made of fungal tissue mounted in sterile tap water. Ten conidia were measured from one conidioma on each of five leaves in the eleven fresh collections. Material in two of the herbarium specimens was very limited, so only twenty-five conidia for each of those two specimens (UC 1138875, UC 1202973) were measured. An analysis of variance (ANOVA) of conidial width for all groups was prepared. The sum of squares simultaneous test procedure (SS-STP) was carried out for: (a) all collections, (b) fresh vs herbarium collections, (c) all fresh collections, (d) fresh collections from Y. brevifolia vs those from other Yucca species, and (e) fresh collections from Y. brevifolia only (Table 1) to test the significance of differences in mean conidial width for these groups. TAXONOMY PLANISTROMELLA Ramaley, gen. nov. Stromata atrocolorata, subepidermalia, immersa, erumpentia, separata vel confluentia, glabra; paries crassus, cupulatus, pars supera in epidermide hospitis affixa, ex aliquot stratts cellularum atroparietum compositus. Stratum supremum stromatis ex columnis cellularum elongatarum compositum. Loculi ascogeni in eodem stromate ac loculi conidiogent vel spermatiogeni vel ambo ezortentes. Locult ovoidet vel globost, unistrati, textura interthecio repletz, omnes ostiolati periphysibus, per columnas ex cellulis elongatis compositis separatt. Asci bitunicati, subclavatt vel prope cylindrati. Ascosporae leves, hyalinae, septatae. Typus generis: Planistromella yuccifoliorum Ramaley Stromata dark, subepidermal, immersed, erumpent, separate or confluent, glabrous; wall thick, cup-shaped, upper portion attached to the host epidermis, composed of several layers of dark-walled cells. The top of a stroma composed 261 of columns of elongated cells. Ascogenous locules developing in the same stroma as the conidial or spermatial locules or both. Locules ovoid to globose, in a single layer, filled with abundant interthecial tissue, each with a periphysate ostiole, separated by columns of elongated cells. Asci bitunicate, slightly clavate or nearly cylindric. Ascospores smooth, hyaline, septate. Planistromella yuccifoliorum Ramaley sp. nov.----- Big el ee UA wks he Stromata fusca vel atra, subepidermalia, tmmersa, erumpentia, amphigena, separata vel confluentia, glabra, 300-550 pm diam ubi separata, usque ad 450 pm elata, 1-5 locularia; paries crassus, cupulatus, pars supera in epidermide hospitis affiza, ex 6-12 stratis cellularum atroparietum compositus. Stratum supremum stromatis ex columnis cellularum elongatarum compositum. Locult ascogeni in eodem stromate ac locult conidiogeni vel spermatiogenit vel ambo exorientes, asct post conidia vel spermatia vel ambo formatt. Loculi ovorder vel globosi, untstrati, omnes ostiolati pertphystbus, per columnas ex cellulis brunneis, elongatis, aliquantum crassiparietibus compostitis separati. Asci bitunicati, subclavati, octosport, 120 x 32 pm ante prolationem ultimam et misstonem ascosporarum. Ascosporae hyalinae, ellipsoideae aspectu uno, curvatae ertremis obtuse rotundatis ubt 90 gradibus versae, 2-septatae, 34-42 x 10-16 pm. Textura intertheciu abunda, filamentosa, copiosa ubi asci vetissimi maturi, fatiscens ab apice locult versus imum. Stromata (Fig. 2, A) dark brown to black, subepidermal, immersed, erumpent, amphigenous, separate or confluent, glabrous 300-550 ym diam when separate, up to 450 pm tall, 1-5 loculate; wall thick, cup-shaped, the upper part attached to the host epidermis, composed of 6-12 layers of dark-walled cells. The top of a stroma composed of columns of elongated cells. | Ascogenous locules developing in the same stroma as a conidiogenous or spermatiogenous locule or both, asci formed after the production of conidia or spermatia or both. Locules (Fig. 1) ovoid to globose, in a single layer, each with a periphysate ostiole, separated by columns of brown, rather thick-walled, elongated cells. Asci (Fig. 2, C) bitunicate, slightly clavate, 8-spored, 120 x 32 ym before the final elongation and ascospore release. Ascospores (Fig. 2, C) hyaline, ellipsoid in one view, curved with bluntly rounded ends when rotated 90 degrees, 2-septate, 34-42 x 10-16 ym. Interthecial tissue abundant, filamentous, present in large quantity when the oldest asci are mature, breaking down at the top of the locule and proceeding downward. Substrate: Leaves of Yucca species. HOLOTYPE: USA. CALIFORNIA: San Bernardino County, Roadside 20 miles east of Baker (Hwy. 91/466), on leaves of Yucca brevifolia Engelmann, 14 April 1960, Isabelle Tavares No. 466, UC 12029738. The collection of this fungus was more than thirty years old, so cultivation was 262 mes, 4 ? Gaffe, : He OA PIECE L AUP IML ISS PRK Tei ASN Dg SANG Wh I VEE OT OSREE Fig. 1. Planistromella yuccifoliorum, detail of an ascoma. Standard line = 50 pm. not possible. Only dead tissues were available for measurement. In the living state, the ascospores and asci could be expected to be larger than the above measurements--up to 18% longer and 25% wider for the ascospores, and up to 20% longer and 26% wider for the asci (1). Anamorph: Kellermania yuccifoliorum Ramaley sp. nov.----- Fig. 3,A-D, Fig. 4, A & B. 263 Conidiomata amphigena, subepidermalia, atra, immersa, erumpentia etiam obtecta in margine per epidermidem, separata vel confluentia, glabra, 250-600 (-800) xm diam ubi separata, usque ad 600 ym elata, unilocularia, ostiolata; parties crassus, cupulatus, in eptdermide hospitis affizus, er 6-12 stratts cellularum atrorum crassiparietum compositus, pallidior versus partem interiorem, et 2-4 stratis cellularum hyalinarum. Stratum supremum contdio- matis ex columnis cellularum aurearum elongatarum compositum. Ostiolum cellulis atripartetibus cinctum, per rumpentem vel disrumpentem epidermidis hospitts in margine interiore parietis cupulati expositum. Ontogenesis conidiorum holoblastica. Conidiophora carentia. Cellulae macroconidiogenae in parietibus lateralibus et basalibus conidiomatis formatae, brevicylindratae, hyalinae, leves, omnes contdium unum acrogenum formantes. Macroconidia anguste ellipsoidea-cylindrata, extremis basalibus obtuse rotundatis, apicibus acutis et appendicibus usque ad 5 pm longis superpositis, plerumque 2-septata, 50-100 x (8-)13-14(-16) pm. Cellulae microconidiogenae in pariete supero conidiomatis et canale ostiolt erorientes. Microconidia acrogena, plus minusve cylindrata, septata, levia, hyalina, 5-10 x 2.5-4 pm. Spermatia in loculo centrali stromatis, cum vel sine conidits, vel in uno vel plus loculis in columnis verticalibus abundantibus cellularum elongatarum ad partetes laterales aliquot conidiomatum affixis formata, omnis loculus ostiolatus vel locult coalescentes et ostiolum solitarium formantes. Cellulae spermatiogenae discretae vel integratae in conidiophoro unicellulario, cylindratae vel elongatae-conicae, phialidicae, 8- 16 r 2-3.5 pm. Spermatia bacillaria, hyalina, aseptata, levia, 3-7 © 1.6-2.5 pm. Conidiomata (Fig. 3, A) amphigenous, subepidermal, dark, immersed, erumpent but a rim remaining covered by the epidermis, separate or confluent, glabrous 250-600 (-800) zm diam when separate, up to 600 ym tall, unilocular, ostiolate; wall thick, cup-shaped, attached to the host epidermis, composed of 6-12 layers of dark, thick-walled cells, lighter toward the interior, and 2-4 layers of hyaline cells. The top layer of a conidioma composed of vertical columns of golden elongated cells. Ostiole surrounded by dark-walled cells, exposed by rupture or breaking away of the host epidermis at the inner edge of the cup-shaped wall. Conidiogenesis holoblastic. Conidiophores absent. Macroconidiogenous cells (Figs. 3, A; 4, A) formed on lateral and basal walls of a conidioma, short- cylindric, hyaline, smooth, each forming a single acrogenous conidium. Macroconidia (Fig. 4, B) narrowly ellipsoid-cylindric, the base bluntly rounded, the apex more pointed and often surmounted by an appendage up to 5 pm long, mostly 2-septate, 50-100 x (8-)13-14(-16) wm. Microconidiogenous cells arising on the upper wall of a conidioma and in the ostiolar channel. Microconidia acrogenous, more or less cylindric, aseptate, smooth-walled, hyaline 5-10 x 2.5-4 pm (Fig. 3, B). Spermatia formed in the central locule of a stroma, with conidia or without, or in one or more locules in the abundant vertical columns of elongated cells attached to the lateral walls of some conidiomata (Fig. 3, D), each locule ostiolate or coalescing and forming one ostiole. Spermatiogenous cells discrete or integrated on a one-celled (usually) conidiophore (Fig. 3, C) cylindric to elongate-conical, phialidic, 8-16 x 2-3.5 ym. Spermatia bacillar, hyaline, smooth, 3-7 x 1.5-2.5 ym (Fig. 3, C). 264 Fig. 2. A, C, Planistromella yucctfoliorum. A. Diagram of stroma with two ascomata sectioned. The left ascoma is drawn in detail in Figure 1; a=ascoma, e=host epidermis, f=columnar filamentous portion of stroma, u=collapsed locule which contained conidia or spermatia or both, w=wall. C. Asci and ascospores. B. Planistromella uniseptata, asci and ascospores. Standard line = A, 175 pm; B, C, 25 pm. 265 Substrate: Leaves of Yucca species. HOLOTYPE: USA. ARIZONA: Mohave County, dirt road 0.3 miles from mile 20, Gem Acres Road turnoff from Highway 40, on leaves of Yucca brevifolia Engelmann, 15 April 1992, Annette Ramaley No. 9208, UC 1475101. Paratypes: (1) USA. CALIFORNIA: Los Angeles County, Claremont, Rancho Santa Ana Botanic Garden, leaves of Yucca baccata Torrey, 26 March 1957, Lee Bonar, UC 1138875; (2) USA. CALIFORNIA: San Bernardino County, roadside 20 miles east of Baker (Highway 91/466), on leaves of Yucca brevifolia Engelmann, Isabelle Tavares No. 466, 14 April 1960, UC 1202973; (3) USA. CALIFORNIA: Riverside County, near Cottonwood Springs, Joshua Tree National Monument, on Yucca schidigera Roezl ex Ortgies, 6 April 1952, Lee Bonar, UC 966401; (4) USA. ARIZONA: Santa Cruz County, mile 282.8 along Interstate Highway 10, leaves of Yucca thornbert McKelvey, 13 April 1992, Annette Ramaley No. 9211, UC 1475100; (5) USA. ARIZONA: Santa Cruz County, mile 283.6 along Interstate Highway 10, leaves of Yucca thornberi McKelvey, 13 April 1992, Annette Ramaley No. 9212, BPI 1112456; (6) USA. UTAH: Washington County, Visitor’s Center, just over border from Arizona along Interstate 15, on leaves of Yucca brevifolia Engelmann, 15 April 1992, Annette Ramaley No. 9207, BPI 1112455. Kellermania yuccifoliorum is distinguished from the other Kellermania species by conidial septation and the size of the apical appendage. K. anomala (Cooke) Hohnel (Fig. 4, C) and K. yuccigena (Fig. 4, E) conidia have long apical appendages (12-32 ym). However, appendages of K. anomala are often difficult to observe without phase optics, and may be absent entirely. They seem pliant, and are sometimes free from the conidia. K. yuccifoliorum conidia (Fig. 4, B) have a short apical appendage (up to 5 ym) when an apical appendage is present. K. yuccigena conidia regularly have one, approximately median septum. K. anomala conidia have two septa, and the middle cell is regularly shorter than either end cell. K. yuccifoliorum conidia have two septa, but the middle cell is generally longer than the end cells. However, K. anomala and K. yuccifoliorum conidia lacking an appendage may be indistinguishable if the middle cell length is atypical for the species (compare conidia in Fig. 4, B & C). Conidia from the fresh collections germinated readily and formed colonies with white aerial, surface, and subsurface mycelium. The reverse of the colonies from some collections was pale pink, from others, white. The majority of the mycelium of colonies of most isolates soon became gray to nearly black. Within a month, conidiomata were formed that produced typical K. yucctfoltorum conidia. Planistromella yuccifoliorum could not be cultivated to confirm its connection with K. yuccifoliorum. The connection is circumstantial but assumed because (a) closely related taxa (9, 10) have similar teleomorph and anamorph phases; (b) many collections have been made throughout the range of the host, and the unilocular representatives of the Kellermania-Piptarthron complex now known from Yucca brevifolia are P. uniloculare, K. yuccifoliorum, and K. anomala. The teleomorph for P. uniloculare, Planistroma obtusilunatum 266 Ramaley, has been confirmed experimentally (10). Two stromata with a perfect state present were found in a collection in which K. anomala was the only unilocular Kellermania-Piptarthron representative . A single ascus with mature ascospores was available. The ascospores were approximately medianly 1- septate; (c) K. yuccifoliorum was the only unilocular representative of the Kellermania-Piptarthron complex present in the collection in which P. yuccifoltorum was found. The average size of fifty conidia from the eleven fresh collections ranged from 66.7-80.1 x 12.4-14.2, X=72.1 x 13.2 wm. Conidia from the herbarium material averaged 61.4 x 10.2 wm, smaller than any of the averages for the fresh conidia. The ANOVA showed extremely significant variation among the fourteen collections (Fs=41.59, a<.001). Results of the SS-STP test for the significance of the contributions to the sum of squares among the groups (SSgroups) by sample means of particular interest are shown in Table 1. The critical value for the sum of squares (CVSS) was computed conservatively using df=13, 120. Even so, SS groups (a), (b), (c), and (e) far exceeded the CVSS, so a<<.001 ‘for these groups. Table 1. Sum of Squares Simultaneous Test Procedure Results for ANOVA of the Width of Conidia from Fresh Collections and Herbarium Collections of Kellermania yucctfoliorum. Groups Analyzed SSgroups (a) all collections 13/636 1515.6262*«« (b) fresh vs herbarium collections 1/648 1206.6935*«« (c) fresh collections only 10/539 = 275.9527 **x (d) fresh: Y. brevifolia vs other sp. 1/548 88.8552** (e) fresh: Y. brevifolia only 1/392 VIIA 33 1oeee *#*, A=.001; **, a=.01 These results show the significant contributions to the total variation by (1) the fresh vs herbarium source of the conidia [(a) and (b)], (2) the different species on which the fungus was found [(c) and (d)], and (3) the plants or site from which a collection was made--the fresh collections from Y. brevifolia were significantly different (e). Conidia from the herbarium collections averaged 14.8% shorter and 22.7% narrower than those from fresh collections, and the walls appeared fairly thick, rather than thin as in the fresh material. Both these characteristics agree with observations from living versus dead ascomycetes (1), and percent difference in length and width of living versus dead conidia is within the range of the percent difference for such measurements in ascospores. These differences account for 267 the extremely large SSgroups for conidia from fresh versus herbarium material [Table 1, (b)], and emphasize the need for caution when drawing taxonomic conclusions from such measurements, at least for taxa with large, thin-walled conidia. Planistromella uniseptata Ramaley, sp. nov.-----Fig. 2, B. Planistromellae yucctfoliorum Ramaley similis sed stromata 250-400 pm diam ubi separata, usque ad 350 pm elata, 1-4 locularia. Asct 105-120 rt 28-36 pm ante prolationem ultimam et misstonem ascosporarum. Ascosporae hyalinae, eztrema obtuse rotundata vel extremum inferius plerumque aliquantum acutum, 1-septatae, 32-48 x 13.5-17.5 pm, cellula inferior 1-1.4x longitudo cellulae superioris. Ubi asct vetissimt maturi, teztura intertheci irregulariter filamentosa, columnae_ dissoctantes et consortiones 2-3-cellulares vel unicellulares formantes. Cellulae irregulariformes, cylindratae, ellipsoidiae, ovatae, vel globosae, plerumque 10-20 xt 5-8 pum. Like Planistromella yuccifoliorum Ramaley, but the stromata 250-400 um diam when separate, up to 350 ym tall, 1-4 loculate. Asci (Fig. 2, B) 105-120 x 28-36 pm before the final elongation and release of the ascospores. Ascospores (Fig. 2, B) hyaline, the ends bluntly rounded or the lower end more often slightly pointed, l-septate, 32-48 x 13.5-17.5 ym, the bottom cell 1-1.4x the length of the top cell. When the oldest asci are mature, interthecial tissue irregularly filamentous, the columns dissociating into 2-3-celled associations or into individual cells which are irregularly shaped, cylindric, ellipsoid, ovate, or spherical, mostly 10-20 x 5-8 pm. Substrate: Leaves of Yucca elata Engelmann. HOLOTYPE: USA. NEW MEXICO: Soccoro County, roadside, west side of Highway 25, mile 105.4, on leaves of Yucca elata Engelmann, 12 April 1992, Annette Ramaley No. 9217, UC 1475102. Planistromella uniseptata was found in small amounts on leaves of Yucca elata Engelmann. The anamorph of the fungus is widely distributed in the United States on Yucca angustifolia Pursh (6), Y. filamentosa Linnaeus (6), Y. glauca Nuttall (6), Y. harrtimaniae Trelease (personal observation), and Y. whipplet Torrey (6). When a systematic search is carried out, P. uniseptata may be expected on these species as well as Y. elata. Colonies arising from asci on PDA had white aerial, surface, and subsurface mycelium with a pink reverse. After 6 weeks, conidiomata which contained typical Kellermania yuccigena conidia had formed. Anamorph: Kellermania yuccigena Ellis & Everhart---Fig. 4, D & E. 268 Kellermania yuccigena has been illustrated and described in modern terms (6, 12). A few additions must be made for a more complete description, but an entire conidioma has not been drawn. In the absence of conidia, the conidio- mata could scarcely, if at all, be distinguished from small conidiomata of K. yuccifoliorum (Fig. 3, A). Conidiomata are 250-300 (-550) 4m diam and up to 450 ym tall. The wall is thick and cup-shaped. As described by Hohnel (4), the top of a conidioma is composed of parallel filaments of elongate golden cells which are very dark-walled in the area surrounding the ostiole. Microconidia are formed on the upper conidiomatal wall and in the ostiolar channel. Conidia are up to 80 ym long, considerably longer than the 50-61ym often cited (6, 13). Part of this difference may be attributed to the irreversible shrinkage of conidia from herbarium specimens (1) which were measured. The amount of filamentous material making up the top of a conidioma is variable. Spermatia are formed in 1-4 separate locules in the filamentous portion of a stroma, or in the central locule with or without accompanying conidia; each locule has an ostiole, or locules coalesce and release spermatia from a single ostiole. Spermatiogenous cells are discrete to integrated on a 1-celled (usually) conidiophore, cylindric to elongate-conical, phialidic, and 5-16 x 2.5-3.2 pm. Spermatia are hyaline, bacillar, and 4-8 x 1.6 um. Conidia germinated readily and formed colonies with white aerial, surface, and subsurface mycelium. The reverse was pink. Within 6 weeks, conidiomata had formed which contained typical Kellermania yuccigena conidia. Planistromella uniseptate and K. yuccigena were therefore considered to be two phases of a single holomorph. DEVELOPMENT Before the host epidermis is broken, a Kellermania conidioma consists of a cup- shaped wall containing maturing macroconidia. The top of a conidioma is made up of columns of elongate golden cells bounded internally by smaller, hyaline cells. As conidiomata mature, the columns of elongate cells lengthen, microconidium formation occurs in and near the ostiolar channel, and the cells that will surround the ostiolar channel become very dark. The microconidio- genous cells look much like periphyses in perithecial ascostromata, but form microconidia as an added feature. The function of microconidia is not known. A macroconidium develops as a simple outgrowth of a holoblastic conidiogenous cell. In conidiomata potentially developing beyond macroconidium formation, columns of cells are attached well down the lateral walls of the stromata (Fig. 3, compare A, f, & D, d, t) reducing the size of the single, central locule. These conidiomata may form spermatiogenous locules, ascogenous locules, or, very rarely in Kellermania yuccigena, additional conidial locules in the columnar filamentous portion. Spermatia are formed in the central locule with or without macroconidia, in small spermatial locules that develop in the lateral upright filaments, or at both sites. Frequently a conidioma forms spermatia in a single, 269 Ane yh Fig. 3. Kellermania yuccifoliorum. A. Small conidioma. B. Microconidia. C. Spermatia and spermatiogenesis. D. Diagrams of stromata. d=columnar filamentous portion of stroma developing into an area of spermatiogenesis, e=host epidermis, f=columnar filamentous portion of stroma, o=dark-walled cells around ostiole or developing ostiole, s=site of spermatiogenesis, t=portion of stroma which may develop ascomata, u=site of conidiogenesis and spermatiogenesis, w=wall. Standard line = A, 80 pm; B, C, 27 wm; D, 200 pm. Fig. 4. A, B, Kellermania yuccifoliorum. A. Conidiogenesis and origin of apical appendage. B. Conidia. C. Kellermania anomala conidia. D, E, Kellermania yuccigena. D. Conidiogenesis and origin of apical appendage. E. Conidia. Standard line = A, D, 27 pm; B, C, E, 40 um. 271 central locule, and then develops 1-4 sites in the columnar filamentous portion where the cells become hyaline and subdivided. Spermatia adhere to the several flexuous hyphae produced at each site. Planistromella develops in pre-formed stromata which have already produced conidia, spermatia, or both. Walls of the central locule are collapsed by expansion of the ascogenous locules. The asci mature among cells of the copious centrum tissues. These elongate cells, arranged in columns, may be laterally compressed between maturing asci, but not much stretched. Dissolution of the sterile cells begins at the upper end of the locule and proceeds downward. They are still abundant as early-formed asci mature. In P. yuccifoliorum the interthecial tissues are clearly filamentous as early-formed asci mature. In P. uniseptata the interthecial tissues are irregularly filamentous, dissociating into single cells or 2-3-celled series as the asci mature. In P. uniseptata the asci elongate greatly, extending beyond the periphysate ostile for ascospore release. Probably living mature P. yuccifoliorum asci elongate similarly, but this could not be ascertained in the herbarium material which was mostly immature. DISCUSSION Beyond its similarity to Planistroma, Planistromella relationships are not clear. (a) Planistroma asci are like those of the Botryosphaeriaceae (P. F. Cannon, personal communication), as are the non-septate ascospores of Planistroma. However, the type, Botryosphaeria dothidea (Mougeot ex Fries) Cesati & De Notaris, has the Pleospora type of centrum development (7) which is not found in Planistromella or Planistroma. (b) Several ascomycetes have 2-septate asco- spores. It is improbable that the two ascospore septa are formed simultaneously in Kiessleriella ocellata (Niessl) Bose or Paraphaeosphaeria michoti (Westendorp) O. Eriksson which have brown ascospores, or Buergenerula Sydow with hyaline ascospores, because of their relationships. Leptoguignardia Miller has hyaline ascospores with two simultaneously-formed septa. None of these fungi has a multilocular stroma, and the ascostromata are separate from the conidiostromata so far as is known, though the two are found in the same material for Leptoguignardia. (c) Planistromella is also distinct from the known teleomorphs for Stagonospora (11) which belong to Leptosphaeria (7 species), Pleospora (2 species), and Didymella (1 species). These fungi have unilocular pseudothecia, pseudoparaphyses that originate above the asci and grow downward to the base of the locule, and, except for Didymella, colored ascospores that are several septate or dictyosporous. According to existing descriptions, ascospore septation is sequential, not simultaneous. The Planistromella anamorph, Kellermania yuccifoliorum, would once have been forced into Stagonospora gigantea. However, the two taxa can be separated easily. Conidia of K. yucctfoliorum are nearly always 2-septate, and commonly have a short apical appendage; Stagonospora gigantea conidia are (3-)4-5(-7) septate, and lack an apical appendage. K. yuccifoliorum forms 272 microconidia near the ostiole and in the ostiolar channel; SS. gigantea has periphysis-like cells present in these areas, but does not form microconidia. The wall of Kellermania is gray-black when young, sometimes becoming dark brown in age, whereas the wall of S. gigantea is brown. The new anamorph taxon was clearly related to Kellermania and Piptarthron by details of the life cycle and structure of the conidiomata. These genera are distinguished by the presence of a single apical appendage on Kellermania conidia. Conidial appendages vary in origin. To place the anamorph properly, it was necessary to determine whether the small conidial appendages of the taxon newly separated from S. gigantea resembled the larger conidial appendages of existing Kellermanza species. Kellermania yucctfoliorum conidia originate as simple outgrowths of the holoblastic conidiogenous cells (Fig. 4, A). The cells elongate, often with little increase in width, and in many of them the apex of the conidium becomes filled with material that cannot be distinguished from the cell wall in unstained, living cells. The conidia then increase in width, become septate, and enlarge somewhat until they are mature. The occluded apex is the ‘apical appendage’ which retains the shape of the conidial initial at the time of appendage formation. The apical cell may be truncate even when conidia are mature, and the appendage becomes distinguishable from the cell wall. Origin of the apical appendage is ‘cellular’ in the sense that it is not material extruded from a cell, but it differs from the ‘cellular’ appendages of Heteropatella, Scolecosporiella, and Chaetoconis which are apical cells drawn out into a filiform appendage. Conidiogenesis was also examined in fresh collections of K. yuccigena (Fig. 4, D) with newly-formed conidiomata. In unstained, untreated preparations, developing conidia elongated to a sharp point bounded by a normal-appearing cell wall. The lumen of the apex filled, and the cytoplasmic contents in the narrowed apex were replaced. The wall of the pointed apex was separable from the surrounding conidial sheath, but could not be distinguished from the contents replacing the cytoplasm in the attentuated apex. No septum was ever seen between the apex and the remainder of the conidium, nor did the. cytoplasmic contents retract from the apex as in Monochaettellopsis (8). During the filling of the apex, small stainable areas persisted for some time. The appendages originated as cellular projections, as do those of conidia of Monochaetiella hyparrheniae Castell (8), but the protoplasmic contents were then replaced. The origin of K. yuccigena appendages seemed identical to, though more elaborate than, the origin of K. yuccifoliorum appendages. The above results are contrary to those of an earlier report (12) which describes Kellermania yuccigena appendages as unstainable material in the apical region of the most immature subulate conidia which becomes increasingly elongated as the conidium matures. The fresh collections were put aside for later study to try to reconcile the differences. When the collections were re-examined several months later, not one sharply pointed conidium could be found with an open 273 narrowed apex. All apices were filled, making it appear that the appendage originated outside the cell wall and increased in size as the conidium developed. The reaction of the fungus to storage had obscured the true origin of the conidial apical appendage. Since the conidial appendages of the new taxon originated like those of K. yuccigena, it was designated a species of Kellermania, and not considered to be a new, related genus. The close relationship of Kellermania and Piptarthron was recognized by Hohnel when he validated the genus Piptarthron (5). The teleomorphs for the two genera are also much alike, but differ consistently in the species studied to this point. Typical aseptate Planistroma ascospores, differing in size, have been found in three of the four Piptarthron species whose teleomorphs have been observed (9 and unpublished). Planistroma obtusilunatum Ramaley has asco- spores that are shaped differently from the other Planistroma species (10), but they are aseptate. The Planistromella teleomorphs for K. yuccigena, and K. yucctfoliorum differ from Planistroma by their possessing septate ascospores. The same is true for the teleomorph of K. anomala. The single observed ascus of that species contained medianly 1-septate ascospores. A simplification would be to include both anamorph genera in the earlier genus, Kellermania, and both teleomorph genera in Planistroma. However, the generic limits are remarkably consistent as they stand. ACKNOWLEDGMENTS I must express my appreciation to the curator of the Herbarium, University of California, for making materials housed at that institution available to me. My thanks also to Francis A. Uecker for the corrections and valuable suggestions made in his review of the manuscript and his correcting of the Latin diagnoses, and to Deborah Berrier for her advice, comments, and encouragement with the statistics. Literature Cited 1. Baral, H. O. 1992. Vital versus Herbarium Taxonomy: Morphological Differences Between Living and Dead Cells of Ascomycetes, and Their Taxonomic Implications. Mycotaxon 44: 333-390. 2. Farr, D. F., Bills, G. F., Chamuris, G. P., and Rossman, A. Y. 1989. Fungi on Plants and Plant Products in the United States. The American Phytopathological Society, 3340 Pilot Knob Road, St. Paul, Minnesota 55121, U.S.A. 3. Heald, F. D., and Wolf, F. A. 1911. New Species of Texas Fungi. Mycologia 3: 522. We 4. Hohnel, F. v. 1915. Fragmente zur Mykologie no. 900. Uber die Gattung Kellermania Ellis and Everhart. Sitzungsber. Kais. Akad. Wiss. Wien, Math-Naturwiss. cl. Abt. 1, 124: 82-84. 274 jae 13. Hohnel, F. v. 1918. Fungi imperfecti. Beitrage zur Kenntnis derselben. Hedwigia 60: 129-209. Morgan-Jones, G., Nag Raj, T. R., and Kendrick, B. 1972. Genera Coelomycetarum. VI. Kellermania. Can. J. Bot. 50: 1641-1648. Parguey-Leduc, A. 1966. Recherches sur L’Ontogénie et L’Anatomie Comparée des Ascocarpes des Pyrénomycétes Ascoloculaires. Ann. Sci. Naturelles, Bot., 12 ser. 7: 505-690. Punithalingam, E. 1992. The Nuclei and Conidial Appendages in Monochaetiellopsis species (Coelomycetes). Nova Hedwigia 54: 255-267. Ramaley, A. W. 1991. Fungi of Yucca baccata. 1. Piptarthron pluriloculare and Its Teleomorph, Planistroma yuccigena. Mycotaxon 42: 63-75. Ramaley, A. W. 1992. Fungi From Yucca baccata 2. Planistroma obtusilunatum sp. nov., and Its Anamorph, Piptarthron uniloculare sp. nov. Mycotaxon 45: 449-460. Sivanesan, A. 1984. The Bitunicate Ascomycetes and Their Anamorphs. A. R. Gantner Verglag K. G., F19490 Vaduz. Sutton, B. C. 1968. Kellermania and Its Generic Segregates. Can. J. Bot. 46: 181-196. Sutton, B. C. 1980. The Coelomycetes. Commonwealth Mycol. Inst., Kew, Surrey, England. MY COTAXON Volume XLVI, pp. 275-281 April-June 1993 4-O-METHYLLIVIDIC ACID, A NEW LICHEN DEPSIDONE JOHN A. ELIX and DEBRA A. VENABLES Department of Chemistry, The Faculties, Australian National University, GPO Box 4, Canberra, ACT, 2601, Australia ABSTRACT : The depsidone, 4-O-methyllividic acid (8-hydroxy-3,4-dimethoxy-1-(2'-oxoheptyl)-6-pentyl- 11-oxo-11H-dibenzo|[b,e][1,4]dioxepin-7-carboxylic acid (11) has been identified in the lichens Hypocenomyce foveata, Hypotrachyna immaculata, H. livida and H. osseoalba and was synthesized and characterized. Introduction Minor biosynthetic variations observed within a lichen genus or a group of related species can produce a large number of structurally related metabolites or a chemosyndrome (Culberson & Culberson 1976). The structural variations within a chemosyndrome take several forms and in the orcinol depsidones (1) these include : i. variations in the length of the polyketide derived side chain R!. ii. the degree of oxidation of this side chain (i.e. CH2COR' or CH2CH)R’). iii. the degree of methylation of the phenolic groups (i.e. R?, R8 = Me or H). iv. the presence/absence of an additional phenolic hydroxy group (or derived methoxy group) in position 4. Lichens of the genus Hypotrachyna, more particularly H. osseoalba (Vain.) Park & Hale, H. livida (Taylor) Hale and H. immaculata (Kurok.) Hale produce such a chemosyndrome, the following members of which have been identified previously; norcolensoic acid (2), colensoic acid (3), hydroxycolensoic acid (4), methoxycolensoic acid (5), physodic acid (6), 3-O- methylphysodic acid (7) [unfortunately this compound was originally termed 4-O-methylphysodic acid, using a now-obsolete numbering system] oxyphysodic acid (8) and lividic acid (9) (Chester & Elix 1981; Culberson 1966; Djura, Sargent et al. 1977a, 1977b; Elix & Engkaninan 1976; Elix & Johnston 1991). Materials and Methods The lichen fragments were freed as far as possible from obvious organic substrate material and extracted with warm acetone for thin-layer chromatography (TLC) or with warm methanol for high performance 276 O Ry O ORg O RgO COoH Rg Re 1 O Ry R30 Rg Ry R3 R4 Re R7 2 Cs5Hi1 H H CsHi; 4H 3 CsHi1 Me H Csi ta abe 4 Cs5H11 Me OH Cs5H11 H 3) Cs5H11 Me OMe CsH11 H 6 CH2COCs5H;; H H Czlivin eee | CH»COCsH;; Me H cH 8 CH»COCsH;; H OH Cs5H}] H 9 CH2COC5H;; Me OH Cs5H11 H OM H 10 CH2COCS5H Me € CsH)1 le} CH2COC5H 1 H OMe Cs5H)1 CH»C.6Hs 12 #CHy»COCsH;; Me OMe Cs5Hj1 CH»2Ce6Hs 277 liquid chromatography (HPLC). Compounds were identified by TLC using the methods standardized for lichen products (Culberson & Ammann, 1979; Culberson & Johnson 1976, 1982; Elix, Johnston & Parker, 1987) and by HPLC with retention index values (RI) calculated from benzoic acid and solorinic acid controls (Feige, Lumbsch, Huneck & Elix, 1993). HPLC used a Spectra System, a Perkin-Elmer HS-5C18 column and a spectrometric detector operating at 254 nm with a flow rate of 1 ml/min. Two solvent systems were used: 1% aqueous orthophosphoric acid and methanol in ratio 3:7 (A) and methanol (B). The run started with 100% A and was raised to 58% B within 15 min., then to 100% B in 30 min. and isocratic elution in 100% B for a further 10 min. The following lichens were studied: Hypocemomyce foveata Timdal AUSTRALIA. Australian Capital Territory. On trunk of Eucalyptus macrorhyncha in dry Eucalyptus woodland, Canberra Nature Park, Bruce Ridge, 35°15'S, 149°05'E, J. A. Elix 27485, 261.1991 (ANUC). New South Wales. At base of trunk of Eucalyptus macrorhyncha in dry Eucalyptus woodland, Great Dividing Range, Tuena-Binda road, 30 km north-west of Crookwell, 34°13'S, 149°20'E, J. A. Elix 27483 , 12.ix.1990 (ANUC). Hypotrachyna immaculata (Kurok.) Hale AUSTRALIA. Australian Capital Territory. On Leptospermum shrubs in Eucalyptus woodland, trail to Booroomba Rocks, 11 km south-west of Tharwa, 35°35'S, 149°02'E, J. A. Elix 4027, 16.ii.1977 (ANUC). REPUBLIC of SOUTH AFRICA. Cape Province. On old oaks, Stormsrivier, Distr. Humansdorp, O. Almborn 4133, 28.viii.1953 (ANUC). Hypotrachyna osseoalba (Vain.) Hale & Park AUSTRALIA. Queensland. On Alphitonia in disturbed rainforest, Lannercost State Forest, Old Mill Road, 36 km WSW of Ingham, 18°45'S, 145°49'E, J. A. Elix 15609, 19.vi.1984 (ANUC). NEW ZEALAND. South Island. On kahikatea in swampy forest, Little Ditch, 8 km north-east of Waimangaroa, J. A. Elix 7273, 26.ii.1980 (ANUC). The following authentic compounds have been synthesised or isolated previously: norcolensoic acid (2) (Chester & Elix, 1981); colensoic acid (3) (Djura & Sargent 1976); hydroxycolensoic acid (4) and methoxycolensoic acid (5) (Djura, Sargent et al. 1977a,b); physodic acid (6), 3- O-methylphysodic acid (7) and oxyphysodic acid (8) (Elix 1975) and lividic acid (9) (Elix & Engkaninan 1976). 278 Synthesis of 4-O-Methyllividic acid (10) Benzyl 4-O-Methyllividate (11) Benzyl 4-methoxyphysodate (12) (Elix & Engkaninan 1976) (17.5 mg, 29.6 umole) was treated with an excess of ethereal diazomethane for a period of 6 hours. The ethereal solution was treated with glacial acetic acid (2 drops) and washed in turn with saturated sodium bicarbonate solution, brine, and dried (MgSOq4). The residue obtained on evaporation of the solvent was crystallized from light petroleum to yield benzyl 4-O- methyllividate (11) (11.6 mg, 65%), as colourless needles m.p. 109.5-111° (Found: mol. wt. 604.2670. 12C35!H49!6Oo, require mol. wt. 604.2672). 1H n.m.r. (CDCl3) 6 0.78-0.91, m, CH2CH3; 1.21-1.62, m, CH2(CH2)3CH3; 2.55, t, COCH2CHy9; 3.26, t) ArCH2CH 9; 3.76, s, 2H, ArCH2CO; 3.89, s, OCH3; 5.37, s, COsCH); 6.56, 6.75; 2s, H2, H9; 7.41, m, C6Hs; 11.19, s; OH. Mass aeheeatss m/z 605 (10%), 604 (M, 29), 513 (3), 469 (6). 4-O-Methyllividic Acid (10) A solution of benzyl 4-O-methyllividate (11) (11.6 mg, 19. umole) in ethyl acetate, containing 10% palladium-on-carbon (3.9 mg) was stirred in an atmosphere of hydrogen at room temperature for 1.5 hours. The reaction mixture was filtered to remove the catalyst and the catalyst washed with ethyl acetate. The combined filtrate was then evaporated under reduced pressure and the residue crystallized from dichloromethane/light petroleum, to afford 4-O-methyllividic acid (10) (5.0 mg, 51%) as colourless needles, m.p. 156.5-159° (Found: mol. wt. 470.2305. 12C271H34!6O7, requires mol. wt. 470.2305). 'H n.m.r. (CDCl3) 6 0.88-0.94, m, CH2CH3; 1.24-1.65, m, CH?2(CH2)3CH3; 2.57, t, COCH2CH?; 3.40, bt, ATCH2CH?; 3.92, s, ArCH2CO; 3:913.96, 25, OCH376.53,16776; 2s, H2, HI; 112-14, b, OL es Mass. spectrum m/z 471 (32%), 470 (M, 100), 293 (16), 292 (42), 277 (38), 263 (16), 221 (12), 207 (14). Results and Discussion The lichen Hypotrachyna immaculata was shown by HPLC and TLC to produce nine phenolic metabolites, eight of which corresponded to known lichen substances. These included the common cortical depsides atranorin and chloroatranorin, and the orcinol depsidones norcolensoic acid (2), colensoic acid (3), physodic acid (6), 3-O-methylphysodic acid (7), oxyphysodic acid (8) and lividic acid (9), all of which were readily identified by comparison with authentic materials (Figure 1). Subsequently we have undertaken the synthesis of 4-O-methyllividic acid (10), and found that the TLC and HPLC behaviour of this compound to be identical with that of the eighth phenolic substance present in H. immaculata. This new depsidone has also been detected in several other lichens which contain the colensoic acid - lividic acid chemosyndrome ie chemosyndrome including Hypocenomyce foveata Timdal and Hypotrachyna osseoalba (Vain.) Park & Hale. 4-O-Methyllividic acid (10) was prepared by hydrogenolysis of benzyl 4-O-methyllividate (11), which in turn was obtained by methylation of benzyl 4-methoxyphysodate (12) (Elix & Engakinan 1976). The standardized chromatographic data for these compounds are listed in Table 1. 7 (ppb: 13 14 Figure 1. H.p.l.c. trace of Hypotrachyna immaculata (J. A. Elix 4027 in ANUC). S = internal standard. ACKNOWLEDGEMENTS We thank the Australian Research Council for generous financial support of this project; Mr. G. Deeble (HPLC), Ms. Caroline Barclay (HPLC) and Ms. S. A. Elix (TLC) who assisted in the determination of the chemistry of many specimens. A Commonwealth Postgraduate Award to (D.V.) is gratefully acknowledged. 280 Table 1: Standardized Chromatographic Data for Hypotrachyna Metabolites Standard Rg values (x 100) were determined in three independent t.l.c. solvent systems: (A) toluene / dioxane / acetic acid (180 : 45 : 5); ( (B*) hexane / t-butyl methyl ether / formic acid (140 : 72 : 18); (C) toluene / acetic acid (170 : 30). HPLC retention index (RI) values (x 100) are relative to solorinic acid and benzoic acid. Compound Rr. A B* RI Norcolensoic Acid (2) 28 27] 31 67.0 Colensoic Acid (3) 4] 66 47 85.0 Hydroxycolensoic Acid (4) 39 50 42 63.6 Methoxycolensoic Acid (5) 44 61 47 77.0 Physodic Acid (6) 25 35 18 63.3 3-O-Methylphysodic Acid (7) 39 45 42 73.3 Oxyphysodic Acid (8) 15 34 13 Sia Lividic Acid (9) o2 37 31 56.8 4-O-Methyllividic Acid (10) 41 46 20 68.6 Atranorin (13) (Standard) 75 ris 79 76.3 Chloroatranorin (14) (Standard) 74 73 81 80.6 LITERATURE CITED Chester, D. O. & Elix, J. A. (1981). New metabolites from Australian lichens. Aust. ]. Chem. 34: 1507-1511. Culberson, C. F. (1966). The structure of a new depsidone from the lichen Parmelia livida. Phytochemistry 5: 815-818. Culberson, C. F. (1972). Improved conditions and new data for the identification of lichen products by a standardized thin-layer chromatographic method. J. Chromatogr. 72: 113-125. 281 Culberson, C. F. & Ammann, K. (1979). Standardmethode zur Dtinnschicht- chromatographie von Flechtensubstanzen. Herzogia 5 : 1-24. Culberson, C. F. & Culberson, W. L. (1976). Chemosyndromic variation in lichens. Syst. Bot. 1: 325-339. Culberson, C. F. & Johnson, A. (1982). Substitution of methyl tert.-butyl ether for diethyl ether in the standardized thin-layer chromatographic method for lichen products. J. Chromatogr. 238: 483-487. Djura, P. & Sargent, M. V. (1976). Depsidone Synthesis. VI. Colensoic acid. Aust. J. Chem. 29: 1069-1077. Djura,-Pisargent, aM V7 Bip) JAG Engkaninan, WU) Huneck, 5.) & Culberson, C. F. (1977a). Depsidone synthesis. VII. Isolation and structure determination of hydroxy- and methoxy-colensoic acids. Synthesis of methyl methoxy-O-methylcolensoate. Aust. J. Chem. 30: 599-607! Djura, P., Sargent, M. V., & Clark, P. D. (1977b). Depsidone synthesis. X. Methoxy- and hydroxy-colensoic acids. Aust. J. Chem. 30: 1545-1551. Elix, J. A. (1975). 2'-O-Methylphysodic acid and hydroxyphysodic acid: two new depsidones from the lichen Hypogymnia billardieri. Aust. J. Chem. 28: 849-858. Elix, J. A. & Engkaninan, U. (1976). The structure of lividic acid, a depsidone from the lichen Parmelia formosana. Aust. J. Chem. 29: 203- 207. Elix, J. A. and Johnston, J. (1991). Additional lichen records for New Zealand. 3. Further representatives in the family Parmeliaceae. Australasian Lichenological Newsletter 28: 7-10. Elix, J. A., Johnston, J. & Parker, J. L. (1987). A Catalogue of Standardized Thin Layer Chromatographic Data and Biosynthetic Relationships for Lichen Substances (Aust. Nat. University, Canberra). Elix, J. A., Johnston, J. & Parker, J. L. (1988). A Computer Program for the Rapid Identification of Lichen Substances. Mycotaxon 31: 89-99. reice,. G. 5.) Lumbsch, H: 17, buneck\S> &jElix J. A.” (1993) The identification of lichen substances by a standardized high-performance liquid chromatographic method. J. Chromatogr. in press. © 4 ry ; wit? me " 4 1! re ‘) ae ey. by ye ¥ pr an AY a. : i leks ; zi ae tin < } Lind gael 8 ia ‘ Be aA Mie ?, PGR ities a ‘ Sastre a: yo i. pine One, oe ‘ | ia me tar Win oe * | A at ve Cate. i (i x i ai yet ‘} ¥ Ne ba i J r 4 ee » A : ‘% ‘ ‘ ‘oa Aes ere kat, ye Pty fay! Lane 7 i te aya ee A) ih PN aN tt r ‘nee. ie j base Hh Ath AON suai 1 ate Be Ss if cans AN, REP ek ee ous Oe aaa NG aicuibt ceetiea \ vate pea ie ree ae me Hes . ( es i. = we "; my Se Ne EAD A “<< # : , : 4 yee Ft, | ioee i abet iat ue ae Peal noel big : ae tier tat etd By | Baey: ae pie i btn ie lige het aby aes ie | hs hes as eters mien ras Maer a Peat ie . - ie” | Oe ee a peaeoc nea Cath dip BO te mi ee a, ‘i r adh , ’ a 4 ; 4 Jat : a pe 5 at] et: zbh al I ' Vict g 4 ie } phe “4c | { eae “rae hae Pe ee COR) wt Sak iain # de it - > | “A ae ne aa; 7 | ase Bias Miisuntay od 4) oo Ms 4%, ‘ ay Ap a | i 4 | , ; ' = te - cs p dee L Aen Skeets, Cae ec ani! eit ' ae ‘ 4 Py hu | | Oui rates! if Breet) Perk ect ae ; as ce ie a | ring Oa : of, sol OE, Pte een nee vi Medan jig ig x | Se ae oe Meee Tee CLAN 7 es yr SITU “% et eee ee , | ts nes tae ue 7 . |__| age ar ape cao Spanien ans ay » 3 7 cm gicitstiaes a ets a a 7 oe ae + hid 6 ale espe See ae (1 ik ? ’ Ay _ ‘ rit fe pias ¥ .! are Dra 2 hi e 6 "hee : OP na eee eT > ’ , A ao ih ete 4 Mus: ie Shake ie et ya ares il Ge AE EON et RRC ent a GE a iy 26 Ea ls part beset aie hee ean ee a Fay bala ayes | ‘ “Vi ies ae j Ne iy hee x sith = Pe 7 ai ry tm es ‘4 °, sau 2a 7 Me a 9 + f view oe = 1h ee) ae : oe ] eR ee | a a > tH ; : a aie ' : 7 Mi hig ; 7 * py ra bit { MY COTAXON Volume XLVII, pp. 283-284 April-June 1993 ANAEROMYCES, AN EARLIER NAME FOR RUMINOMYCES Y. W. HO Department of Biology, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia. D. J. S. BARR Centre for Land and Biological Resources Research, Central Experimental Farm, Research Branch, Agriculture Canada, Ont. Canada. N. ABDULLAH and S. JALALUDIN Department of Animal Science, Universiti Pertanian Malaysia, 43400 Serdang, Selangor Malaysia. ABSTRACT Ruminomyces Ho is placed into synonymy with Anaeromyces Breton and comparison is made between A. mucronatus and A. elegans (Ho) Ho comb. nov. There were two taxa of anaerobic polycentric fungi described in 1990 from the rumen of cattle (Bos taurus). One was from France and the other from Australia. The one from France was named Anaeromyces mucronatus Breton while the other from Australia was called Ruminomyces elegans Ho. In our opinion the two generic diagnoses are essentially similar and the two genera are synonymous. Thus, there is no justification for retaining them as two separate genera. The generic name Anaeromyces has priority as it was published July 19, 1990 whereas Ruminomyces was published August 15, 1990. However, the species are morphologically distinct and should be retained as separate taxa. The following combination is therefore proposed: Anaeromyces elegans (Ho) Ho comb. nov. = Ruminomyces elegans Ho (Ho etal. 1990). Mycotaxon 38 : 398 - 404. 284 Although the morphology and size of the sporangia and zoospores of A. elegans and A. mucronatus are somewhat similar, the rhizomycelia of the two species are clearly different. The rhizomycelium of A. elegans is composed of much- branched, narrow to wide hyphae ranging from 0.5 - 10 um wide. Many of the wide hyphae are sausage-shaped with constrictions at regular or irregular intervals. Some hyphae also produce lobed or bead-like structures, 1.5 - 14.5 um wide x 1.5 - 12.5 um high. Frequently, fine pegs (about 0.5 um wide) are produced from the centre of these lobed or bead-liked structures. The characteristics of these structures appear to be similar to the penetrating structures “appressoria" produced by anaerobic fungi colonising grass and straw fragments in the rumen of cattle and swamp buffalo (Ho et al/., 1988a, b; 1991). The hyphae of A. mucronatus, though with constrictions, do not possess the lobed or bead-like structures. Although the difference may eventually prove to be inconsequential, in our opinion, it is preferable to retain the two species until the cultures are compared under similar growing conditions and analysed by molecular analysis. REFERENCES Breton, A., Bernalier, A., Dusser, M., Fonty, G., Gaillard-Martinie, B. & Guillot, J. (1990). Anaeromyces mucronatus nov. gen., nov. sp. A new strictly anaerobic rumen fungus with polycentric thallus. FEMS Microbiology Letters 70,177 - 182. Ho, Y.W., Abdullah, N. & Jalaludin, S. (1988a) Penetrating structures of anaerobic rumen fungi in cattle and swamp buffalo. Journal of General Microbiology 134, 177 - 181. Ho, Y. W., Abdullah, N. & Jalaludin, S. (1988b), Colonization of guinea grass by anaerobic rumen fungi in swamp buffalo and cattle. Animal Feed Science and Technology 22, 161 - 171. Ho, Y. W., Bauchop, T., Abdullah, N. & Jalaludin, S. (1990). Ruminomyces elegans gen. et sp. nov., a polycentric anaerobic rumen fungus from cattle. Mycotaxon 38, 397 - 405. Ho, Y. W., Abdullah, N. & Jalaludin, S. (1991). Fungal colonization of rice straw and palm press fibre in the rumen of cattle and buffalo. Animal Feed Science and Technology 34, 311 - 321. MY COTAXON Volume XLVII, pp. 285-293 April-June 1993 A NEW SPECIES OF PIROMYCES FROM THE RUMEN OF DEER IN MALAYSIA Y.W.HO Department of Biology, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia. D.J.S. BARR Centre for Land and Biological Resources Research, Central Experimental Farm, Research Branch, Agriculture Canada, Ottawa, Ont. Canada. N. ABDULLAH, S. JALALUDIN Department of Animal Science, Universiti Pertanian Malaysia, 43400 Serdang, Selangor, Malaysia. H. KUDO Tropical Agriculture Research Centre, Tsukuba, Ibaraki, Japan. ABSTRACT Piromyces minutus, a new species of anaerobic fungus from the rumen of deer is described and illustrated. The fungus is characterised by small sporangia, each with a wide apical discharge pore, persistent sporangial wall and straight, unbranched main rhizoid terminating in a sparsely-branched rhizoidal system. The name Piromonas communis Liebetanz was established in 1910 for a uniflagellated rumen organism presumed at that time to be a flagellated protozoan. Later, Orpin (1977) discovered a rumen fungal isolate with a vegetative phase which he associated with Liebetanz’s Piromonas communis, but his isolate was clearly a Chytridiomycete. Because it is not possible to know the true nature of the organism Liebetanz observed, Gold et al. (1988) established a new 286 generic name, Piromyces, for Orpin’s fungus and the description was later emended by Li et al. (1990). The name Piromonas could perhaps have been validated and retained as it had been used in the literature up to 1988. However, Piromyces is now generally accepted. There are two other species of Piromonas that need mention. These are Piromonas minima Liebetanz and Piromonas maxima Liebetanz, but we regard these as ambiguous names. The descrip- tions were based entirely on zoospore size and morphology (Liebetanz, 1910), and although it is possible that these were chytrids, it is net possible to verify or relate them to any known rumen fungal species. At present, four species have been formally described and included in the genus Piromyces. They are P. communis Gold (Gold et al., 1988, = Piromonas communis Sensu Orpin, 1977), P. mae Li (Li et al., 1990), P- dumbonica Li (Li et al., 1990) and P. rhizinflata Breton (Breton et al., 1991). Recently, two morphological forms of Piromyces were isolated over a two-month period from the rumen contents of Sika or Japanese deer (Cervus nippon) fed guinea grass (Panicum maximum) in Malaysia. One of the morphological forms was similar to P. communis but the other had conspicuously smaller sporangia with persistent wall following release of zoospores through a wide apical pore and is thought to be a new species as it is distinctly different from the four Piromyces species described. The method for isolating the fungi from the rumen contents was similar to that described by Ho & Bauchop (1991) except that glucose agar instead of cellulose agar was used in the roll tubes. The media for maintaining and culturing the isolates were glucose sloppy agar, cellulose sloppy agar and liquid medium with Whatman No. 1 filter paper or rice straw as carbon source (Ho & Bauchop, 1991). Piromyces minutus Ho sp. nov. Figs. 1-14 Thallus monocentricus eucarpicus. Sporangium omnino endo- genum, plerumque parvum denique ellipsoidale vel pyriforme vel globosum, 8-25 um latum, 8.5-28 um longum, unico poro apicali emiitenti lato et pariete sporangli persistenti, sed interdum grandius denique globosum vel parum irregulare, 40-80 um diametro, 1-2 (uno vel duobus) poris emittentibus. Axis rhizoidei singularis, aliquando 2-4 (duo vel quatuor). Rhizoideum principale sporangiorum parvorum nonramosum, systeme rhizoidali parce ramoso terminans. Zoosporae globosae 5.7-7.5 um diametro, uno interdum duobus raro quatuor 287 flagellis usque ad 31 um longis. Sporangia perdurantia ignota. Species obligate anaerobia. Thallus monocentric and eucarpic consisting of a sporangium and filamentous rhizoidal system. Sporangia strictly endogenous, predominantly small, ellipsoidal, pyriform or globose, 8 - 25 um wide x 8.5 - 28 um long with one wide, apical discharge pore and persistent sporangial wall but occasionally larger, globose or slightly irregular, 40 - 80 um diameter with one or two discharge pores. Rhizoidal axis single, occasionally two to four. Main rhizoid of small sporangia usually unbranched, terminating in a sparely-branched rhizoidal system. Zoospores globose, 5.5 - 7.5 um in diameter with one flagellum, occasionally two, rarely four, up to 31 um long. Resting spores unknown. Obligate anaerobe. From rumen contents of Sika or Japanese deer (Cervus nippon), Universiti Pertanian Malaysia, Serdang, Malaysia, 12 May, 1992, Y.W. Ho and D.J.S. Barr, D2 (UPM) holotypus. The zoospores of P. minutus are globose (Fig. 1), 5.5- 7.5 um in diameter when actively moving or stationary but occasionally irregular and amoeboid just before encystment. They are usually monoflagellated, occasionally biflagellated and rarely quadriflagellated. The flagellum is up to 31 um long including a short whiplash. Shed flagella frequently possess a bead-like structure (0.5-1.5 umin diameter) at one end. Similar flagellar structure has been reported by Barr et al. (1989) and Ho et al. (1993). Germination of zoospores produces strictly endogenous thallus development in all the four types of culture media used and in cultures of different ages, including old cultures 4-5 days old. The encysted zoospore expands uniformly and produces a germ tube (Fig. 2) which develops into a main rhizoid. Two types of rhizoidal systems are formed. In one type the main rhizoid is often straight and unbranched for some length (up to 225 um) before branching to form a sparse rhizoidal system (Fig. 3). It is narrow, 1.5 - 4.5 um in diameter, usually quite even over its length but occasionally with swellings and tightly constricted points (Figs. 4). The ends of the rhizoids are straight with moderately blunt tips. Usually a single main rhizoid is produced but occasionally two or more are formed (Figs. 5-6). This type of rhizoidal system is predominantly formed in all the four culture media used. In the other type of rhizoidal system, the main rhizoid is much wider, 5.5 - 11.5 um in diameter, sometimes coiled and more densely branched (Figs. 7-9). The ends of the rhizoids are usually straight Fig. 1. Zoospore of Pminutus. Fig. 2. A germinating zoospore. Fig. 3. Rhizoidal system showing straight and unbranched main rhizoid terminating in sparsely-branched rhizoids. Diameter of the main rhizoid is even without constrictions. Fig. 4. Main rhizoid with swellings and constrictions. Figs. 5-6. Rhizoidal system with two and three main rhizoids respectively. Bar = 20um. 289 and moderately blunt but occasionally they are rounded and slightly expanded resembling small pegs (Fig. 10). At maturity, the rhizoidal system is separated from the sporangium by a septum at the neck (the point between the sporangium and main rhizoid) or at the base of the sporangium. The sporangium develops from the expansion of the encysted zoospore. Sporangium shape is variable. Small sporangia are predo- minantly formed. They are mostly ellipsoidal to pyriform, 8 - 25 um wide x 8.5 - 28 um long, occasionally globose. They possess the rhizoidal system with long, straight main rhizoid terminating with sparse branches. Zoospores are discharged following dissolution of a wide apical portion of the sporangial wall (Fig. 11). After discharge, a persistent cup-shaped portion of the sporangial wall with a wide apical pore or opening remains (Fig. 12). In all cultures, there are invariably a small number of larger sporangia which are globose, up to 80 um in diameter, or occasionally slightly irregular. These larger sporangia frequently possess a denser rhizoidal system and occasionally with a coiled main rhizoid. Zoospore discharge is through one or two wide pores or openings (Fig. 13). The sporangial wall is smooth (as seen after zoospore release), two-layered and possibly thicker than in other species of Piromyces. There is an outer layer that is continuous with the main rhizoidal wall and an inner layer continuous with the septum. In sporangia following discharge, the two layers are occasionally separated (Fig. 14), and the septum can be seen as part of the inner layer. The persistence of the wall following discharge can be considered a distinctive characteristic of this species but the configuration of the layers, and continuity with the septum, cannot be ascertained until more species of both aerobic and anaerobic chytrids are studied with transmission electron micros- copy. There are some differences in septum morphology between different species of chytrids. Powell (1974) reported that in the aerobic chytrid Entophylyctis variabilis Powell, a transverse septum forms by centripetal accumulation of material on the walls of the rhizoids where they join the sporangium (i.e. the port). In Triparticalcar arcticum Barr there is a ring of material, different in composition to the wall, that partially closes the port (Barr, 1970). In Spizellomyces spp. the neck port is open (Barr, 1984). In contrast, Heath et al. (1983) reported that in Neocallimastix frontalis a septum is laid down at sporogenesis, and that the septum is continuous with the sporangial wall. A similar continuity of septum and inner wall can be seen in the cellulolytic, aerobic chytrid Rhizophlyctis rosea (de Bary & Woronin) Fischer 290 Figs. 7-9. P. minutus with wide main rhizoid and more densely- branched rhizoidal system. The main rhizoid is sometimes coiled. Note the ends of the rhizoids are straight and moderately blunt. Fig. 10. Rhizoidal system with densely-branched rhizoids and tips of lateral rhizoids are rounded and slightly expanded. Bar = 20 um. Fig. 11. Zoospores are released following the dissolution of a wide apical portion of the sporangial wall. Fig. 12. A persistent cup- shaped portion of the sporangium wall with a wide apical pore or opening after discharge of zoospores. Fig. 13. An empty sporangium with two wide pores or openings (arrows) through which zoospores are discharged. Fig. 14. An empty sporangium after release of zoospores showing a two-layed wall. The inner wall including the septum (arrows) is detached from the outer wall which is continuous with the main rhizoidal wall. Bar = 20um. 292 (unpublished). The septum in E. variabilis has plasmodesmata whereas the septa in N. frontalis and R. rosea apparently do not. Further studies of walls and septa in the Chytridiomycetes may reveal homologies between the aerobic and anaerobic rumen taxa. The presence of predominantly small sporangia each with a wide apical pore or opening is the conspicuous feature of P. minutus that distinguishes it from the other four species. In P. communis the zoospores are released following overall dissolution of the sporangial wall. For P. mae there are papillae that are thought to be products of localised wall softening possibly similar to the more complex papillae in other Spizellomycetales. Although rhizoid morphology is highly variable within species of Chytridiomycetes, the straight, unbranched main rhizoid terminating in a sparsely-branched rhizoidal system is very characteristic of P minutus. This species also has strictly endogenous sporangium which separates it from P. communis. All these charac- teristics separate P. minutus from the other Piromyces species described and thus require it to be assigned to a new species. Other isolates examined: Five other isolates obtained from the rumen contents of Timorensis deer (Cervus timorensis) at Universiti Pertanian Malaysia, Serdang, Malaysia, November - December, 1991, were also examined and their morphological characteristics conform to the above description. ACKNOWLEDGEMENTS We thank Dr. J. Bisset for kindly providing the Latin translation of the diagnosis, and Professor A. Nawawi for reviewing the manuscript. We would also like to extend our thanks to Dr. M.K. Vidyadaran for providing the Sika and Timorensis deer. REFERENCES Barr, D.J.S. (1970). Phlyctochytrium arcticum n. sp. (Chytridiales) morphology and physiology. Can. J. Bot. 48, 2279-2283. Barr, D.J.S. (1984). The classification of Spizellomyces, Gaertnerio- myces, Triparticalcar and Kochiomyces (Spizellomycetales, Chytridiomycetes). Can. J. Bot. 62, 1171-1201. Barr, D.J.S., Kudo, H., Jakober, K.D. & Cheng, K.-J. (1989). Morphology and development of rumen fungi: Neocallimastix sp., Piromyces communis and Orpinomyces bovis gen. nov., sp. nov. Can. J. Bot. 67, 2815-2824. is, Breton, A., Dusser, M., Gaillard-Martinie, B., Guillot, J., Millet, L. & Prensier, G. (1991). Piromyces rhizinflata nov. sp., a strictly anaerobic fungus from faeces of the Saharian ass : a morpho- logical, metabolic and ultrastructural study. FEMS Microbiol. Lett. 82, 1-8. Gold, J.J., Heath, |.B. & Bauchop, T. (1988). Ultrastructural description of a new chytrid genus of caecum anaerobe, Caecomyces equi gen. nov. sp. nov., assigned to the Neocallimasticaceae. BioSystems 21, 403-415. Heath, |.B., Bauchop, T. & Skipp, R.A. (1983). Assignment of the rumen anaerobe Neocallimastix frontalis to the Spizellomyce- tales (Chytridiomycetes) on the basis of its polyflagellate zoospore ultrastructure. Can. J. Bot. 61, 295-307. Ho, Y.W. & Bauchop, T. (1991). Morphology of three polycentric rumen fungi and description of a procedure for the induction of zoosporogenesis and release of zoospores in cultures. J. Gen. Microbiol. 137, 213-217. Ho, Y.W., Barr, D.J.S., Abdullah, N., Jalaludin, S. & Kudo, H. (19983). Neocallimastix variabilis, a new species of anaerobic fungus from the rumen of cattle. Mycotaxon 46, 241-258. Li, J., Heath, |1.B. & Bauchop, T. (1990). Piromyces mae and Piromyces dumbonica, two new species of uniflagellate anaerobic chytridiomycete fungi from the hindgut of the horse and elephant. Can. J. Bot. 68, 1021-1033. Liebetanz, E. (1910). Die parasitischen protozoen der weiderkauer- magens. Arch. Protistenkunde 19, 19-80. Orpin, C.G. (1977). The rumen flagellate Piromonas communis: its life-history and invasion of plant material in the rumen. J. Gen. Microbiol. 99, 107-117. Powell, M.J. (1974). Fine structure of plasmodesmata in a chytrid. Mycologia 66, 606-614. pg ee Ys o ee ft ‘ iy | ie a ~ f ‘" a < ; a ' *y wid pbs Hith at sae x) a Oy 7 Te a t 2 7 we nH ine ren ak iy ray he ot ra i: ae At ly ef é perk aveain et! wart) Pa ~~ 2 a ch fh i) a Eu he a eteo he a ray res ees eat ca 9 Rey ied al Wel y e r r * ca ie | babel oh Aes? ps Cond oe itt i. ye, ‘i 1 cy Ai i, 7 Wa RH a is Mn « ee i i: = a = : es i. Ve } Fj bY : “| i aN : pl mel Ay ay v4 is es ¢ i ; 1 ‘ bw ; i Aes 3 sh MAAN e aden RS cere yew in thet sane BRAT I: a Ne yee Deed Fa east wy si raed Wel ee i males be dine i = Sis ST ee ee es me oe ~ = = = oe — — tere i= - Orisa RE Gi chines te ier 1h ee es i a: oe ae ies 7 a ‘ “ | | peg fa ; a = bea Naa, : wah “ity i 7 = 4 V2) Be ae z ‘it ; nate Gigs cea ae ‘ } ‘ee Jaane Nie | alte aii phi bat yolehines — bate rata i i a va y Pe ie ai a age Cpr é F As ai el | Rice Ae 1D, as i ee Me é bs “ge Aa jana Fg arcane cs orate de 1 i} ) P Ais oH ‘im, 4 r mal G of hy f ap Hi We, A eh ‘ sist ; hee } i > 4 7! ‘ ? a 4 : Spy et i i / VS el hie an sin H pace ai, moieys fe eens co ; a va i, Hi bute iy \ ti | Be i na hat Bertin hy pla fi as on ia - tiash : i ey si } Sona ie OE fig al Mus G ie t em Tas a ' fe Ny) sah pee 7 Jy MY COTAXON Volume XLVI, pp. 295-357 April-June 1993 TYPE SPECIMENS OF LICHENS AND LICHENICOLOUS FUNGI IN THE CANADIAN MUSEUM OF NATURE (CANL) PAK YAU WONG Canadian Museum of Nature, P.O. Box 3443, Station 'D', Ottawa, Ontario K1P 6P4, Canada ABSTRACT Type specimens of 737 lichen taxa and lichenicolous fungi (82 holotypes, 418 isotypes, 1 lectotype, 3 lectoparatypes, 22 isolectotypes, 13 syntypes, 13 isosyntypes, 176 paratypes, 6 isoparatypes and 3 isoneotypes) in the lichen herbarium of the Canadian Museum of Nature (CANL) are listed. The name of the taxon, authority, literature citation, category of type, locality information, collector, collector's number, collecting date and CANL accession number are recorded for each taxon, as well as, where applicable, exsiccat tittle and number, and currently accepted name. With the recent reorganization of the Canadian Museum of Nature, a decision was made to upgrade the level of care for the herbarium and to register all the type specimens in the lichen collections (CANL).84Since thiss collection sis*ithe largest in Canada and is one of the most important North American lichen herbaria, the job was given the highest priority. The present catalogue lists in alphabetical order all type specimens of lichens and lichenicolous fungi preserved in the National Lichen Herbarium (CANL), which is housed at the Canadian Museum of Nature. Authority abbreviations for the epithets mainly follow Egan (1987) and its supplements (Egan 1989, 1990). References for the original descriptions are abbreviated according to Bridson (1991). The decision on whether or not a particular specimen is actually a type was made in the standard way, i.e., the label data were compared with the protologue, and annotations of the original author and subsequent annotators were considered. In some cases, the type status was indicated in some way on the labels, but this was not considered to be definitive. If the status of the type is in question, e.g., because of a discrepancy in the data, a query mark (?) is placed after the kind of type in the catalogue. The label data, including locality information, collector, collector's number and collecting 296 date are cited more or less as originally written on the packet, with some standardisation, but the names of countries are given in English reflecting current political boundaries in so far as possible. The CANL accession number on the packet is also presented. If the type is part of a published exsiccata, the name of the series (with its author or distributor) and number are also given at the end of the entry. If paratypes occur in addition to other types or when there is more than one paratype specimen, only locality, collector, collector's number and CANL number are listed. No attempt has been made to revise all the types by including current synonyms, but where the type is known to be synonymous with another name, the currently accepted name is given. I hope that this catalogue will be of assistance to lichenologists who need to locate type specimens required for their research. It may also help curators in other herbaria locate types in their collections which may have gone unnoticed. This catalogue is not intended to be an authorative list of the type status of each specimen, nor can it be a definitive list of all the types now in CANL. As mentioned above, further study may reveal that some specimens in the list may not be types at all, and undoubtedly, some type specimens exist in CANL that have been overlooked. Each specialist must examine the evidence in order to decide the status of the listed specimens and other CANL material not in the list. Finally, it should be noted that in no case is any new typification of any epithet being proposed in this catalogue. Acarospora canadensis H. Magn., Kongl. Svenska Vetensk. Acad. Hand lo 73 232'"(1929)". ISOTYPE. Canada: Ontario. Near the Hog's Back, Rideau River. On limestone rocks. J. Macoun 3646, 2 May 1897 (CANL 11285). Acarospora punae Lamb, Lilloa 14: 231 (1948). ISOTYPE. Argentina: Prov. Tucuman, Valle de Tafi, western Slope of Cumbre Potrerillo. On the overhung side of a large schistose rock. I.M. Lamb 5290, 20 November 1947 (CANL aa kee Bef) Be Acolium bolanderi Tuck., Lich. of California Baier eereog). ISOLECTOTYPE. United States: California, Oakland Hills. H.N. Bolander 56, 1864 (CANL 68772). Reliq. Tuck. No. 101. = Thelomma mammosum (Hepp in Hartung) Massal. Alectoria ambigua Mot., Bryologist 67: 17 (1964). ISOPARATYPES. Canada: Newfoundland, Trinity North District, 2.5 miles NNW of Northern Bight Sta. Exposed rock outcrop in old fire-barren heath. T. Ahti 592, 15 August 1956. (CANL 52083); Placentia West, W of Swift Current, 2 miles NW of Long Pond. On big boulder in treeless heath. T. Ahti S70 7.16 297 August 1956 (CANL 32344). = Bryoria trichodes (Michx.) Brodo & D. Hawksw. subsp. americana (Mot.) Brodo & D. Hawksw. Alectoria arctica Elenk. & Savicz, Trudy Glavn. Bot. Sada 32: 730(1912).. ISOTYPE. Russia: Novaya Zemlya, sinus Krestovaja. J.V. Palibin, 1901 (CANL 104197). = Evernia perfragilis Llano Alectoria cornicularioides P. Jg@rg., Bryologist 78: 77 (1975). ISOTYPE. China: Prov. Shensi, Kuan-tou-san. Among mosses. G. Giraldi, 1896 (CANL 60512). = Bryoria cornicularioides (P. Jgrq.) Brodo & D. Hawksw. Alectoria corymbosa Hue, Expéd. Antarct. Franc. 9.0.3 29 O Ope bys 12 (1908). ISOSYNTYPES. Antarctica: W. Graham Land (Palmer Peninsula) Wandel (Booth) Island. Sur les roches. Expéd. Antarct. Fran¢. 272; 277 and 299, 25 November 1904 (CANL 5707, 5708). = Catillaria corymbosa (Hue) Lamb Alectoria fuscescens Gyelnik, Nytt Mag. Naturvidensk 70: 55 (1932). ISOLECTOTYPES. Finland: Tavastia austr., Hollola, ad truncos Pini locis apricioribus in silva. J.P. Norrlin, 18 September 1882 (CANL 29907 & 32345). Nyl. & Norrl.: Lich. Fenn. Exs. No. 466. = Bryoria fuscescens (Gyelnik) Brodo & D. Hawksw. Alectoria glabra Mot., Fragm. Florist. Geobot. 6: 448 (1960). ISOTYPE. United States: Washington, Clallum Co., Hurricane Ridge, Olympic Peninsula. On trunk of Abies lasiocarpa. B.I. Brown & W.C. Muenscher 129, 24 July 1950 (CANL 28849). =Bryoria glabra (Mot.) Brodo & D. Hawksw. Alectoria imshaugii Brodo & D. Hawksw., Opera Bot. 42: 59 (1977). HOLOTYPE. United States: Idaho, Benewah County, ca. 4.5 miles NW of St. Joe, top of St. Joe Baldy, near BLM Campground Tingley Spring area. On Pseudotsuga menziesii in open conifer stand. G.J. Schroeder L1944, 19 July 1971 (CANL 38827). PARATYPES. United States: Calicornia, I.M. Brodo 20489 & 20502. (CANL 50581 & 50582). Oregon. L.H. Pike 1475 (CANL 42730). Alectoria mexicana Brodo & D. Hawksw., Opera Bot. 42: 62 (1977). HOLOTYPE. Mexico: Oaxaca, Sierra de San Felipe, in pino. On pines. C.G. Pringle 195, 3 June 1894 (CANL 16582). PARATYPES. Mexico: Oaxaca. J. Beharrel (CANL 55334, 55335 & 55342). Alectoria stigmata Bystr., Fragm. Florist. Geobot. 20: 255 298 1974). vaneeea United States: Alaska, Central Pacific Coast District, Kenai Peninsula, Creek Ridge. H. Krog, 28 May 1957 (CANL 54995). = Alectoria sarmentosa (Ach.) Ach. subsp. sarmentosa. Amagdalaria consentiens var. japonica M. Inoue, J. Hattori Bot. Lab. 56: 325 (1984). ISOTYPE. Japan: Honshu, Pref. Nagano, Kiso-gun, Mitake-mura, Mt. Ontake. On rock. M. Inoue 11474, 17 August 1976 (CANL 92651). Amygdalaria continua Brodo & Hertel, Herzogia 7: 505 (1987). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Moresby Island, Takakia Lake. At edge of water partially submerged. I.M. Brodo 10880A, 4 July 1967 (CANL 70598) PARATYPES. Moresby Island. I.M. Brodo 10880B, 10880D, 12804, 12819, 13960, 14288. (CANL 70599, 70601, 70605, 70553, 36494 & 70603). Amygdalaria haidensis Brodo & Hertel, Herzogia 7: 508 (1987). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Moresby Island, Tasu. On mined mountain behind Tasu camp, forest line. On summit rocks. I.M. Brodo 12784, 28 July 1967 (CANL 70594). PARATYPES. Moresby Island. I.M. Brodo 10834, 10933 & 14289. (CANL 87857, 70604 & 70595). Anaptychia fragilissima Kurok., Beih. Nova Hedwigia 6: 60 (1962). ISOTYPE. Japan: Prov. Kii, Mt. Koya. On rocks. S. Kurokawa 60238, 17-18 December 1960 (CANL 20012). Kurokawa: Lich. Rar. Crit EXs NO. 3. Anaptychia japonica var. reagens Kurok., J. Jap. Bot. 35: 354 (1960). ISOTYPE. Japan: Honshu, Prov. Sagami, Mt. Kintoki, Hakone. On trunk of trees. S. Kurokawa 53064, 26 April 1958 (CANL 26391). Kurokawa: Lich. Rar. Crit. Exs. No. 53. Anthracothecium japonicum Kashiw. & Kurok., J. Jap. Bot. 56: 308 (1981). ISOTYPE. Japan: Hokkaido, Prov. Ishikari, Noborikawa, Yubari- City. On bark of Acer. H. Shibuichi 6067, 4 August 1980 (CANL 78691). Kurokawa: Lich. Rar. Crit. Exs. No. 502. Anzia afromontana R. Sant. in Moberg, Moberg: Lich. Sel. Exs. Ups., Thunbergia 2: 1 (1986). ISOTYPE. Tanzania: Arusha Prov., Mt. Meru, ca. 2 km N of Kitoto Camp. On Stoebe kilimandscharica. R. Santesson 22949, 7 January 1971 (CANL 95077). Moberg: Lich. Sel. Exs. Ups. No. Ls Arthopyrenia submuriformis R. Harris, Michigan Bot. 12: 15 (1973). ISOTYPE. United States: Pennsylvania, West Chester. On bark Pe pe) of hickory. F. Windle, July 1898 (CANL 19709). Cummings: Dec. N. Amer. Lich. No. 299. = Strigula submuriformis (R. Harris) R. Harris Arthopyrenia willeyana R. Harris, Michigan Bot. 12: 16 (1973). ISOTYPE. United States: Michigan, Iosco County. In jackpine- aspen woods around Corsair State Forest Campground. R.C. Harris 818, 18 September 1965 (CANL 37273). = Anisomeridium nyssaiegenum (Ellis & Everh.) R. Harris. Aspicilia quartzitica W. Weber, Bryologist 74: 183 (1971). ISOTYPE. United States: Colorado, Boulder County, outer foothills of the Front Range at Boulder. on resistant quartzite rocks of the Lyons formation, base of the first "Flatiron" on east face of Green Mountain. W.A. Weber, 24 January 1971 (CANL 52114). Weber: Lich. Exs. Colo. No. 364. Bacidia colchica Vézda, Folia Geobot. Phytotax., Praha 14: 203 (1979). ISOTYPE. Russia: Transcaucasia, Colchis. Distr. Gagra, in valle angusta rivi Zo Ekvara. Ad folia Buxi colchicae. A. Vézda, 13 June 1978 (CANL 73823). Vézda: Lich. Sel. Exs. No. 1660. Bacidia lisowskii Vézda, Folia Geobot. Phytotax., Praha 15: 87 (1980). ISOTYPE. Zaire: Haut-Zaire, Kisangani, prope vicum Batiambale. Ad folia arborum. S. Lisowski, 27 October 1976 (CANL 73852). Vézda: Lich. Sel. Exs. No. 1690 Bacidia nivalis Follm., Philippia 4: 30 (1978). ISOTYPE. United States: Washington, Whatcom Co., fleckférmig an lichtoffenen, etwas windgeschiitzten, langfristig schneebedeckten Kulm- und Neigungsflachen niederer Felsblicke und bodennaber Gesteinstrtimmer im Sporastatietum testudineae Frey, im Bereich der Sommerschneegrenze am Nordosthang des Mount Baker liber dem Austinpass. G. Follmann, July 1969 (CANL 69925). Follmann: Lich. Exs. Sel. No. 262. Bacidia scutellifera Vézda, Folia Geobot. Phytotax., Praha 10: 421 (1975). ISOTYPE. Tanzania: Distr. Morogoro, montes Nguru, in monte Mafulumula supra vicum Mnembule. Epiphylla in pluviisilva montana. T. Pécs 6438, 20-21 August 1971 (CANL 57290). Vézda: Lich. Sel. Exs. No. 1310. Bacidia spirospora var. patagonica Lamb, Farlowia 4: 453 (1955). HOLOTYPE. Argentina: Patagonia Prov. Chubut, Lago Cisnes near Lago Menendez. On trunk of Colletia at edge of the forest. I.M. Lamb 5933, 5.11.1950 (CANL 6024). Bacidia vasakii Vézda, Folia Geobot. Phytotax., Praha 18: 64 (1983). ISOTYPE. Russia: Colchis (Transcaucasia occid.), distr. 300 Adler, in valle angusta rivi Kudepsta, 5 km ab ostio in mare. Ad folia Buxi colchicae. A. Vézda, 19 June 1979 (CANL 85982). Vézda: Lich. Sel. Exs. No. 1869. Bacidia violascens Kalb & Vézda, Folia Geobot. Phytotax., Praha 15: 309 (1980). ISOTYPE. United States: Hawaii Insulae, Insula Kauai. Ad ramulos arboris (Prosopis sp. cult.). O. Degener 34240, 29 February 1977 (CANL 76382). Vézda: Lich. Sel. Exs. No. 1733. Baeomyces weberi Thomson, Bryologist 73: 632 (1970). ISOTYPE. Papua New Guinea: Morobe District, Mount Kaindi near Papua Wau. On clay banks of road-cuts not far below summit. W.A. Weber, 18 June 1968 (CANL 33497). Weber: Lich. Exs. Coloe No? 73.3137 Bahianora poeltii Kalb, Kalb: Lich. Neotr. Fasc. 8: 4 (1984). ISOTYPE. Brazil: Bahia: Chapada Diamantina, Serra do Tombador; etwa 1 km vor der Stadt Morro do ChapeG. Am Grunde von Grdasern in einer Caatinga. K. Kalb, 20 July 1980 (CANL 89418). Kalb: Lich. Neotr. No. 302. Bilimbia finkii Vainio, Mycologia 21: 35 (1929). ISOTYPE. Puerto Rico: Rio Piedras. Open field on post. B. Fink 534, 12 January 1915 (CANL 5847). = Bacidia finkii (Vainio) Zahlbr Bryoria carlottae Brodo & D. Hawksw., Opera Bot.: 42: 101 (1977). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Graham Island, two mi. SE of Port Clements, S of the Tlell- Port Clements Road. I.M. Brodo 18096, 11 July 1971 (CANL 38266). PARATYPES. Canada: British Columbia, Queen Charlotte Islands. I.M. Brodo 9900, 10657B, 12879, 18093, 18499 & 18503B (CANL 45536, 28902, 45490, 38264, 38265 & 42514). Bryoria friabilis Brodo & D. Hawksw., Opera Bot.: 118 (1977). ISOTYPE. United States: Washington, Gate. On oak trees. A.S. Foster, 14 October 1911 (CANL 36151). PARATYPES. British Columbia. I.M. Brodo 8179 (CANL 52607); J. Macoun s.n. & 77 (CANL 16156 & 16162); Quebec. I.M. Brodo 18678 & 18691D (CANL 37596 & 37587). Bryoria pikei Brodo & D. Hawksw., Opera Bot. 42: 125 (1977). HOLOTYPE. United States: Oregon, Marion Co.. Epiphyte in an old fruit orchard in a clearing in a douglas fir forest near Winter Falls in Silver Creek Falls State Park E of Salem. On young douglas fir. L. Pike 2475, 9 April 1972 (CANL 38271). PARATYPES. Canada: British Columbia, Queen Charlotte Islands. I.M. Brodo 12874A, 13839 & 18144A (CANL 45543, 45510 & 38383); Nova Scotia. J. Speer 73-24 (CANL 45159); United States: Oregon. I.M. Brodo 20954 (CANL 99034); L.H. Pike 1110 (CANL 48810); Washington. I.M. Brodo 15438b (CANL 32327); L.H. Pike 844 (CANL 48821. 301 Bryoria pseudocapillaris Brodo & D. Hawksw., Opera Bot. 42: AA Sal @ EMA pe HOLOTYPE. United States: Oregon, Curry County, Cape Blanco, 8 miles N of Port Orford. On Sitka spruce at headland. I.M. Brodo 20539, 28 June 1974 (CANL 50596). Bryoria salazinica Brodo & D. Hawksw., Opera Bot. 42: 130 (1977). ISOTYPE. United States: Massachusetts, Springfield. W.G. Farlow, May 1878 (CANL 36152). PARATYPES. Canada: Prince Edward Island. J. Fabiszewski (CANL 31266); R.R. Ireland 10298 (CANL 35527). Bryoria spiralifera Brodo & D. Hawksw., Opera Bot.: 42: 131 (1977). HOLOTYPE. United States: California, Humboldt County. Pine forest near Manila. On Pinus contorta. S. Dowty 137, 22 January 1972 (CANL 38403). PARATYPES. United States: California. Brodo 20501, 20509 & 20511A (CANL 50580, 50594 & 50578). Buellia excellens H. Magn., Medd. Gdteb. Bot. Trdadg. 17: 69 (1947). ISOTYPE. Argentina: Prov. Salta, Quebrada de San Lorenzo. M. Digilio-Grassi 335, 1946 (CANL 18205). = Rhizocarpon compositum Lamb Buellia galapagona W. Weber, Bryologist 74: 188 (1971). ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz (Indefatigable I.), Academy Bay. On lava rocks along the shore of the bay just above high tide mark. W.A. Weber, 30 January 1964 (CANL 33508). Weber: Lich. Exs. Colo. No. 344. Buellia imshaugii Hafellner, Beih. Nova Hedwigia 62: 58 (1979). HOLOTYPE. Canada: South Saskatchewan. J. Macoun, 13 July 1879 (CANL 19326). ISOTYPE. (CANL 19327). Buellia multispora Kalb & Vézda, Folia Geobot. Phytotax., Praha 14: 203 (1979). ISOTYPES. United States: Hawaii Insulae, Oahu, Keawaula Valley. On living trunks of exotic Leucaena. O. & I. Degener 34257, 16 March 1977 (CANL 73863). Vézda: Lich. Sel. Exs. No. 1670; (CANL 102697). Kalb: Lich. Neotr. No. 408. Buellia rivas-martinezii Barreno & Crespo, Philippia 2: 283 (1975). ISOTYPE. Spain: Prov. Madrid, zerstreut an stark geneigten kristallinen Gipsabbriichen in licht-und windoffenen Trockenrasenlticken im Lecideion gypsicolae Crespo et Barr., 620 m. SO, Cerros Yesiferos de Valdemoro nérdlich Aranjiez. S. Rivas-Martinez & A. Crespo, March 1972 (CANL 57049). Follmann: Lich. Exs. Sel. No. 143. Buellia tephrodes Lamb, Farlowia 4: 468 (1955). 302 HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel Huapi, Isla Victoria. On rocks at shore of lake, about 4 m above water level. I.M. Lamb 5855 p.p. (CANL 18365). Buelliella eximia Kalb & Hafellner, Kalb: Lich. Neotr. Fasc. 11: 6 (1990). ISOTYPE. Venezuela: Lara Prov., Distr. Torres, Etwa 35 km E von Barquisimeto. In einer trockenen Steppenlandschaft, parasitisch auf Pyxine cocoes (Sw.) Nyl. K. & A. Kalb, 10 August 1989 (CANL 102740). Kalb: Lich. Neotr. No. 452. Buelliella physciicola Poelt & Hafellner, Beih. Nova Hedwigia 623), 15S VOL979 )\. ISOTYPE. Czechoslovakia: Moravia austro-occidentalis, in valle fluminis Rokytna supra urbem MoravskYyY Krumlov. In thallo Physciae sciastrae parasitice vigens, in rupibus conglomeraticis. J. Poelt & A. Vézda, 16 April 1974 (CAN 73837). Vézda: Lich. Sel. Exs. No. 1675. Byssoloma amazonicum Kalb & Vézda, Nova Hedwigia 51: 437 (1990). ISOTYPES. Brazil: Amazonas, Regenwdlder am Rio Negro, zwischen 100-200 km oberhalb von Manaus. Foliicola in pluviisilva. K. Kaib, 14-18 October 1980 (CANL 103355). Vézda: Lich Sel. Exs. No. 2427; (CANL 103753). Kalb: Lich. Neotr. No. 482. Byssoloma anomalum Kalb & Vézda, Nova Hedwigia 51: 438 (1990). ISOTYPE. Brazil: Amazonas, Regenwdlder am Rio Negro, zwischen 100-200 km oberhalb von Manaus. K. Kalb, 14-18 October 1980 (CANL 103754). Kalb: Lich. Neotr. No. 483. Calicium adspersum subsp. australe Tibell, Publ. Herb. Univ. Uppsala 12: 1 (1984). ISOTYPE. Australia: Tasmania, Ben Lomond National Park, 32 km E of Evansdale, along Ben Lomond Road, 1 km NW of Carr Villa. On decorticated, dead but still standing trunk of Eucalyptus sp. L. Tibell 11465, 14 March 1981 (CANL 87449). Tibell: Caliciales Exs. No. 76. Calicium constrictum Tibell, Lichenologist 14: 223 (1982). ISOTYPE. Costa Rica: Puntarenas, on old Camino Real, 3.5 km SE of Paso Real. On bark of roadside fence-post. L. Tibell 8233, 30 December 1978 (CANL 87450). Tibell: Caliciales Exs. NOW Lic Calicium cryptocroceum Tibell, Publ. Herb. Univ. Uppsala 10: L(1982)% ISOTYPE. New Zealand: Canterbury, 9.5 km SSE of Arthur's Pass, Bealy Spur, along track to Grasmere Hut. On decorticated stump of Nothofagus solanderi v. cliffortioides. L. Tibell 10084, 17 December 1980 (CANL 82033). Tibell: Caliciales Exs. No. 51. Calicium curtisii var. splendidula G. K. Merrill, Bryologist 303 LAS 1L07 C1909) ISOTYPE. United States: Maine, Knox County, Rockland. On Rhus typhina. G.K. Merrill, 3 May 1909 (CANL 20271). Merrill: Tacn. Exse No. 724% = Phaeocalicium curtisii (Tuck.) Tibell. Calicium fuscipes Tuck., Genera Lichenum p. 240 (1872). SYNTYPE. United States: New Jersey, Closter. C.A. Austin, November 1864 (CANL 68717). Reliq. Tuck. No. 22. = Mycocalicium fuscipes (Tuck.) Fink Calicium obscurum G. K. Merrill, Bryologist 12: 107 (1909). ISOTYPE. United States: Maine, Rockland. On dead fungus. G.K. Merrill, 5 September 1909 (CANL 20337). Merrill: Lich. Exs. NOs. 92% = Chaenotheca brunneola (Ach.) Mull. Arg. Calicium parvum Tibell, Symb. Bot. Upsal. 21: 84 (1975). ISOTYPE. Canada: Ontario, Renfrew County, Petawawa Forest Experiment Station, not far from Mill Lake. On trunk of Picea in a swamp. L. Tibell 4638, 21 August 1972 (CANL 55417). Caloplaca bisagnonis B. de Lesd., Bull. Soc. Bot. France 98: he asf Ge toh i es ISOTYPE. Italy: Genova, Val Bisagno, Prato. Sbarbaro, 21 April 1951 (CANL 17465). Caloplaca brattiae W. Weber, Graphis Scripta 2: 168 (1989). ISOTYPE. United States: California, Santa Barbara County, Channel Islands, Santa Cruz Island. West end of the island. On top of the rocky headland just above the spray zone. W.A. Weber & C. Bratt, 8 January 1986 (CANL 97959). Weber: Lich. Exs. Colo. No. 660. Caloplaca britannica R. Sant., Lichenologist 24: 2 (1992). ISOTYPE. Scotland: Caithness (V. C. 109), Crosskirk Bay, c. 10 km W of Thurso. On seashore rocks. R. Santesson 20410, 18 July 1969 (CANL 104124). Caloplaca citrina var. arcis Poelt & Vézda, Vézda: Lich. Sel. Exs. Fasc. 99: 6 (1990). ISOTYPE. Austria: Stiria, Distr. Feldbach, Riegersburg. Ad saxa andesitica sub arcem. G. Kantvilas, H. Mayrhofer & A. Vézda, 22 September 1990 (CANL 103398). Vézda: Lich. Sel. Exs. No. 2470. Caloplaca fraxinea Lamb, Rep. (Annual) Natl. Mus. Canada, Bill ewe ces 042"( 1954)", HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, at Baddeck. On black ash bark in a swamp. J. Macoun 1737, 8 July 1898 (CANL 17744). Caloplaca hensseniana Kalb, Kalb: Lich. Neotr. Fasc. 11: 7 (1990). ISOTYPE. Venezuela: Lara Prov. Distr. Iribarreno, etwa 15 km E von Barquisimeto. Epiphytisch in trockener Steppen- 304 Vegetation, untermischt mit Cacteen. K. & A. Kalb, 20 August 1989 (CANL 102742). Kalb: Lich. Neotr. No. 454. Caloplaca litoricola Brodo, Bryologist 87: 98 (1984). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Graham Island, Tow Hill. On shoreline rocks. I.M. Brodo 9905, 16 June 1967 (CANL 85435). Brodo: Lich. Canad. Exs. No. 156. Caloplaca marina subsp. americana Arup, Bryologist 95: 158 (1992). ISOTYPE. United States: Oregon, Lincoln County, Rocky Creek Wayside State Park, just south of Whale Cove. On volcanic seashore rocks. U. Arup 89189, 19 April 1989 (CANL 103721). Caloplaca sbarbaronis B. de Lesd., Bull. Soc. Bot. France 96: 173 (1949). ISOTYPE. Italy: Liguria orientalis, Bonassola. Rupicola prope mare. Sbarbaro, February 1949 (CANL 17859). Caloplaca sorediata var. tenuis H. Magn., Ark. Bot. 33A: 134 (1946). SYNTYPE. Sweden: Lycksele Lappmark, par. Stensele, Kyrkberget. Under overhanging rocks. A.H. Magnusson 7841, 5 July 1924 (CANL 17873). = Xanthoria sorediata (Vainio) Poelt Caloplaca spotornonis B. de Lesd., Bull. Soc. Bot. France 100: 177 (1953). ISOTYPE. Italy: Savona, Spotorno, loco "Monte". Sbarbaro, December 1951 (CANL: 17876). Caloplaca trabicola H. Magn., Medd. Gdteb. Bot. Tradg. 18: 219 (1950). PARATYPE. Uruguay: Florida Prov., Estancia, 25 de Agosto. Sobre postes de alambrado. H.S. Osorio 1752, 22 November 1949 (CANL 17887). Candelariella faginea Nimis & Poelt, Nova Hedwigia 49: 276 (1989). ISOTYPE. Italy: Calabria, Serra del Prete, loco Massiccio del Pollino dicto. Ad corticem Fagi. P.L. Nimis & J. Poelt, 10. July 1988. (CANL 100651). Vézda: Lich. Sel. Exs. No. 2377. Candelariella hudsonica Hakul., Ann. Bot. Soc. Zool.-Bot. Fenn. "Vanamo" 27: 49 (1954). HOLOTYPE. Canada: Northwest Territories, Franklin District. On earth, north shore of Hudson Strait. R. Bell 1660, August 1897 (CANL 12961). = Candelariella canadensis H. Magn. Candelariella lambii Hakul., Ann. Bot. Soc. Zool.-Bot. Fenn. "Vanamo" 27: 49 (1954). HOLOTYPE. Argentina: Prov. Tucumen, Valle de Tafi, west slope of Cumbre Potrerillo. On the top of a block of schistose rock in alpine pasture. I.M. Lamb 5413, 24 November 1947 (CANL 12962). 305 Candelariella plumbea Poelt & Vézda, Folia Geobot. Phytotax., Praha 11: 88 (1976). ISOPARATYPE. Austria: Karawanken, Karnten, Kanzianiberg bei Finkenstein SE Villach, W- bis S-sei- tige Abbritiche. J. Poelt, 10 February 1974 (CANL 73768). Poelt: Pl. Graecenses No. 137. Catapyrenium caeruleopulvinum Thomson, Bryologist 90: 30,1987. ISOTYPE. United States: California, San Bernardino, 2 miles SW of Parker Dam near the Colorado River. On soil. T.H. Nash 8449, 26 February 1974 (CANL 52409). PARATYPE. United States: Arizona. T7.H. Nash 8047. (CANL 52394). Catillaria italica B. de Lesd., Bull. Soc. Bot. France 97: 170 (1950). ISOTYPE. Italy: Rapallo (Genova), + Loco opaco’_ infra Montallegro, ad maceriam. Sbarbaro, November 1949 (CANL 5758). Catillaria melanopotamica Lamb, Farlowia 4: 445 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel Huapi, Isla Victoria. On trunk of Nothofagus dombeyi. I.M. Lamb 5815, 26 January 1950 (CANL 5769). PARATYPE. Argentina: Isla Victoria. I.M. Lamb 5836 (CANL 5768). Cetraria alaskana cC. cCulb. & Culb., Bryologist 69: 200 (1966). ISOTYPE. United States: Alaska, Along the Pitmegea River, 15 miles upstream from Cape Sabine. On wet tundra. J.W. Thomson, 10-17 July 1958 (CANL 20583). Thomson: Lich. Arct. No. 13. = Cetrelia alaskana (C. Culb. & Culb.) Culb. & C. Culb. Cetraria australiensis W. Weber ex Karnef., Bot. Not. 130: yee ek: be ISOTYPE. Australia: New South Wales, Snowy Mountains, Kosciusko State Park. W.A. Weber & D. McVean, 2 February 1968 (CANL 52204). Weber: Lich. Exs. Colo. No. 454. Cetraria fendleri f. coralligera W. Weber, Univ. Colorado Stud., Ser. Biol. 10: 11 (1963). ISOTYPE. United States: Colorado, Larimer County, Lumpy Ridge nachst Twin Owls an der Route zum Gem Lake, 1.5 Meilen N Lake Estes. R.A. Anderson, 27 August 1962 (CANL 71724). Crypt. Exs. Vindob. No. 4439. = C. coralligera (W. Weber) Hale Cetraria iberica Crespo & Barreno, Anales Jard. Bot. Madrid 37: 205 (1980). ISOTYPE. Spain: Guadalajara, Tamaj6én. Ad Cistus ladaniferus. A. Crespo & E. Barreno, May 1977 (CANL 78851). Vézda: Lich. Sel. Exs. No. 1752. Cetraria islandica var. arborialis Zahlbr., Cat. Lich. Univ. 306 6: 333 (1930). LECTOTYPE. Canada: British Columbia, Glacier, Marion Lake. On bushes. J. Macoun, 23 August 1904 (CANL 15550). = C. subalpina Imsh. Cetraria subalpina Imsh., Mycologia 42: 746 (1950). ISOTYPE. United States: Washington, Mt. Rainier National Park, Mt. Wow. On branches of small shrubs. H.A. Imshaug 1876, 21 August 1948 (CANL 15549). PARATYPES. Mt. Rainier National Park. H.A. Imshaug 261 & 267 (CANL 15551, 15552); Cataract Falls. H.A. Imshaug 510 (CANL 24532). Chaenothecopsis nana Tibell, Publ. Herb. Univ. Uppsala 4: 4 (1979). PARATYPE. Sweden: Varmland, Norra Finnskoga par., N of Digerfallet. On trunk of Picea abies. S. Sundell 13382, 16 August 1979 (CANL 71126). Tibell: Caliciales Exs. No. 35. Chaenothecopsis nigropedata Tibell, Symb. Bot. Upsal. 27: 135 (1987). ISOTYPE. New Zealand: South Island, Southland, Longwood State Forest, 19 km NNW of Riverton, along Pourakino River, N of Pourakino Campground. On decorticated stump. L. Tibell 10428, 16 January 1981 (CANL 99793). Tibell: Caliciales Exs. No. 160. Chaenothecopsis sagenidii Tibell, Symb. Bot. Upsal. 27: 148 (1987). ISOTYPE. New Zealand: Otago, 15 km WNW of Owaka, Catlins Forest, along Catlins River, S of the junction with Chloris stream. On still corticated parts of decaying stump of Nothofagus menziesii. L. Tibell 10282, 11 January 1981 (CANL 99795). Tibell: Caliciales Exs. No. 162. Chaenothecopsis tasmanica Tibell, Publ. Herb. Univ. Uppsala 16: 6 (1985). ISOTYPE. Australia: Tasmania, Mt. Field National Park, W shore of Lake Dobson. On dead leaves of Richea pandanifolia. Lb. .Tibeli 11227, (¢S March’ ©1981) '*-(CANL” °89742))."" *Tibely: Caliciales Exs. No. 118. Charcotia rufidula Hue, Bull. Soc. Bot. France 62: 17 (1915). ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth (Wandel) Island. Deux Exped. Antarct. France 123, October 1908. (CANL 10604). = Arthonia rufidula (Hue) D. Hawksw., R. Sant. & @vstedal (1991). Cheiromycina petri D. Hawksw & Poelt, Lichenologist 22: 219 (1990). ISOTYPE. Austria: Styria, in collibus "Oststeierisches Hiigelland", distr. Feldbach, Lamberg, in pago Petersdorf II. Ad corticem trunci Betulae. J. Poelt & H. Pittoni, 27 November 1987 (CANL 103356), Vézda: Lich. Sel. Exs. No. 2428. 307 Cladina conspicua Ahti, Ann. Bot. Fenn. 23: 224 (1986). ISOTYPE. Canada: Newfoundland, Humber East District, ca. 2.4 km N of Gaff Topsail, timberline heath. 7. Ahti 2997, 27 June 1956 (CANL 96933). PARATYPE. Canada: Newfoundland, St. Mary District. T. Ahti 9461. (CANL 96934). Cladonia alpestris f. aberrans des Abb., Bull. Soc. Sci. Bretagne 16, Fasc. hors sér. 2: 93 (1939). ISOLECTOTYPE. United States: Alaska, St. Paul Island. J. Macoun, 26 June 1897, (CANL 6540). Macoun: Canad. Lich. No. LVL; Cladonia anitae Culb. & C. Culb., Mycologia 74: 663 (1982). ISOTYPE. United States: North Carolina, Onslow Co., inter Folkstone et Holly Ridge. Ad terram arenariam. W.L. Culberson 18539, 1 April 1981 (CANL 85967). Vézda: Lich. Sel. Exs. No. 1854. Cladonia arbuscula subsp. beringiana Ahti, Ann. Bot. Soc. Z001.-Bot. Fenn. "Vanamo" 32: 109 (1961). ISOTYPE. Canada: Northwest Territories, Mackenzie District, Great Bear Lake, at head of Great Bear River where it flows out of Great Bear Lake. A.A. Lindsey 417, 23 July 1951 (CANL 9766). = Cladina arbuscula (Wallr.) Hale & Culb. subsp. beringiana (Ahti) Golubk. -Cladonia boliviana Ahti, Ann. Bot. Soc. Zool.-Bot. Fenn. wVanamno™ (3227 °PS i (1L96L)s ISOTYPE. Bolivia: La Paz, 10,000 ft. M. Bang 20a (20 on holotype), 1889. (CANL 59522). = Cladina boliviana (Ahti) Ahti Cladonia convoluta var. vagans Follm., Philippia 2: 208 (1975). ISOTYPE. Spain: Prov. Teruel. Zwergstrauchheide in der Sierra de la Costera unweit Fuentes Calientes. G. Follmann, July 1973 (CANL 57051). Follmann: Lich. Exs. Sel. No 145. Cladonia crispata var. infundibulifera f. albopunctata G. kK. Merrill, Bryologist 11: 110 (1908). SYNTYPES. Canada: Yukon, Hunker Creek. In bogs. J. Macoun 104 & 108, 28 July 1902 (CANL 7411 & 7382). = C. crispata (Ach.) Flotow var. cetrariiformis (Delise) Vainio (fide T. Ahti) Cladonia favillicola Trass, Investigationes Naturae Orientis Extremis (Tallinn) p. 198 (1963). ISOTYPE. Russia: Kamczatka, 20 km a Kozyrevsk in SO, ad flumen Pahtza, in lariceto-ledose, ad laricis truncum prolapsum crasso strato ex favilla vulcanica et aculeis laricis tectum. H. Trass, 23 August 1960 (CANL 57970). = C. botrytes (K. Hagen) Willd. Cladonia gracilis subsp. vulnerata Ahti, Ann. Bot. Fenn. 17: 308 207 (1980). ISOTYPE. United States: Alaska, Chugach Mts., Blueberry Lake Campground, Mile 23, Richardson Hwy. In montane tundra. T. Ahti & J.W. Thomson 23642, 25-26 July 1967 (CANL 75105). PARATYPES. Canada: British Columbia. I.M. Brodo 9760 & 18156. (CANL 31098 & 61622). Cladonia hedbergii Ahti, Lichenologist 9: 4 (1977). ISOTYPE. Kenya: Central Prov., Nanyuki Distr., Mt. Kenya, National Park Road (Naro Moru Track). On the ground in a wet slope in the Hagenia-Hypericum zone. R. Santesson "32495" (22040 in protologue), 23 January 1970 (CANL 99586). Moberg: Lich. Sel. Exs. Upsal. No. 56. Cladonia kauaiensis G. K. Merrill, Ark. Bot. 31A(6): 28 (1944). ISOLECTOTYPE. United States: Hawaiian Islands, Kauai. On earth. A.A. Heller, 10-16 September 1895 (CANL 20334). Merrill: Lich. Exs. No. 89. Cladonia labradorica Ahti & Brodo, Bryologist 84: 238 (1981). HOLOTYPE. Canada: Québec, Post-de-la-Baleine, 8 milles de la mer (Baie d'Hudson). A. Vachon, 20 August 1974 (CANL 71500). PARATYPES. Canada: Quebec. J.P. Ducruc 73-4062 (CANL 49135); G. Lemieux 21399 (CANL 70052); J.L. Lethiecgq QFB-E 4495 & 4497 (CANL 48848 & 48881). Cladonia libifera Savicz, Novosti Sist. NizSih Rast. 1965: 167 (1965). ISOTYPE. Russia: Sibiria orientalis, Respublica autonoma Jakutensis, in valle fluminis Aldan, prope Chandyga apud fontem "Tjoply kljucz' in lariceto ad truncum putridum. L.N. Tjulina, 1949 (CANL 48266). = C. pocillum (Ach.) O. Rich.? (fide T. Ahti) Cladonia magyarica Vainio, Fl. Hung. Exs. No. 715 (1927). ISOLECTOTYPE. Hungary: Comit, Pest Prov., "Bugaci nagyerddé" prope oppidum Kecskemét. Ad arenam mobilem in silva. G. TimkO, May 1924 (CANL 19843). Cladonia norvegica Tonsberg & Holien, Nordic J. Bot. 4: 79 (1984). ISOTYPE. Norway: S@r-Tr@ndelag, Melhus, inter montem Loasen et rivulum Loa. Ad basin abscisam arboris (Picea abies). T. Teansberg 6870, 5 June 1982 (CANL 89184). Vézda: Lich. Sel. Exs. No. 1978. Cladonia patagonica A. Evans, Rev. Bryol. Lichénol. 24: 135 (1955). ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menéndez, W. end. I.M. Lamb 5886, 2 February 1950 (CANL 9081). = C. squamosa (Scop.) Hoffm. (barbatic acid chemotype) (fide TRAnt sw 992) Cladonia physodalica Elix, Lich. Austral. Exs. Fasc. 9, No. 204 (1990). 309 ISOTYPE. Australia: Queensland, Main Coast Range, Mt. Lewis Track, Mary Creek, 15 km NNW of Mt. Molloy. On soil on large boulder. J.A. Elix 16947, 30 June 1984 (CANL 102793). Elix: Lich. Austral. Exs. No. 204. = C. poeciloclada des Abb. (physodalic acid chemotype) (fide T. Ahti 1992). Cladonia rangiferina f. caerulescens Schade, Ber. Deutsch. Bot. Ges. 69: 284 (1956). ISOTYPE. Canada: Manitoba, Oxford Lake. H.J. Scoggan 60, 3 July 1949. (CANL 9394). = Cladina stygia (Fr.) Ahti Cladonia rangiferina f. leucosticta G. K. Merrll, Bryologist 11: 109 (1908). SYNTYPES. Canada: Yukon, Hunter Creek. In a bog. J. Macoun 112 & 113 (CANL 9141 & 9140). = C. wainii Savicz (fide T. Ahti) Cladonia subchordalis A. Evans, Rev. Bryol. Lichénol. 24: 133 (1955). ISOTYPE. Argentina: Patagonia, Prov. Chubut, w. end of Lago Menéndez, Torrecillas Stream. On mossy soil between morainic boulders. I.M. Lamb 5917, 4 February 1950 (CANL 9671). Cladonia terrae-novae Ahti, Arch. Soc. Zool.-Bot. Fenn. "Vanamo" 14: 131 (1960). ISOTYPE. Canada: Newfoundland: Placentia West District, 4 miles NW of Long Pond. T. Ahti 108, 16 August 1956 (CANL 58385). = Cladina terrae-novae (Ahti) Hale & Culb. Cladonia terrae-novae f. cinerascens Ahti, Ann. Bot. Soc. Zool.-Bot. Fenn. "Vanamo" 32: 82 (1961). ISOTYPE. Canada: Newfoundland, Placentia West District, 2 mi. NW of Long Pond. T. Ahti 7, 17 August 1956 (CANL 58408). = Cladina terrae-novae (Ahti) Hale & Culb. f. cinerascens (Ahti) Brodo Cladonia thomsonii Ahti, Bryologist 81: 334 (1978). ISOPARATYPE. United States: Northern Alaska, on the point of Point Barrow. J.W. Thomson, S. Shushan & A.J. Sharp, 18 July 1958 (CANL 20591). Thomson: Lich. Arct. No. 21. Cliostomum luteolum Gowan, Mycologia 82: 769 (1990). HOLOTYPE. Canada: New Brunswick, Albert Co., Fundy National Park, Point Wolfe. On Picea rubens. S.P. Gowan 3836, 1 August 1980 (CANL 82486). Cliostomum vitellinum Gowan, Mycologia 82: 769 (1990). PARATYPE. Canada: New Brunswick, Westmoreland Co., near Sackville, Johnson's Mills. On trunk of Abies balsamea. M. Fredericksen 78008, 1 February 1978 (CANL 67236). Coccocarpia stellata Tuck., Proc. Amer. Acad. Arts. 5: 402 (1862). 310 ISOTYPE. United States: South Carolina, Santee Canal. H.W. Ravenel 250 (CANL 68742). Reliq. Tuck. No. 63. Coccotrema maritimum Brodo, Bryologist 76: 263 (1973). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Moresby Island, Jedway. On shoreline rocks. I.M. Brodo 12459, 25 July 1967 (CANL 30581). PARATYPES. Canada: British Columbia. I.M. Brodo 14355 (CANL 30862); K.E. Ohisson 1011, 1023) 1050, ) 1059, 21/3 i216 2448; (CANL 32299, 32300, 32295) 32296732298 ,32297) &°32301)7 G.F. Otto 1504 (CANL 36507); M.J. Shchepanek 253 (CANL 30876). Collema microptychium Tuck., Lich. Calif.: 35 (1866). ISOLECTOTYPE. United States: Massachusetts, Amherst, Parttrigg's Swamp. E. Tuckerman, 1861 (CANL 23165). = C. leptaleum Tuck. Coniocybe amabilis Tibell, Publ. Herb. Univ. Uppsala 10: 6 (1982). ISOTYPE. New Zealand: Wellington, Tongariro National Park, 3 km NE of Ohakune, along Mountain Road. On decorticated stump of Dacrydium cupressinum. L. Tibell 13605, 3 June 1981 (CANL 82046). Tibell: Caliciales Exs. No. 65. Conotremopsis weberiana Vézda, Folia Geobot. Phytotax., Praha 1281411997) 2 ISOTYPE. Australia: Tasmania, Lake St. Clair National Park. On saplings near shore of Lake St. Clair at beginning of trail to Mt. Hugel. W.A. Weber & D. McVean 49678, 24 February 1968 (CANL 92385). Dactylina beringica Bird & Thomson, Canad. J. Bot. 56: 1612 (1978). ISOTYPE. Canada: British Columbia, Summit Lake. Mile 392.5, Alaska Hwy, slope of Mt. George. J.W. Thomson & T. Ahti 16553, 13 August 1967 (CANL 59778). Dermatocarpon corticola ("corticolum") Rd&dsdnen, Anales Soc. Ci. Argent. 128: 147 (1939). ISOTYPE. Uruguay: Dept. Artigas, Sucio. On bark of trees. W.G. Herter, 1934 (CANL 1063). Dermatocarpon gorzegnoense Servit, Ann. Mus. Civico Storia Nat. Giacomo Doria 66: 242 (1953). ISOTYPE. Italy: Piemonte, Langhe, Gorzegno. Sbarbaro, September 1951 (CANL 1081). Dermatocarpon vagans Imsh., Mycologia 42: 753 (1950). ISOTYPE. United States: Wyoming, NE corner of Yellowstone National Park. On arid soil. H.A. Imshaug 15, 23 June 1948 (CANL 1185). Dimerella chiodectonoides Kalb, Kalb: Lich. Neotr. Fasc. 10: 7 (1988). ISOTYPE. Ecuador: Azuay Prov., etwa 35 km siidlich von Cuenca. 311 In sehr dichten, feuchten Waldresten in einem Graspdramo. K. & A. Kalb, 26 August 1987 (CANL 102706). Kalb: Lich. Neotr. No. 417. Dimerella degeneri Kalb & Vézda, Folia Geobot. Phytotax., Prana 115253107 (1980). ISOTYPE. United States: Hawaii Insulae, Insula Oahu, Kaukonahua Gulch prope territorium Universitatis. Ad corticem arboris (Ficus sp. cult.). O. et I. Degener 34269, 17 May 1977 (CANL 76378). Vézda: Lich. Sel. Exs. No. 1729. Dimerella frederici Kalb, Folia Geobot. Phytotax., Praha 15: 310 (1980). ISOTYPES. United States: Hawaii, island Hawaii, Kilauea, Kau Desert. On bark of naturalized Samanea saman. O. et I. Degener 34230, 1 February 1977 (CANL 76379, 77581, 77633, and VODs \ovanveZda isi Chace se ExXGs NOt 6173074 Poelt:) PL. Graecenses No. 200; Follmann: Lich. Exs. Sel. No. 343; Kalb: hich.’ Neotr.s)No. 419, Dimerella isidiigera Vézda & Osorio, Vézda: Lich. Sel. Exs. Masco 4 2762 7(1989)).. ISOTYPE. Uruguay: Montevideo, Parque Rivera. Ad corticem Cupressi loco umbroso. H.S. Osorio, 12 October 1988 (CANL 100603). Vézda: Lich. Sel. Exs. No. 2329. Dimerella pocsii Vézda & Farkas, Folia Geobot. Phytotax., Prahay232) 193.7(1988)* ISOTYPE. Tanzania: Tanga regio, Usambara Orientalis, reservatum naturae Kihuhwi Forest dictum, secus trametem inter Kihuhwi et Wamkoro. Foliicola in pluviisilva. E. Farkas 86228, 3 November 1986 (CANL 97332). Vézda: Lich. Sel. Exs. Nowe2230. Dimerella subdentata Vézda & Thor, Vézda: Lich. Sel. Exs. Fasc. 94: 3 (1989). ISOTYPE. Australia: Queensland, circa 15 km ad occidentem et septentriones a Tully prope litorem Mission, secus Stony Creek. Ad corticem arboris in pluviisilva. G. Thor 5721, 28 November 1985 (CANL 100607). Vézda: Lich. Sel. Exs. No. 2333. Dimerella subfallaciosa Vézda & Farkas, Vézda: Lich. Sel. Exo. erasc .0:98's)2-7(01990). ISOTYPE. Tanzania: Regio Morogoro, montes Nguru, in valle "Dikurura", 3 km ad orientem a "Mhonda Mission". In pluviisilva montana. E. Farkas 89100, 22 March 1989 (CANL 103357). Vézda: Lich. Sel. Exs. No. 2429. Dimerella usambarensis Vézda & Farkas, Folia Geobot. Pauytotax., Praha 23, 196 9(1988).. ISOTYPE. Tanzania: Tango regio, Usambara Orientalis, reservatum naturae Marvera Forest dictun, 6 km ad septentriones et orientem ab Amani. Foliicola. E. Farkas & T. Pécs 86228, 12 December (November in protologue) 1986 (CANL 67333) .)VézdasiLichwesel J iExe si Na. i2230". Die Diploschistes montevidensis H. Magn., Medd. Gdteb. Bot. Trddg. 18: 223 (1950). ISOTYPE. Uruguay: Montevideo, Pajas Blancas. Sobre la tierra. H.S. Osorio 1621, 20 February 1949 (CANL 2236). = D. conceptionis Vainio Diploschistes sbarbaronis B. de Lesd., Bull. Soc. Bot. France 99: 147 (1952). ISOTYPE. Italy: Toscana, Siena, Latericola in villa Solaia (Monteliscai). Sbarbaro, May 1951 (CANL 2241). = D. actinostomus (Pers. ex Ach.) Zahlbr. Diploschistes straminescens Zahlbr., Mycologia 22: 71 (1930). ISOLECTOTYPE. Puerto Rico: Mayaguez. Open road-side on clay bank. B. Fink 1016, 18 December 1915 (CANL 2279). = D. hypoleucus Zahlbr. Dirina californica Tuck., Lich. Calif. p. 422 (1866). ISOLECTOTYPE. United States: California, Oakland. H.N. Bolander 129, 1866 (CANL 68755). = Schismatomma californicum (Tuck.) Zahlbr. Dolichocarpus chilensis R. Sant., Svensk Bot. Tidskr. 43: 552 (1949). ISOTYPE. Chile: Prov. Coquimbo, Coquimbo. On spines of Cereus in dry hill near the sea-shore. R. Santesson 2620a, 1 August 1940 (CANL 2113). Endocarpon mailae Lamb, Lilloa 14: 213 (1948). PARATYPE. Argentina: Prov. Tucuman, Sierra de Javiev, E. side. On rocks at roadside. I.M. Lamb 5096, 14 September 1947 (CANL 1197). Endocarpon tuckermanii Rav. ex Mont., Sylloge Gener. Spec. Cryptog.: 359 (1856). ISOTYPE. United States: South Carolina, Santee Canal. On Carya sp. among mosses. H.W. Ravenel 138 (CANL 1180). Reliq. Tuck. No. 42. = Catapyrenium tuckermanii (Rav. ex Mont.) Thomson Evernia perfragilis Llano, J. Wash. Acad. Sci. 41: 199 (1951). (see Alectoria arctica Elenk. & Savicz). Everniastrum fragile Sipman, Proc. Kon. Ned. Akad. Wetensch. Ser. C. 83: 348 (1980). ISOTYPE. Colombia: Caldas, below hotel Termales, Nevado del Ruiz. On grazed secondary shrub vegetation. H. Sipman & H. Valencia 10512, 4 February 1979 (CANL 91358). Hale: Lich. Amer. Exs. No. 235. Fellhanera endopurpurea Hafellner & Vézda, Nova Hedwigia 52: 76 (1991). ISOTYPE. Australia: New South Wales, Border Ranges National Park, Wiangaree, Brindle Creek. Foliicola in silva (Nothofagus moorei). J. Hafellner, P. Merotsy & R. Rogers, 30 313 August 1986 (CANL 103382). Vézda: Lich. Sel. Exs. No. 2454. Fulgensia canariensis Follm. & Poelt, Philippia 4: 370 (1981). ISOTYPE. Canary Islands: Tenerife, kleinfeldrig auf lichtoffenen, weitgehend ebenen oder massig geneigten flachgriindigen Léssinseln zwischen Hartlaubstrauchern auf Bergriicken im Diploschistetum albescentis Klem. G. Follmann, March 1980 (CANL 77635). Follmann: Lich. Exs. Sel. No. 345. Glyphis achariana Tuck., Amer. J. Sci. Arts, ser. 2, 25: 429 (1858). ISOSYNTYPE. United States: South Carolina, Santee Canal. H.W. Ravenel, 4 February 1858 (CANL 68731). Reliq. Tuck. No. 45. Gonohymenia cribellifera ssp. macrocarpa Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 6 (1990). ISOTYPE. Canary Islands: Fuerteventura, Sukkulentenhalbwtiste auf basisch-kristallinem Gestein, Passstrasse zwischen Betancuria und Pajara nahe Vega del Rio de las Palmas; auf sonnenexponierten Sickerwasserflachen bei 550-580 m. A. Henssen 30731la, 31.1/1.2. 1986 (CANL 102543). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 37. Gonohymenia lusitanica Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 7 :(1990). ISOTYPE. Portugal: Baixo-Alentejo, Garique der Serra da Arrabida; auf Sickkerwasserstreifen und in Mulden auf §S exponierten Kalkfelsen in Meeresnahe. A. Henssen & D. Schafer 21775g, 9 August 1971 (CANL 102545). Henssen: Lich. Cyanoph Fungi Sax. Exs. No. 39. Gonohymenia undulata Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc.,.2: 5 (1990). ISOTYPE. Canary Islands: Fuerteventura, Sukkulentenhalbwiste auf basisch-kristallinem Gestein, Passstrasse zwischen Betancuria und Pajara nahe Vega del Rio de las Palmas; auf sonnenexponierten Sickerwasserflachen bei 550-580 m. A. Henssen 30736b, 31.1/1.2.1986 (CANL 102540). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 35. PARATYPE. Canary Islands. A. Henssen 32048a (CANL 102541). Graphina aibonitensis Fink, Mycologia 19: 215 (1927). ISOTYPE. Puerto Rico: Aibonito. In open woods on bark. B. Fink 2017, 1 July 1916 (CANL 2085). Graphina muscicola Kalb, Kalb: Lich. Neotr. Fasc. 10: 8 (1988). ISOTYPE. Ecuador: Napo, Bei Sarayacu, zwischen Tena und Baeza. Ad einem feuchten Erdanriss, tiber Moosen. K. & A. Kalb, 10.9.1987 (CANL 102711). Kalb: Lich. Neotr. No. 422. Graphis subvirginalis Nyl., Bull. Torrey Bot. Club 16: 106 (1889). ISOTYPE. United States: Florida, Fort George. On Ilex. W.W. Calkins, 1888 (CANL 2094). 314 = Graphina subvirginalis (Nyl.) Mull. Arg. Gyalectidium colchicum Vézda, Folia Geobot. Phytotax., Praha 18: 58 (1983). ISOTYPE. Russia: Colchis (Transcaucasia occid.) distr. Lazarevskoje: in faucibus rivi Dagomys Zapadnyi, supra vicum Treja Rota. Ad folia Buxi colchicae. A. Vézda, 25 June 1979 (CANL 85979). Vézda: Lich. Sel. Exs. No. 1866. Gyalideopsis alnicola Noble & Vézda, Folia Geobot. Phytotax., Bohemoslov. 14: 62 (1979). ISOTYPES. Canada: British Columbia, Saltspring Island, road to Mount Maxwell. On Alnus rubra. W.J. Noble 5811A, 3 (30 in protologue) June 1976 (CANL 75390 & 75391). Gyalideopsis kalbii Vézda, Mitt. Bot. Staatssamml. Mtinchen 19: 153 (1983). ISOTYPES. Brazil: Sado Paulo, Ilha de SAo Sebastido, etwa 130 km 6stlich von S&ao Paulo. Osthang des Morro das Tacas; tber Moosen, an einem Wegabstich. K. Kalb & J. Poelt, 7 July 1979 (CANL 89171 & 89439). Vézda: Lich. Sel. Exs. No. 1965; Kalb: Lich. Neotr. No. 323. Gyalideopsis trapperi Kalb & Vézda, Biblioth. Lichenol. 29: 49 (1988). ISOTYPE. Ecuador: Napo, Bei Sarayacu, zwischen Tena und Baeza. Als Erstbesiedler auf herumliegenden Urgesteinsbloécken. K. & A. Kalb, 10.9.1987. (CANL 102712). Kalb: Lich. Neotr. No. 423. Gyalideopsis vezdae Kalb, Kalb: Lich. Neotr. Fasc. 6: 10 (1983). ISOTYPE. Brazil: Mato Grosso, Serra dos Coroados, Buriti, Naturschutzgebiet der evangelischen Schule von Buriti, etwa 6 km sltidwestlich von Buriti. In einem submontanen Regenwald. K. Kalb, 8 July 1980 (CANL 84778). Kalb: Lich. Neotr. No. 2297 Gyrophoropsis dwaliensis R&dsdnen, Arch. Soc. Zool.-Bot. Fenn. "Vanamo" 6: 80 (1952). ISOTYPE. India: N. W. Himalaya, Dwali, Almora District. 8500 ft. D.D. & A.M. Awasthi, 21 May 1950 (CANL 10771). | = Umbilicaria badia Frey Haematomma inexpectatum Kalb, Kalb: Lich. Neotr. Fasc. 12: 11 (1991). ISOTYPE. Mexico: Baja California Sur. Along Rte 22; 16 km W of Ciudad Constitui6én, Southern Magdalena Region of the Socnoran Desert, sandy coastal plain. On bark of deciduous trees. K. & A. Kalb & T. Nash, 1 January 1991 (CANL 103777). Kalb: Lich. Neotr. No. 506. Haematomma pustulatum Brodo & Culb., Bryologist 89: 203 (1986). ISOTYPE. United States: North Carolina, Orange Co., Hillsborough, Duke Forest (Quarry Division); near the quarry. 315 On Acer rubrum. W.L. Culberson & C.F. Culberson 19701, 22 November 1984 (CANL 96379). Vézda: Lich. Sel. Exs. No. 2152. Hafellia fosteri Imsh. & Sheard, Bryologist 95: 85 (1992). PARATYPE. Canada: British Columbia, Discovery Island, 3 km east of Oak Bay, Victoria. On driftwood. A.M. Crossley & W.J. Noble 5335a, 3 August 1975 (CANL 102584). Heppia uruguayensis Rasdnen, Arch. Soc. Zool.-Bot. Fenn. "Vanamo" 2: 48 (1947). HOLOTYPE. Uruguay: Dept. PaysandtG. On rocky banks of river Queguay near Paysandda. I.M. Lamb 3107, 10 March 1946 (CANL 983). = Thelidium umbilicatum Th. Fr. Heterodermia desertorum Kalb, Kalb: Lich. Neotr. Fasc. 8: 10 (1984). ISOTYPE. Peru: Lima, Ktistenwliste bei Huacho, etwa 150 km nérdlich von Lima. An kleinen Felsbrocken in der Nebelzone. K. Kalb & G. Plébst, 28 July 1979 (CANL 89440). Kalb: Lich. Neotr. °No: 324. Heterodermia sitchensis Goward & Noble ex Goward, Bryologist 87: 366 (1984). ISOTYPE. Canada: British Columbia, Vancouver Island, 12 km SE of Tofino, Schooner Cove. On twigs of Picea sitchensis. T. Goward, Kirkvold & McGrenere 83-326, 30 March 1983 (CANL 95990). Heterothecium pachycheilum Tuck., Synops. N. Amer. Lich. 2: 56 (1888). ISOSYNTYPE. United States: South Carolina, Santee Canal. H.W. Ravenel, 1857 (CANL 68775). Reliq. Tuck. No. 106. = Megalospora pachycheila (Tuck.) Sipman Homothecium sorediosum Henssen, Bot. Not. 132: 271 (1979). ISOTYPE. Chile: Malleco, Parque Nacional de cContulmo, Stidbuchenwald mit Nothofagus obliqua; auf Erde und briichigem Sandstein einer Wegbéschung. A. Henssen, G. Vobis & J. Red6én 24271, 22 February 1973 (CANL 102536). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 31. Hubbsia lumbricoides W. Weber, Svensk Bot. Tidskr. 59: 61 (1965). ISOTYPE. Mexico: Baja California, Isla Guadalupe. On seaward- facing basalt cliffs at Melpomene Cove, southernmost tip of the island. W.A. Weber & C.J. McCoy, 21 April 1963 (CANL 20874). Weber: Lich. Exs. Colo. No. 85. = H. californica (Rasdnen) W. Beber Huilia circumnigrata Vézda, Folia Geobot. Phytotax. Praha 14: 205 (1979). ISOTYPE. Jugoslavia: Montenegro, Sinus Kotor dictus, supra pagum Morin. Ad saxa quartzitica. A. Vézda, 18 July 1975 (CANL 73828). Vézda: Lich. Sel. Exs. No. 1665. 316 Hydrothyria venosa J. Russell, Proc. Essex Inst. 1: 188 (1853). ISOSYNTYPE. United States: Vermont, Brattleboro. In rivulets from Wantasquit Mt. J.L. Russell, 16 July 1851 (CANL 23259). Relig sy LUCK eaN On 1, Hyperphyscia mobergii Kalb, Kalb: Lich. Neotr. Fasc. 11: 9 (1990). ISOTYPE. Venezuela: Falcon, Distr. Acosta, Zwischen Boca da Tocuyo und San Lorenzo, etwa 20 km NW von Chichiriviche. K. & A. Kalb, 24 August 1989 (CANL 102751). Kalb: Lich. Neotr. No. 463. Hypocenomyce leucococca R. Sant. in Moberg, Publ. Herb. Univ. Uppsala, Thunbergia 2: 3 (1986). ISOTYPE. Sweden: Harjedalen Prov., Tanndas par., c. 1 km E of Ramundbergets Fjdllgard. On the trunk of a birch. R. Santesson 27901, 5 August 1977 (CANL 95082). Moberg: Lich. Sel. Exs.:Upss No.6. Hypogymnia heterophylla Pike, Mycotaxon 16: 157 (1982). ISOTYPE. United States: California, Mendocino Co., Pygmy Forest Reserve, Van Damme State Park. On stunted conifers. M.E. Hale 49365, 9 June 1977 (CANL 91360). Hale: Lich. Amer. Exs.: No. 237... Hypogymnia hokkaidensis Kurok., Mem. Natl. Sci. Mus. (Tokyo) 427620 (1970). ISOTYPE. Japan: Hokkaido, Prov. Nemuro, Ochiishi. On trunk of Picea glehnii. S. Kurokawa 65826, 2 September 1965 (CANL 45315). Kurokawa: Lich. Rar: Crit. Exs: No. 167. Hypogymnia kiboensis Dodge, Ann. Missouri Bot. Gard. 46: 50 (1959). ISONEOTYPE. Tanzania: Kilimanjaro, above Peter's Hut. In the upper part of the Philippia region, on "frost-lifted ground". O. Hedberg 1354, 23 June 1948 (CANL 99605). Moberg: Lich. Sel. Exs. Ups. No. 75. = Xanthoparmelia kiboensis ( Dodge) Krog & Swinscow Hypogymnia mollis Pike & Hale, Mycotaxon 16: 161 (1982). ISOTYPE. United States: California, San Luis Obispo Co., Los Osos Oak Reserve, near Los Osos. On low shrubs in sandy area. M.E. Hale 57768, 31 July 1980 (CANL 91361). Hale: Lich. Amer. Exs. No’ 238. Hypogymnia oceanica Goward, Bryologist 91: 229 (1988). HOLOTYPE. Canada: British Columbia, Vancouver Island, 5 km SW of Sutton Pass. Trunk and branches of Pinus contorta. T. Goward 83-302, 20 March 1983 (CANL 87115). PARATYPE. Canada: British Columbia. I. M. Brodo 18066. (CANL 69916). Involucrothele ligurica Servit, Webbia 10: 443 (1954). ISOTYPE. Italy: Liguria occid., Spotorno (Savona), Loco Lajolo. Maceriicola. Sbarbaro, March 1948 (CANL 985). 317 Ionaspis ventosa P. Jgorg. & R. Sant., Nord. J. Bot. 9: 433 (1989). ISOTYPE. Sweden: Hdrjedalen Prov., Tannds par., Mt. Réstvalen (c. 5 km W of Mt. Stora Mittakldppen). On pebbles at top of the mountain. R. Santesson 32679, 14 August 1989 (CANL 102824). Moberg: Lich. Sel. Exs. Ups. No. 85. Jenmania osorioi Henssen, Lichenologist 5: 447 (1973). ISOTYPE. Uruguay: Salto, Uruguay River, Salto Grande Island. Auf dem oberen Teil eines grossen Steins unterhalb der Hochwasserlinie. H. Osorio 6675, 4-6 February 1974 (CANL 102583). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 29. Kalbiana brasiliensis Henssen, Kalb: Lich. Neotr. Fasc. 10: 9 (1988). ISOTYPE. Brazil: Bahia, Chapada Diamantina, Serra do Tombador, etwa 1 km vor der Stadt Morro de Chapea. Am Rand von natiirlichen, kleinen Wasserbecken, an Sandstein. K. Kalb, 20 July 1980 (CANL 102715). Kalb: Lich. Neotr. No. 426. Lecanactis premnea var. chloroconia Tuck., Gen. Lich. p. 194 (1872). ISOSYNTYPE. United States: New Hampshire, Mt. Washington. E. Tuckerman, September 1848 (CANL 68776). Reliq. Tuck. No. 107. = L. chloroconia Tuck. Lecanactis subdryophila Follm. & Vézda, Philippia 3: 271 (1977). ISOTYPE. Chile: Prov. Aconcagua. Kieternpflanzung im Windschatten der Isla Seca in der Bucht von Zapallar. G. Follmann, January 1965 (CANL 67506). Follmann: Lich. Exs. Sel. No. 247. Lecania constricta W. Weber, Weber: Lich. Exs. Colo. Fasc. P60 V3. (L984) . ISOTYPE. United States: Colorado, Boulder Co., Outer foothills of the Front Range, "The Flatirons", on southwest edge of City of Boulder. On bark of Populus angustifolia just S of mouth of Gregory Canyon. W.A. Weber, J.W. Sheard & R.A. Anderson, 5 September 1981 (CANL 88658). Weber: Lich. Exs. Colo. iNo. 614. Lecania macrocarpa B. de Lesd., Bull. Soc. Bot. France 99: 146 (1952). ISOTYPE. Italy: Gorzegno, Le Langhe (Piemonte). Sbarbaro 319, September 1951 (CANL 12830). Lecania sbarbaronis B. de Lesd., Bull. Soc. Bot. France 97: 170 (1950). ISOTYPE. Italy: Varazze per Invrea. Maceriicola in via publica. Sbarbaro, June 1949 (CANL 12836). Lecanora aeruginascens H. Magn., Medd. Géteb. Bot. Trddg. 18: 26) 41950). ISOTYPE. Uruguay: Montevideo, Carrasco, Cafiada de las Cantberas. Sobre rocas expuestas al sol. H. Osorio, 17 318 January 1948 (CANL 11644). Lecanora austrooceanica Hertel & Leuck., Hertel: Lecideaceae Exs. Fasc. “12:2 (1990). ISOTYPE. New Zealand: Southland. Bluff, S of Ocean Beach. Coastal rocks. H. Hertel 30808, 12 November 1985 (CANL 102505). Hertel: Lecideaceae Exs. No. 224. Lecanora bolanderi Tuck., Proc. Amer. Acad. Arts 6: 266 (1864). LECTOPARATYPE. United States: California, Marin County. H.N. Bolander (CANL 68831). = Cladidium bolanderi (Tuck.) Ryan Lecanora caesiorubella subsp. merrillii Imsh. & Brodo, Nova Hedwigia 12: 28 (1966). ISOTYPE. United States: California, Berkeley. M.A. Howe, February 1893 (CANL 19545). Cummings: Dec. N. Amer. Lich. No. ABS YS). Lecanora caesiosora Poelt, Denkschr. Regensb. Bot. Ges 26: 82 (1966). (see Lecanora soralifera H. Magn.). Lecanora californica Brodo, Beih. Nova Hedwigia 79: 107 (1984). HOLOTYPE. United States: California, San Mateo Co. Rocks near the sea. M.A. Howe, February 1893 (CANL 19543). Cummings: Dec. N. Amer. Lich. No. 131. PARATYPES. United States: California. I.M. Brodo 20455 (CANL 76211); Oregon. I.M. Brodo (CANL 77027). Lecanora christoi W. Weber, Bryologist 78: 206 (1975). ISOTYPE. United States: Colorado, Garfield Co. 8 km N of Rifle. Sandstone ridge on west side of Rifle Creek along south edge of Grand Hogback 3.2 km SW of Rifle Gap. On talus blocks. W.A. Weber, G. Kunkel & J. Munger, 12 June 1974 (CANL 52208). Weber: Lich. Exs. Colo. No. 458. Lecanora cinereofusca var. appalachensis Brodo, Beih. Nova Hedwigia 79: 116 (1984). HOLOTYPE. United States: Arkansas, Newton Co., Lost Valley State Park, 22 miles SW of Harrison. On sandstone block at edge of brook. I.M. Brodo 23512, 11 August 1979 (CANL 76592). PARATYPE. Canada: Nova Scotia. J. Macoun (CANL 11825). Lecanora concinna ("concinnum") Thomson, Bryologist 75: 358 (1972). ISOTYPE. United States: Alaska, north end at Franklin Bluffs on the Sagavanirktok River. On smooth boulders on gravel bench. J.W. Thomson, S. Shushan & J. Koranda, 1958 (CANL 44988). Thomson: Lich. Arct. No. 95. = a species of Aspicilia Lecanora congesta Clauz. & Vézda, Portugaliae Acta Biol., ser. ‘By (9% -332 (1969). O19 PARATYPE. France: Gallia, Bouches-du-Rhéne, Marseille, insula Maire. Ad saxa calcarea maritima, in septentrionem spectantia. G. Clauzade, June 1968 (CANL 55614). Vézda: Lich. Sel. Exs. No. 779. Lecanora freyi Poelt, Mitt. Bot. Staatssamml. Miinchen 2: 464 (1958). ISOTYPE. Austria: Bregenzer Wald, Ifenstock, Vorarlberg (Kleines Walsertal): Kleiner Gipfel am H&ahlekopf bei etwa 2000 m, stidseitig auf Kieselkalk. J. Poelt, July 1958 (CANL 52798). Poelt: Lich. Alpium No. 73. Lecanora fuliginosa Brodo, Beih. Nova Hedwigia 79: 124 (1984). HOLOTYPE. Canada: Ontario, Thunder Bay District, Sibley Peninsula, 1.3 miles east of Silver Islet. On exposed rock on bluff. I.M. Brodo 13735, 13 June 1968 (CANL 77033). PARATYPES. Canada: New Brunswick. S.P. Gowan 3588 (CANL 82233); Ontario. I.M. Brodo 5941 (CANL 22789), A. Henssen 14029c (CANL 77237); Quebec. I.M. Brodo 18841 (CANL 38740), E. Lepage 16766 (CANL 34969); Northwest Territories. J.wW. Thomson 12087 (CANL 23472). Lecanora homalea H. Magn., Medd. Géteb. Bot. Trddg. 18: 217 (1950). PARATYPE. Uruguay: Montevideo, Carrasco, Cafiada de las Canteras. Sobre postes de madera. H. Osorio 1761, 17 January 1948 (CANL 12081). Lecanora imshaugii Brodo, Beih. Nova Hedwigia 79: 137 (1984). Holotype. Canada: Ontario, Ottawa, Rockcliffe. On a pine trunk. J. Macoun 345, 16 April 1896 (CANL 11626). PARATYPES. Canada: Ontario. J. Macoun 347 (CANL 12443); United States: Delaware. I.M. Brodo 9558 (CANL 21882), Virginia. I.M. Brodo 9435B (CANL 21780). Lecanora insolata H. Magn., Kongl. Svenska Vetenskapsakad. Hand tge1 727-137 3(1939).. ISOTYPE. Sweden: Vdastergétland, G6dteborg, Delsj6karr. On steep, sunny rocks in the heath. A.H. Magnusson, 11-10-1937 (CANL 20146). Magnusson: Lich. Sel. Scand. Exs. No. 323. Lecanora kutakii Senft, Preslia 4: 26 (1926). ISOTYPE. Czechoslovakia: Sudeti occid., KrkonoSe, in valle Obfi dil loco dicto Kiesberg. V. Kuf&k, 1921 (CANL 33384). = L. gisleri Poelt & Ullrich Lecanora luteovernalis Brodo, Bryologist 84: 521 (1981). HOLOTYPE. Canada: Northwest Territories, Dist. of Franklin, Bathurst Island, Polar Bear Pass area. I.M. Brodo & R.R. Ireland 19435, 23 July 1973 (CANL 59491). PARATYPES. Canada: Yukon. G.W. & G.G. Douglas 8895 (CANL 56763); G.W. Scotter 19127, 19252, 19257, 19487, 19525, 19530 & 19531 (CANL 50183, 49740, 49739, 54313, 54315, 54316 & 54317); Northwest Territories. P. Barrett 0807. (CANL 34299); G.R. Brassard 4253B (CANL 28932); I.M. Brodo 19388 (CANL 320 53565); S.D. MacDonald (CANL 53033); G.R. Parker SP-70-184C (CANL 33112); R.M. Schuster 35941 (CANL 5572); G.W. Scotter 18487 (CANL 70135); J.W. Thomson 8689 (CANL 45361). Lecanora mellea W. Weber, Bryologist 78: 208 (1975). ISOTYPE. United States: California, Sacramento Co., SE of Sacramento between Slough House and Bridge House. On sloping faces of rock outcrops. W.A. Weber & S. Kipp, 24 March 1974 (CANL 52212). Weber: Lich. Exs. Colo. No. 462. Lecanora microbola Lamb, Rep. (Annual) Natl. Mus. Canada, 1952-53. Bull. 132: 291 (1954). HOLOTYPE. Canada: Nova Scotia, Cape Breton Co., Cape Breton Island, Louisbourg. On large stone near seashore. I.M. Lamb 6897, 19 July 1952 (CANL 12151). Lecanora oleae ("Yolea") Reich. & Galun, Bull. Res. Council Israel Sect. D, Bot. 6D: 238 (1958). SYNTYPE. Israel: Thalpioth (Jerusalem). On Olea europea. I. Reichert 8(1), 16 October 1933 (CANL 32294). Lecanora opiniconensis Brodo, Mycotaxon 26: 309 (1986). HOLOTYPE. Canada: Ontario, Leeds Co., Snake Island in Lake Opinicon. Queen's University Biological Station, Chaffeys Locks. I.M. Brodo 25117, 2 February 1985 (CANL 92112). PARATYPES. Canada: Ontario. I.M. Brodo 456, 13575B, 20277, 23432B & 23543 (CANL 23295, 79030, 53929, 72543 & 80250); R.F. Cain 26672 (CANL 44587); H.L. Dickson 154 (CANL 57666) ; C.E. Garton 5467, 6089 & 22268 (CANL 12295, 12304 & 87731); C.M. Wetmore 28714 & 29679 (CANL 85169 & 85259). Quebec. I.M. Brodo 21015 (CANL 54173). United States: Maine. S. Selva 990 (CANL 98807). Michigan. C.M. Wetmore 47222 (CANL 90696). Minnesota. C.M. Wetmore 34532 & 35014 (CANL 75807 & 75810). S. Dakota. C.M. Wetmore 8760 (CANL 12150). Wisconsin. T.S. & B.A. Cochrane 5601 (CANL 45366). Lecanora orae-frigidae R. Sant., Lich. Sweden Norway, p.145 (1984). (see Lecidea sorediata Lynge). Lecanora pachysoma Ryan & Poelt, Bryologist 92: 518 (1989). PARATYPES. Mexico. T.H. Nash 8804. (CANL 52389); W.A. Weber & C.J. McCoy. (CANL 20876). Weber: Lich. Exs. Colo. No. 87. Lecanora perplexa Brodo, Beih. Nova Hedwigia 79: 148 (1984). HOLOTYPE. Canada: Quebec, Gatineau Co., Gatineau Park, King Mountain area off Mountain Road, 3 mi. NW of Notch Road intersection. On limestone cliff. I.M. Brodo 21192, 6 October 1975 (CANL 67214). PARATYPES. Canada: Ontario. I.M. Brodo 6070 (CANL 22769), A. Henssen 14276 (CANL 77235), J. Macoun? (CANL 11624); Quebec. I.M. Brodo 23585 (CANL 73901); United States: Arkansas. I.M. Brodo 13504 (CANL 76596). Lecanora pseudopinguis W. Weber, Mycotaxon 13: 102 (1981). ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz. On 321 rocks just above high tide. W.A. Weber & J. Lanier, 10 April 1976 (CANL 58215). Weber: Lich. Exs. Colo. No. 500. Lecanora pycnocarpa H. Magn., Bot. Not. 1939: 310 (1939). ISOTYPE. Sweden: Uppland, Vallentuna, Abydal. On Populus tremula. A.H. Magnusson, 12 June 1934 (CANL 20132). Magnusson: Lich. Sel. Scand. Exs. No. 305. Lecanora salicicola H. Magn., Bot. Not. 1939: 311 (1939). ISOTYPE. Sweden: Jamtland, Are, Skalstugan. On branches of dying Salix. C. Stenholm, 28 July 1933 (CANL 20133). Magnusson: Lich. Sel. Scand. Exs. No. 306. Lecanora soralifera H. Magn., Bot. Not. 1937: 135 (1937). ISOTYPE. Sweden: Vastergétland, Partille, northwest of Tultered. On stone fence. A.H. Magnusson, 27 August 1936 (CANL 20111). Magnusson: Lich. Sel. Scand. Exs. No. 270. = L. caesiosora Poelt Lecanora superfluens H. Magn., Acta Horti Gothob. 19: 46 (1952). ISOTYPE. Canada: Northwest Territories, Baffin Island. Head of Clyde Fiord. On wet soil near large boulders. M.E. Hale 416, 26 June 1950 (CANL 12472). = L. geophila (Th. Fr.) Poelt Lecanora texana W. Weber, Mycotaxon 13: 103 (1981). ISOTYPE. United States: Texas, Gillespie Co. (not "Llano Co. -as given on the label), Texas Hill Country, trail to Balanced Rock, 6.4 km N of Fredericksberg. On vertical N-facing of large blocks. W.A. Weber, 29 April 1974 (CANL 52201). Weber: Rich..Exs.,.Colo.iNo. 451. Lecanora thallophila H. Magn., Bot. Not. 107: 196 (1954). ISOTYPE. United States: Utah, San Juan Co., Devil's Canyon. On Dermatocarpon miniatum on dry exposed sandstone. S. Flowers 476, 21 June 1952 (CANL 71524). Lecanora utahensis H. Magn., Acta Horti Gothob. 19: 39 (1952). ISOTYPE. United States: Utah, Wayne Co., Ekker's Ranch. On dry exposed sandstone. S. Flowers 3107, 20 May 1951 (CANL 12503)%% Lecanora weberi Ryan, Mycotaxon 36: 10 (1989). ISOTYPE. United States: Colorado, Boulder Co., Boulder Mountain Parks, just S of city of Boulder. On scattered low fine-grained sandstone or quartzite boulders. W.A. Weber, 10 August 1986 (CANL 97984). Weber: Lich. Colo. Exs. No. 685. Lecidea baffiniana H. Magn., Acta Horti Gothob. 19: 43 (1952). ISOTYPE. Canada: Northwest Territories, Baffin Island, Cape Searle. On phonolite at the bird cliffs. M.E. Hale 391, 17-18 August 1950 (CANL 4786). 322 Lecidea carbonoidea Thomson, Bryologist 75: 356 (1972). ISOTYPE. United States: Northern Alaska, along the Pitmegea River, 15 miles upstream from Cape Sabine. On top of ridge. J.W. Thomson, 10-17 July 1958 (CANL 44987). Thomson: Lich. Arct. No. 94. = Immersaria carbonoidea (Thomson) Esnault & Roux Lecidea grassiana H. Magn., Medd. Gdéteborgs Bot. Trddg. 17: 60 (1947). ISOTYPE. Argentina: Prov. Salta, Quebrada de San Lorenzo. M. Digilio-Grassi 336, 1946 (CANL 5061). Lecidea gypsicola Llimona, Vézda: Lich. Sel. Exs. Fasc. 47: She GLI LES) is ISOTYPE. Spain: Las Bardenas, Caparosso. Ad gypsum compactum locis septentr. vel occident. spectantibus. X. Llimona, 24 September 1972 (CANL 77198). Hertel: Lecideaceae Exs. No. 45. Lecidea (Psora) hyotheja Lamb, Farlowia 4: 444 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near Futalaufquen, On soil in crevices of an almost vertical rock on the shore of the lake. I.M. Lamb 5870, 1 February 1950 (CANL 5072). PARATYPE. Argentina: Lago Cisnes. I.M. Lamb 5932 (CANL 5071). Lecidea leproloma R. Sant., Lich. Sel. Exs. Upsal., Thunbergia 2: 6 (1986). ISOTYPE. Sweden: Hadrjedalen Prov., Tann&s par., Mt Gruvvalen, 0.2 km SSW of Lake Glimsj6n (c. 1.8 km NE of Ramundbergets Fjallgard). On a vertical rock wall. R. Santesson 31390, 19 August 1985 (CANL 95090). Moberg: Lich. Sel. Exs. Ups. No. 14. Lecidea patagonica Lamb, Farlowia 4: 440 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Pathway from Lago Frias to the Frias Glacier. On a stone beside the path in the forest. I.M. Lamb 6029, 13 November 1950. (CANL 5290). PARATYPE. Argentina: Lake Nahuel Huapi. I.M. Lamb 5818 (CANL 5289). Lecidea santensis Tuck., Amer J. Sci. Arts ser. 2, 25: 428 (1858). SYNTYPE. United States: South Carolina. H.W. Ravenel (CANL 23244)|. Relig. “Tuck. No. 15. = Phyllopsora corallina (Eschw.) MUll. Arg. Lecidea shushanii Thomson, Bryologist 75: 356 (1972). ISOTYPE. United States: Northern Alaska, valley of the Okpilak River at Okpilak Lake near Mt. Michelson. On huge boulder at top of ridge. J.W. Thomson & S. Shushan 9316, 3 August 1958 (CANL 44982). Lecidea sorediata Lynge, Novaya Zemlya 43: 117 (1928). ISOLECTOTYPE. Russia: Novaya Zemlya, Nordre Kristovii holmen. B. Lynge, 14 August 1921. (CANL 5430). = Lecanora orae-frigidae R. Sant. 323 Lecidea ullrichii Hertel, Hertel: Lecideaceae Exs. Fasc. 10: 4 (1988). ISOTYPE. Germany: Niedersachsen, Harz Ramelsberg bei Goslar. H. Ullrich, 26 October 1982 (CANL 96965). Hertel: Lecideaceae Exs. No. 190. Lempholemma dispansum H. Magn., Bot. Not. 1939: 302 (1939). ISOTYPE. Sweden: Dalsland, Backe, Karud. On sunny, slightly irrigated, calcareous rock. S. Bergstrom & A.H. Magnusson, 24 June 1938 (CANL 20128). Magnusson: Lich. Sel. Scand. Exs. No. 298. Leproloma diffusum var. chrysodetoides Laundon, Lichenologist 21: 18 (1989). ISOTYPE. Sweden: Vdarmland Prov., Burtjdarn par., on the western slope of the hill between Irettondetjdrnen and Svarttjdrnen. Over mosses on perpendicular rock. L.E. Muhr 598, 8 November 1977 (CANL 99591). Moberg: Lich. Sel. Exs. Ups. No. 61. Leptogium caesiellum Tuck., Synopsis N. Amer. Lich. 1: 156 (1882). ISOTYPE. United States: Illinois, Athens. EF. Hall, 1876 (CANL 23261). eReliq .etTucka No1w53.% = L. byssinum (Hoffm.) Zwackh ex Nyl. Letharia togashii Asah., J. Jap. Bot. 27: 295 (1952). ISOTYPE. Japan: Honshu. Prov. Kai, Oshino-mura, Minami-Tsuru- gun. On twigs of Picea polita. M. Togashi, 11 August 1952 (CANL 26404). Kurokawa: Lich. Rar. Crit. Exs. No. 66. Letharia wandelensis Hue, Expedit. Antarct. Fran¢. 1903-1905, P7568 61908 ),. ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth (Wandel) Island. Expedit. Antarct. Fran¢. 1903-1905, No. 277 & 299 (CANL 16959). Lichen candelarius L., Spec. Plant 2: 1141, 1753. ISONEOTYPE. Sweden: Oland Prov., Boda par., Mensalvaret. On the top of granitic boulders. R. Santesson & T. Tonsberg 30905, 3 August 1983 (CANL 95101). Moberg: Lich. Sel. Exs. Ups. No 25. = Xanthoria candelaria (L.) Th. Fr. Lichen salazinus Bory, Voyage Quatre Iles d'Afrique 3: 106 (1804). ISOLECTOTYPE. Africa: Réunion (Bourbon) Island. Bory de St. Vincent (CANL 75659). = Stereocaulon salazinum (Bory) Fée Lichenoconium parasiticum D. Hawksw., Persoonia 9: 178 (1975 \e ISOTYPE. Denia: Sjaellandia, Herstedvester. In apotheciis Lecanorae conizaeoidis. M.S. Christiansen 11707, 20 April (July in protologue) 1944 (CANL 69519). Vézda: Lich. Sel. Exs. No. 1600. 324 Lichenostigma maureri Hafellner, Herzogia 6 : 299 (1982). PARATYPE. Italy: Siidtirol, Mendelgebirge SW von Bozen, Mte. Roén, Larchen-Fichtenwald auf der N-Seite. J. Hafellner, October 1976 (CANL 91566). Poelt: Pl. Graecenses No. 319. Lichina macrospora Henssen, Biidel & Wessels, Mycotaxon 22: 171 (1985). ISOTYPE. South Africa: Ost-Transvaal, Graskop 2430DD, Panorama Falls, Lookout Point. Auf Sandsteinplatten am Rande von Rinnsalen. A. Henssen & D. Wessels 28405a, 8 June 1982 (CANL 102537). Henssen: Lich. Cyanoph. Fungi sax. Exs. No. B2e Lichina polycarpa Henssen, Lichenologist 5: 450 (1973). PARATYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel Huapi, Isla Victoria. On semi-decomposed rock at side of lake. I.M. Lamb 5822 (=5821), 26 January 1950 (CANL 2358). Lichina rosulans Henssen, Lichenologist 4: 95 (1969). ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel Huapi, Isla Victoria. On rocks at shore of lake. I.M. Lamb 5847 (=5857), 28 January 1950. (CANL 27353) Lobaria fulva Hoffm., Deutschl. Flora, p. 159, 1796. ISONEOTYPE. Germany: Bavaria, Berchtesgadener Land, Untersch6nau, Astenreitweg-Zulehenweg. Ad truncum Aceris pseudoplatani. R. Ttirk & H. Wunder, 30 April 1991 (CANL 103428). Vézda: Lich. Sel. Exs. No. 2500. = Xanthoria fulva (Hoff.) Poelt & Petutschnig Lobaria fuscotomentosa Yoshim., J. Hattori Bot. Lab. 34: 311 (1971). ISOTYPE. Japan: Honshu, Prov. Echigo. On trail between the summit and Akayu, Mt. Naeba. On trunk of trees. S. Kurokawa 57194, 17 August 1957 (CANL 45320). Kurokawa: Lich. Rar. CYTG. EXs.-NOvelL 7 2. Lobaria gyrophorica Yoshim., J. Hattori Bot. Lab. 34: 275 (1971). ISOTYPE. Taiwan: Prov. Ilan, Piyanan Pass. On trees. S. Kurokawa 794, 16 January 1964 (CANL 68480). Kurokawa: Lich. Rare Crit. albxa.NOw 365" Lobaria tuberculata Yoshim., J. Hattori Bot. Lab. 34: 280 (1971). ISOTYPE. Japan: Hokkaido, Prov. Nemuro, Ochiishi. On trunk of Salix sp. S. Kurokawa 65767, 1 September 1965 (CANL 45326). Kurokawa: Lich. Rar. Crit. Exs. No 178. Maronella laricina M. Steiner, Osterr. Bot. Z. 106: 441 (1959). ISOTYPE. Austria: Wipptal, Tirol, Steinach-Mauern. Fussteil, NW-Flanke von Larix decidua an einem Rinnsal in Fettwiesen. M. Steiner, August 1958 (CANL 52819). Poelt & Steiner: Lich. Alpium No. 102. 325 Mazosia pilosa Kalb & Vézda, Folia Geobot. Phytotax., Praha 23: 203 (1988). ISOTYPES. Brazil: Amazonas, Manaus, in ripa fluminis Rio Negro, 100-200 km supra Manaus. Foliicola in pluviisilva. K. Kalb, 14 October 1980 (CANL 97329 & 102720). Vézda: Lich. Sel. Exs. No. 2227; Kalb: Lich. Neotr. No. 431. Mazosia pseudobambusae Kalb & Vézda, Folia Geobot. Phytotax., Praha 23: 207 (1988). ISOTYPE. Brazil: Sdo Paulo, Ilha de Sdo Sebastiao, 130 km ad orientem a Sdo Paulo. Folliicola in pluviisilva. K. Kalb, J. Poelt & H. Sipman, 6 July 1980 (CANL 97330). Vézda: Lich. Sel. Exs. No. 2228. Megalospora foersteri Kalb, Kalb: Lich. Neotr. Fasc. 11: 10 (1990). ISOTYPE. Venezuela: Merida, Distr. Libertador, Pico Espejo, SE von Mérida, Loma Redonda. Uber feuchten Moosen auf Lichtungen eines kleinen Restwdldchens. K. & A. Kalb, 10 August 1989 (CANL 102753). Kalb: Lich. Neotr. No. 465. Melanaria macounii Lamb, Rep. (Annual) Natl. Mus. Canada, 1952-53 BULL 1323328 6n41954) ¢ HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton Island, Franey Mountain trail near Ingonish. On trunk of Fagus. I.M. Lamb 6985, 6 August 1952 (CANL 11613). = Pertusaria macounii (Lamb) Dibben Melanaria macounii var. meizotoca Lamb, Rep. (Annual) Natl. Mus eCanada,.2952=53' abu) Lay 6132 2628 a Glo54) HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, Baddeck. JseMacoun, 1953) p.pw.W,).9) July) 1898 (CANL' {11479),. = Pertusaria macounii (Lamb) Dibben Melaspilea fuscolimitata Fink, Mycologia 19: 211 (1927). ISOTYPE. Puerto Rico: Yauco, Dry Hill. On bark. B. Fink 1480, 29 December 1915 (CANL 1974). Micarea muhrii Coppins, Bull. Brit. Mus. (Nat. Hist.), Bot. 11(2): 160 (1983). ISOTYPE. Sweden: Vermlandia, par. Lungsund, Punbd&dcken. Loco aperto ad truncum decorticatum supra rivum. L.E. Muhr 285 (2851 in protologue), 15 July 1980 (CANL 89160). Vézda: Lich. Sel. Exs. No. 1954. = Micaria vulpinaris (Nyl.) Muhr Microthelia atramentea Norman, Bot. Not. 1867: 88 (1867). ISOTYPE. Norway: Finmark, Tanafjord, Lille Lerpollen. J.M. Norman, 1866 (CANL 93368). Muhria urceolata P. Jgrg., Notes Roy. Bot. Gard. Edinburgh 44: 583 (1987). ISOTYPE. Sweden: Vdrmland. Dalby, Gravbdcken, 6 km N of Syssleback. L.E. Muhr 4208, 6 August 1981 (CANL 95302). Mycoglaena quercicola R. Harris, Michigan Bot. 12: 30 (1973). 326 ISOTYPE. United States: Iowa, Bremer Co., Iowa. B. Fink, 1895 (CANL 1266). Nephroma chubutense Lamb, Farlowia 4: 438 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Chubut, W end of Lago Menéndez in the lower reaches of the Torrecillas stream. On trunk of a small tree. I.M. Lamb 5916, 4 February 1950 (CANL TOZ25 7)es Nephroma kuehnemannii Lamb, Farlowia 4: 436 (1955). ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menéndez, Alerzal del Norte. O. Ktihnemann 5485, 1941 (CANL 3751). Nephroma lepidophyllum f. hypomelaenum ("hypomelaena") Rasdnen, Farlowia 4: 439 (1955). ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias, pathway alongside the Frias stream on trunk of Nothofagus pumilio. I.M. Lamb 6010, 12 February 1950 (CANL 3785). Nephroma occultum Wetm., Bryologist 83: 243, !980. ISOTYPE. United States: Oregon, Lane Co., 7 miles NE of Blue River, H.J. Andrews Experimental Forest, 2 miles NE of the junction of Blue River and Lookout Creek. On old growth Pseudotsuga menziesii branches. S. Sundberg 120, 19 February 1978 (CANL 75860). Neuropogon rohmederi Lamb, Lilloa 14: 158 (1948). ISOTYPE. Argentina: Chubut, Lago Futalaufquén. G. Rohmeder (Herb. Crypt. Inst. Lillo no. 2275), 1945. (CANL 17205). = Usnea perpusilla (Lamb) F. J. Walker Neuropogon rohmederi f. ushuaiensis Lamb, Lilloa 14: 160 (1948). ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, ca. 20 km ENE of Ushuaia. On rocks. R. Santesson 640a p.p., 1940 (CANL 17209). = Usnea perpusilla (Lamb) F. J. Walker Neuropogon trachycarpus f. elatior Lamb, Lilloa 14: 157 (1948). ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, southern slope, above Las Cotorras (about 20 km ENE of Ushuaia). On rocks in the alpine region. R. Santesson 640b, 1940 (CANL 17258). = Usnea trachycarpa (Stirt.) MUll. Arg. Ochrolechia gowardii Brodo, Canad. J. Bot. 69: 746 (1991). HOLOTYPE. Canada: British Columbia, Wells Gray Park, Wavy Mt. near Murtle Lake. On branch of Abies lasiocarpa. T. Goward 86-39-5, 13 August 1980 (CANL 96072). Ochrolechia juvenalis Brodo, Canad. J. Bot. 69: 749 (1991). HOLOTYPE. United States: California, Mendocino Co., Fort Bragg. On twigs of Pinus muricata. I.M. Brodo 25301, 13 July 1986 (CANL 94666). 327 Ochrolechia montana Brodo, Canad. J. Bot. 69: 753 (1991). HOLOTYPE. United States: Montana, Lake Co., near Soup Creek, Swan Range. On bark. B. McCune 8854, 26 August 1877 (CANL 77559). Ochrolechia pseudopallescens Brodo, Canad. J. Bot. 69: 755 (1991). HOLOTYPE. Canada: Quebec, Gatineau Co., near Eardley in Picea mariana bog between Ramsay Lake and Lac Hawley. On Picea mariana twigs. I.M. Brodo 16280, 2 October 1969 (CANL 34613). Ochrolechia subisidiata Brodo, Canad. J. Bot. 69: 759 (1991). HOLOTYPE. United States: Arizona, Gila Co., McFadden Peak, 15 mi. S of Young. On dead Juniperus branch. T.H. Nash 11166 (CANL 58113). Ochrolechia trochophora var. pruinirosella Brodo, Canad. J. Bot. 69: 763 (1991). PARATYPES. Canada: New Brunswick. Gowan 3129 (CANL 81779); Ontario. I.M. Brodo 14680 & 16130 (CANL 27752 & 60142); Quebec, I.M. Brodo 9630, 16895 & 25290 (CANL 21653, 72040 & 93690); Prince Edward Island. J. Fabiszewski 7C & 9A (CANL 9329984933 00)«: Ochrolechia tuckermanii Vers., Beih. Nova Hedwigia 1: 97 (1962). ISOTYPE. United States: Massachusetts, Manchester. E. Tuckerman, 1839. (CANL 23277). Reliq. Tuck. No. 114 ("No. 14" in protologue). = O. yasudae Vainio Omphalodina bullata Follm. & Crespo, Philippia 3: 24 (1976). ISOTYPE. Spain: Prov. Toledo, Koloniebildend an halbschattigen, luftfeuchten, teils lberhangenden, verhdltnismassig glatten Silikatfeisen. G. Follmann & A. Crespo, 1975. (CANL 58316). Follmann: Lich. Exs. Sel. No. 174. = Rhizoplaca bullata (Follm. & Crespo) Leuck. & Poelt. Opegrapha albidoatra Fink, Mycologia 19: 210 (1927). PARATYPE. Puerto Rico: Naranjito. In woods on bark. B. Fink 298, 27 November 1915 (CANL 1903). Opegrapha astraea Tuck., Lich. Calif. p. 33 (1866). SYNTYPES. United States: South Carolina, Santee Canal. On Acer rubrum. H.W. Ravenel 466, 472 (CANL 68781, 68782). Reliq. Tuck. Nos. 119a, 119b. Opegrapha pertusariicola Coppins & P. James, Lichenologist 11: 164'-(1979). ISOTYPE. Great Britain (England): Caledonia, Mid-Perthshire, Aberfeldy, Glen Lyon, prope Woodend. In thallo Pertusariae leucoplacae vigens, ad corticem Coryli. P. James & B.J. Coppins 3834, 11 September 1976 (CANL 73861). Vézda: Lich. Sel. Exs. No. 1699. 328 Opegrapha subabnormis Fink, Mycologia 19: 210 (1927). ISOTYPE. Puerto Rico: Mayaguez. In open field on bark. B. Fink 1245, 33 December 1915 (CANL 1946). Oropogon colibor Essl., Syst. Bot. Monogr. 28: 60 (1989). ISOTYPE. Costa Rica: Cartago Prov., about 20.4 km SE of El Palme. On tree bark. F. Almeda 2143B, 27 December 1973 (CANL 98282). Oropogon diffractaicus Essl., Syst. Bot. Monogr. 28: 61 (1989). ISOTYPE. West Indies: Dominican Republic, Sierra de la Pelonia, Maciso Central, Cordillera Central. On small shrub in open. C.M. Wetmore & H.A. Imshaug 3652, 9 August 1958 (CANL 16620). Oropogon mexicanus Essl., Syst. Bot. Monogr. 28: 92 (1989). ISOTYPE. Mexico: State of Oaxaca, Trail leading to Cerro San Felipe. On conifers. M.E. Hale 20706, 1 April 1960 (CANL 31064). Pannaria leucostictoides Ohlsson, Bryologist 76: 379 (1973). ISOTYPE. Canada: British Columbia, Queen Charlotte Islands, Moresby Island, Skidegate Lake. On branches of Malus. I.M. Brodo 11085, 9 July 1967 (CANL 29174). Parmelia alabamensis Hale & McCull., Bryologist 71: 44 (1968). ISOTYPE. United States: Alabama, St. Clair Co., 6 miles E of Leeds. On sandstone outcrops. M.E. Hale 24072, June 1966 (CANL 31055). = Paraparmelia alabamensis (Hale & McCull.) Elix & Johnston Parmelia albertana Ahti, Bryologist 72: 236 (1969). ISOTYPE. Canada: Alberta, near SW tip of Big Lake, ca. 10 mi. NW of Edmonton City. On Populus balsamifera. T. ‘Ahti LItAas 7 July 1967 (CANL 28560). = Melanelia albertana (Ahti) Essl. Parmelia ambigua var. halei Tuck., Synops. N. Amer. Lich. 1: 66 (1882). ISOSYNTYPE. United States: South Carolina. On Pinus mitis. H.W. Ravenel (CANL 68744). Reliq. Tuck. No. 65. = Parmeliopsis subambigua Gyelnik Parmelia chlorochroa Tuck., Proc. Amer. Acad. Arts 4: 383 (1860). ISOSYNTYPE. United States: North Dakota, Missouri River. F.V. Hayden, 1859-60 (CANL 68784). Reliq. Tuck. No. 121. = Xanthoparmelia chlorochroa (Tuck.) Hale Parmelia crinita f. varians G. K. Merrill, Bryologist 11: 95 (1908). HOLOTYPE. Canada: Ontario, Carleton Place. On elm trees ina Swamp. J. Macoun 3828, 12 May 1900 (CANL 13348). = Parmotrema subtinctorium (Zahlbr.) Hale 329 Parmelia cryptoxanthoides Kurok., Bull. Natl. Sci. Mus. Tokyo. 17: 297 (1974). ISOTYPE. Brazil: Parana, Jardim Paraizo, 9 km west of Curitiba. On trees. S. Kurokawa 8245, 9 July 1971 (CANL 57029). Kurokawa: Lich. Rar. Crit. Exs. No. 232. Parmelia elabens Kurok., Bull. Natl. Sci. Mus. Tokyo. 17: 298 (1974). ISOTYPE. Brazil: Parana, Jardim Paraizo, 9 km west of Curitiba. On trees. S. Kurokawa 8253, 9 July 1971 (CANL 57030). Kurokawa: Lich. Rar. Crit. Exs. No. 233. Parmelia dubia var. scrobiculata B. de Lesd., Bull. Soc. Bot. BVance 499: 145941952) :. ISOTYPE. Italy: Liguria sabatia, Spotorno, tinicola loco "Collina". Sbarbaro, 19 December 1951 (CANL 13700). = Punctelia perreticulata (Rdsdnen) Wilhelm & Ladd Parmelia frondifera G. K. Merrill, Bryologist 11: 91 (1908). HOLOTYPE. Canada: Ontario, Essex Co., Point Pelee. On cedar bark. J. Macoun 3627, 27 May 1901 (CANL 13193). = Punctelia bolliana (Miill. Arg.) Krog Parmelia hygrophila Goward & Ahti, Ann. Bot. Fenn. 20: 9 (1983). ISOTYPE. Canada: British Columbia, Kokanee Creek Provincial Park, 17 km east of Nelson. On trunk of Betula papyrifera. T. Goward 81-1601, 25 July 1981 (CANL 86317). Parmelia gloriosa Kurok., Stud. Cryptog. Papua New Guinea p. 135 (1979). ISOTYPE. Papua New Guinea: Central District, Mt. Albert Adward, around Abios Hut. On bark of Pandanus sp. S. Kurokawa 9113, 31 October 1975 (CANL 68483). Kurokawa: Lich. Rar. Crit Exs.)No,.°'368. Parmelia omphalodes subsp. glacialis Skult, Ann. Bot. Fenn. Zee 2cuLe (1985)\, HOLOTYPE. Canada: Northwest Territories, Dist. of Franklin, Prince Patrick Island, Mould Bay. On damp polygon soil. S.D. MacDonald, 24 July 1952 (CANL 13656). PARATYPES. Canada: Northwest Territories. L. C. Bliss 2. (CANL 70510); M. Kuc 6, AA-14, AK-11, AK-12, AK-13, AM-24, AN-15, LC-13 & LD-11 (CANL 29811, 29854, 29871, 29818, 29787, 29850, 29840, 29774 & 29779); S.D. MacDonald (CANL 13647); D.B.O. Savile 4102 (CANL 46004); D.W. Smith (CANL 63481). = Parmelia skultii Hale Parmelia pacifica Kurok., Mem. Natl. Sci. Mus. Tokyo 11: 28 (1978). ISOTYPE. Japan: Bonin Islands, Hahajima Island, En route between Oki-mura and Cape Minami-zaki. On bark of Pandanus boninensis. S. Kurokawa 77014, 3 August 1977 (CANL 68485). Kurokawa: Lich. Rar. Crit. Exs. No. 370. Parmelia physodes var. enteromorpha f. rugosa G. K. Merrill, 330 Bryologist 11: 86 (1908). HOLOTYPE. Canada: British Columbia, near MacLeodes Lake. J. Macoun 696, 27 June 1875 (CANL 14331). = Hypogymnia rugosa (G. K. Merrill) Pike Parmelia physodes f. subisidioides G. K. Merrill, Bryologist Dis 86m (1908). SYNTYPE. Canada: Alberta. On old logs at Laggan. J. Macoun 3635, 26 June 1904 (CANL 14298). = Hypogymnia austerodes (Nyl.) Rdasdnen Parmelia ralla Brusse, Mycotaxon 27: 241 (1986). ISOTYPE. South Africa: Swartberg range, Seven Weeks poort, 8.6 km NW of Amalienstein. On Table Mountain sandstone boulders. F. Brusse 4935, 10 February 1986 (CANL 97358). Parmelia reptans Kurok., Austral. J. Bot. 21: 137 (1973). ISOTYPE. Australia: Victoria, 6 miles west of Red Cliffs along the Werrimull Road. On soil. S. Kurokawa 6621, 8 December 1965 (CANL 68487). Kurokawa: Lich Rar. Crit. Exs. NOSUE3 7 25 = Xanthoparmelia reptans (Kurok.) Elix & Johnston Parmelia saximontana R. Anderson & W. Weber, Bryologist 65: 23081962) e. ISOTYPE. United States: Colorado, Boulder Co., Dakota Group just NW of Boulder. On sandstone. R.A. Anderson & W.A. Weber L-30072, 6 April 1962 (CANL 20831). Weber: Lich. Exs. Colo. Now (41. = Melanelia tominii (Oxner) Essl. Parmelia semansiana Culb. & C. Culb., Mycologia 72: 128 (1980). ISOTYPE. United States: Arkansas, Polk Co., 5 km ad septentriones versus a Mena, in loco dicto Blue Haze Vista prope viam dictam Talimena Skyline Drive. Ad saxa in silva. W.L. Culberson & C.F. Culberson 16074, 20 August 1972 (CANL 76386). Vézda: Lich Sel. Exs. No. 1737. = Punctelia semansiana (Culb. & C. Culb.) Krog Parmelia spinibarbis Kurok., Bull. Natl. Sci. Mus. Tokyo. 17: 299 8( 197A). ISOTYPE. Brazil: Rio de Janeiro, Petrépolis. On trees. S. Kurokawa 8348, 17 July 1971 (CANL 57035). Kurokawa: Lich. Rar. Crity Exs sj) Now 2387 Parmelia squarrosa Hale, Phytologia 22: 29 (1971). ISOTYPE. United States: Virginia, Madison Co., Shenandoah National Park. On Quercus alba. M.E. Hale 36949, 30 April 1979 (CANL 38279). Hale: Lich. Amer. Exs. No. 142. Parmelia subdistorta Kuroki 2) Hattorr Bot... bab: 32-7 ie (1969). ISOTYPE. Australia: Victoria, 6 miles west of Red Cliffs along the Werrimull Road. On soil. S. Kurokawa 6617, 8 December 1965 (CANL 45332). Kurokawa: Lich. Rar. Crit. Exs. 331 No.4, 184. = Xanthoparmelia subdistorta (Kurok.) Hale Parmelia sulcifera Kurok., Bull. Natl. Sci. Mus. Tokyo. Ser. Bees: 371982) ISOTYPE. Australia: New South Wales, 9 miles E of Cooma on Numeralla Road. On rocks. S. Kurokawa 6450, 2 December 1965 (CANL 87340). Kurokawa: Lich. Rar. Crit. Exs. No. 583. = Xanthoparmelia sulcifera (Kurok.) Hale Parmelia trabeculata Ahti, Acta Bot. Fenn. 70: 54 (1966). PARATYPE. Canada: Ontario, Kenora District, Winisk. On trunks of Populus balsamifera. T. Ahti 4669, 7 August 1958 (CANL 14265). = Melanelia trabeculata (Ahti) Essl. Parmelia virginica Hale, Contr. U.S. Natl. Herb. 36(4): 186 (1964). ISOTYPE. United States: Virginia, Page Co., Hawksbill, Shenandoah National Park. On fir trees. M.E. Hale 21592, 26. June 1962 (CANL 31054). = Hypotrachyna virginica (Hale) Hale Parmeliella concinna Lamb, Farlowia 4: 431 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near Futalaufquén. On a carbonised trunk in open forest. I.M. Lamb 5879, 1 November 1950 (CANL 2629). Parmeliella granulata Lamb, Farlowia 4: 429 (1955). HOLOTYPE. Argentina: Prov. Neuquén, Parque Nacional Lanén, Lago Quillen. O. Kuhnemann 791 p.p., 10 November 1943 (CANL 2841). Parmotrema demethylmicrophyllinicum Elix, Lich. Austral. Exs. PASC... 9,NO.a.14 . (1986 ).. ISOTYPE. Australia: Queensland, Haning Pinnacle (Dempseys Peak), 7 km E of Mt. Garnet. On granite rocks. J.A. Elix & H. Streimann 16844, 29 June 1984 (CANL 95337). Elix: Lich. Austral. Exs. No. 114. Parmotrema queenslandense Elix, Lich. Austral. Exs. Fasc. 5, No. 116 (1986). ISOTYPE. Australia: Queensland, near the summit of Mt. Leswell, 32 km S of Cooktown. On granite rocks. J.A. Elix & He Streimann 17365, 5 July, 1984..(CANL 95339) .. Elax: Lich, Austral. Exs. No. 116. Parmotrema indicum Hale, Mycotaxon 5: 436 (1977). ISOTYPE. India: Tamil Nadu, Silver Cascade, Kodaikanal, Palni Hills. Ad saxa in vicinitate fluminis. M.E. Hale 43874, January 1975 (CANL 69492). Vézda: Lich. Sel. Exs. No. 1573. Peltigera kristinssonii Vitik., Ann. Bot. Fenn. 22: 291 (1985). PARATYPE. United States: Colorado, Boulder Co., Boulder Canyon. W.A. Weber, 23 September 1975 (CANL 58199). Weber: 332 Lich. -Exs./Colo. Now 484. Peltigera malacea f. hasimotoi Asah. & Inum., Acta Phytotax. Geobot. 12: 8 (1943). ISOTYPE. Japan: Prov. Etchu, Mt. Kaminotake. Over mosses. Hasimoto & Asahina, 24 July 1936 (CANL 20052). Kurokawa: Lich. Rar. Crit. Exs: No. 43%. Peltigera pacifica Vitik., Ann. Bot. Fenn. 22: 294 (1985). ISOTYPE. Canada: British Columbia, 10 km N of Kitsumkalum Lake on E branch road. Terricolous. T. Goward 81-2025, 24 August 1981 (CANL 104553). Peltula cylindrica Wetm., Ann. Missouri Bot. Gard. 57: 182 (1970). ISOTYPE. United States: Georgia, Dekalb Co., Mt. Arabia, 20 km E von Atlanta. C.M. Wetmore 19264, 1969 (CANL 102538). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 33. Pertusaria gymnospora Kantvilas, Lichenologist 22: 296 (1990). PARATYPE. Australia: Tasmania, forest around Lake Judd, SW Tasmania. On dead wood, soil, rotten leaves. G. Bratt 73/900, 28 July 1973 (CANL 58194). Weber: Lich. Exs. Colo. No. 479. Pertusaria neoscotica Lamb, Rep. (Annual) Natl. Mus. Canada, 1952-537, Bulls 1323"284')5(1954)% HOLOTYPE. Canada: Nova Scotia, Cape Breton Island, Baddeck. On trunk of trees. J. Macoun 1953, p.p. 9 July 1898 (CANL 11479). Pertusaria paramerae A. Crespo & Vézda, Anales Jord. Bot. Madrid 41: 252, 1984 (1985). ISOTYPE. Spain: Prov. Soria, prope vicum judes, Paramera. Ad corticem Juniperi thuriferae. A. Crespo & A. Vézda, 24 May 1983 (CANL 89185). Vézda: Lich. Sel. Exs, No. 1979. Pertusaria santamonicae Dibben, Publ. Biol. Geol. Milwaukee Public Mus. 5: 67 (1980). PARATYPE. United States: California, San Mateo Co., San Francisco, below Spillway of San Andres Lake. W. Jordan 769, 13 September 1967 (CANL 27071). Pertusaria suboculata Brodo & Dibben, Publ. Biol. Geol. Milwaukee Public Mus. 5: 72 (1980). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Moresby Island, Tasu. On Alnus sinusta. I.M. Brodo 12825, 28 July 1967 (CANL 30938). PARATYPES. Canada: moresby Island. I.M. Brodo 10896, 10980 & 14210 (CANL 29029, 27485 & 36493); United States: Oregon. C.E. Fellows 32934 (CANL 38171). Pertusaria subpupillaris Vézda, Vézda: Lich. Sel. Exs. Fasc. 20: 5 (1966). ISOTYPE. United States: California, Mendocino Co., Caspar. On branches of Pinus muricata. H. Sharsmith & I. Tavares 725, 25 333 July 1961 (CANL 36511). Vézda: Lich. Sel. Exs. No. 491. = P. glaucomela (Tuck.) Nyl. Pertusaria subvelata G. kK. Merrill, Bryologist 11: 111 (1908). ISOTYPE. United States: Alaska, Skagway. On birch bark. J. Macoun 87, 4 September 1902 (CANL 11603). = P. panyrga (Ach.) Massal. Pertusaria sulcata Dibben, Publ. Biol. Mulwaukee Public Mus. 62073) (1980)- ISOTYPES. United States: Maine, North Lubec. On rail fences. C.E. Cummings & E.A. Teller, 3 July 1893 (CANL 19548 & 68955). Cummings: Dec. N. Amer. Lich. No. 136 & Lich. Boreali-Amer. No. 55. Pertusaria victoriana Lamb, Farlowia 4: 463 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lake Nahuel Huapi, Isla Victoria. On a large rock. I.M. Lamb 5817, 26 January 1950 (CANL 10590). Phaeocalicium asciiforme Tibell, Symb. Bot. Upsal. 27(1): 200 (1987). ISOTYPE. New Zealand: South Island, Westland, Arthur's Pass National Park, 3 km NNE of Otira. On branches of Coprosma sp. L. Tibell 9944b (CANL 99801). Tibell: Caliciales Exs. No. 168. Phaeopeccania australiensis Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 10 (1990). ISOTYPE. Australia: Kiistenebene zwischen Denham und Nanga; in Vertiefungen auf horizontalen Felsplatten aus Muschelkalk. A. Henssen 31366a, 31 August 1987 (CANL 102556). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 49. Phloeopeccania hispanica Henssen & Willems, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 10 (1990). ISOTYPE. Spain: Andalusien, Sierra de Caho de Gata, Las Negras, Nordhang am Meeresufer. A. Henssen & G. Willems 33008a, 2-4 January 1989 (CANL 102555). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 48. Phaeophyscia spinellosa Kashiw., Bull. Natl. Sci. Mus. Tokyo. Ser. B, 10: 46 (1984). ISOTYPE. Japan: Honshu, Prov. Suruga, Fujieda-city. On rocks. K. Sugiyama, 10 August 1974 (CANL 87342). Kurokawa: Lich. Rar. “Cri. xs. No. 585. Phlyctis pulveracea Lamb, Lilloa 26: 425 (1953), HOLOTYPE. Uruguay: Dept. Florida, La Palma. On bark. W.G. Herter 99430, 1938 (CANL 12933). Physcia duplicorticata W. Weber & Thomson, Mycotaxon 3: 102 (1975). ISOTYPE. United States: California, Marin Co., 1.6 km E of junction Nicasio-Point Reyes road E of Inverness. On trunk 334 and branches of Umbellularia californica. W.A. Weber & G. Kunkel, 12 March 1975 (CANL 58191). Weber: Lich. Exs. Colo. No. 476. Physcia hirsuta var. echinella Poelt, Mitt. Bot. Staatssamml. Muinchen 4: 298 (1962). ISOTYPE. Italy: Appennino Ligure, Liguria, A un muro un poco ombroso, sopra Fabiano presso La Spezia. C. & J. Poelt, April 1962 (CANL 52894). Poelt: Lich. Alpium No 179. = Phaeophyscia cernohorskyi (Nadv.) Essl. Pilophorus cereolus var. hallii Tuck., Syn. North Amer. Lich. ies) (ake i ISOTYPE. United States: Oregon. E. Hall, 1871 (CANL 68789). Reliqy Tuck .4No. (1275 = P. clavatus Th. Fr. Placodium coralloides Tuck., Proc. Amer. Acad. Arts 6: 287 (1866). ISOTYPE? United States: California. On coastal rocks near San Francisco. H.N. Bolander, 1865 (CANL 68790). Relig. Tuck. No. L2or = Caloplaca coralloides (Tuck.) Hult. Placopsis alphoplacoides Lamb, Lilloa 13: 234 (1947). ISOTYPE. New Zealand: South Island, Western Bot. Distr., Otira Gorge, rata belt. On earth. G.E. & G.Du Rietz 2667, 15 April 1927 (CANL 12558). = P. trachyderma (Krempelh.) P. James Placopsis alphoplacoides var. clavifera Lamb, Lilloa 13: 236 (1947). ISOTYPE. New Zealand: South Island, Western Bot. Distr., Otira Gorge. On earth. G.E. & G.Du Rietz 2668, 15 April 1927 (CANL 12630). = P. trachyderma (Kremp.) P. James var. clavifera (Lamb) P. James in Mark Placopsis contortuplicata f. fuegiensis Lamb, Lilloa 13: 276 (1947). ISOTYPE. Argentina: Tierra del Fuego, Sierra Alvear, ca. 20 km ENE of Ushuaia. On rocks in the alpine region. R. Santesson 901, 7 November 1940 (CANL 12560). Placopsis dusenii Lamb, Lilloa 13: 215 (1947). ISOTYPE. Chile: Magallanes, Isla Desolaci6én, Puerta Angosto. P. Dusén 199 p.p., 1896 (CANL 12568). Placopsis gelidoides Du Rietz ex Lamb, Lilloa 13: 210 (1947). ISOTYPE. New Zealand: South Island, Cass, Mt. Misery, in upper subalpine belt. G.E. & G.Du Rietz 1468:15, 9 January 1927 (CANL 12608). Placopsis lateritioides Lamb, Lilloa 13: 222 (1947). ISOTYPE. New Zealand: South Island, Cass, Mt. Misery, in upper subalpine belt. On rocks. G.E. & G.Du Rietz 1468:16, 9 335 January 1927 (CANL 12609). Placopsis perrugosa f. activa Lamb, Lilloa 13: 272 (1947). ISOTYPES. Chile: Magallanes, Isla Riesco, Mina Elena. On a stone block. R. Santesson 2023a, 29 April 1940 (CANL 12614 & 12621). Placopsis salazina Lamb, Lilloa 13: 259 (1947). ISOTYPE. New Zealand: Southland, Mt. Barber, above Deep Cove. On boulder. G.E. & G.Du Rietz 2063:2, 3 March 1927 (CANL 12635). Plagiocarpa septemseptata R. Harris, Michigan Bot. 12: 37 CL972)e ISOTYPE. United States: North Carolina, Transylvania Co., Pisgah National Forest. On Fraxinus. R.C. Harris 3373, 29 May 1967 (CANL 37270). Polyblastia cucurbitula Thomson, Bryologist 91: 89 (1988). PARATYPES. Canada: British Columbia. Brodo 21737 & 22417 (CANL 87604 & 86571). Polyblastia sbarbaronis Servit, Ann. Mus. Civico Storia Nat. Giacomo Doria 66: 241 (1953). ISOTYPE. Italy: Toscana, Vallombrosa, Saltino. Sbarbaro, July 1937 (CANL 1003). Polyblastiopsis quercicola Brodo, Bull. New York State Mus. mSCioeSury. 410: 147 (1968)*. ISOTYPE. United States: New York, Suffolk Co., Shorehen, Saint Joseph's Villa. N. Country Road. On Quercus alba. I.M. BEOdOVZ65 19d OuULy 1961 (CANE! 2277). = Julella fallaciosa (Stizenb. ex Arnold) R. Harris Polycauliona coralligera Hue, Expédit. Antarct. France 1903- 05. p.10 (1908). ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Booth (Wandel) Island. Expédit. Antarct. France 1903-05, No. 277, 299. 1904 (CANL 17955). = Xanthoria candelaria (L.) Th. Fr. Porina mangiferae Vainio, Univ. Calif. Publ. Bot. 12(1): 14 (1924). ISOTYPE. Tahiti: Society Islands, Fautaua Valley. On leaves of Mangifera. W.A. Setchell and H.E. Parks 5091, p.p., 3 June 1922 (CANL 103926). = P. corruscans (Rehm) R. Sant. Porocyphus kalbarrensis Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 9 (1990). ISOTYPE. Australia: Kalbarri National Park, Red Bluff. Sandstein. A. Henssen 31358a, 29 August 1987 (CANL 102552). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 46. Porpidia carlottiana Gowan, Bryologist 92: 39 (1989). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, 336 Graham Island, Port Lewis. On rock. I.M. Brodo 10489, 26 June 1967 (CANL 27433). Porpidia thomsonii Gowan, Bryologist 92: 54 (1989). ISOTYPE. Canada: Northwest Territories, District of Keewatin, North end of Lake Ennadai. On boulders. J.W. Thomson 20416, 22 July 1960 (CANL 78044). Protoparmelia loricata Poelt & Vézda, Vézda: Lich. Sel. Exs. Fasc. 92: .5 (1989); ISOTYPE. Austria: Tirolia orientalis, alpes dicti Hohe Tauern, in valle Umbaltai. J. Poelt & R. Tiirk, 29 August 1988 (CANL 99233). Vézda: Lich. Sel. Exs. No. 2292. Pseudocyphellaria anomala Brodo & Ahti, Mycotaxon 28: 95 (1987). HOLOTYPE. United States: Washington. On trees. A.S. Foster, April 1908 (CANL 20272). Pseudocyphellaria exanthematica Lamb, Farlowia 4: 435 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias. On mossy trunk of Coihue. I.M. Lamb 5986, 11 November 1950 (CANL 3313). Pseudocyphellaria hirsuta f. leucosticta Lamb, Farlowia 4: 436 (1955). ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Futalaufquen. O. Ktihnemann 5449, 1941 (CANL 3334). Pseudocyphellaria rainierensis Imsh., Mycologia 42: 749 (1950). ISOTYPE. United States: Washington, Mt. Rainier National Park, Ohanapecosh River above Panther Creek. H.A. Imshaug 598, 23 July 1948 (CANL 3369). Pseudoparmelia crystallicola Kalb & Hale, Kalb: Lich. Neotr. Fasc., 7: 14 (1983). ISOTYPE. Brazil: Minas Gerais, Serra do Espinhacgo. Bei Diamantina, in einer offenen Felssteppenlandschaft. K. Kalb & G. Plébst, 17 July 1978 (CANL 85932). Kalb: Lich. Neotr. No. 294. Psilolechia leprosa Coppins & Purvis, Lichenologist 19: 35 (1987). PARATYPE. Sweden: Harjedalen Prov., Tanndas par., Mt Gruvvalen, 0.1 km NE of Lake Glimsj6n. On the walls and roof of a cave in the old copper mine. R. Santesson 31128, 10 August 1984 (CANL 95091). Moberg: Lich. Sel. Exs. Ups. No. 16. Psora cerebriformis W. Weber, Mycotaxon 13: 104 (1981). ISOTYPE. United States: Colorado, Montrose Co., 4 mi. E of Bedrock, on gypsum knolls, floor of Paradox Valley. S. Shushan, R. Anderson & W.A. Weber 25501, 30 May 1960 (CANL 20814). Weber: Lich, Exs. Colo. No. 24. 337 Psora montana Timdal, Bryologist 89: 266 (1986). ISOTYPE. United States: Colorado, Grand Co., Tonahutu Creek Trail on NE end of Big Meadows. R.A. Anderson 2386, 6 July 1962 (CANL 124) Psora pacifica Timdal, Bryologist 89: 268 (1986). ISOTYPE. United States: California, Santa Barbara Co., Channel Islands, Santa Cruz Island, upper end of Islay Canyon. On bare earth. W.A. Weber & C. Bratt, 8 January 1986 (CANL 97963). Weber: Lich. Exs. Colo. No. 664. Psora texana W. Weber, Mycotaxon 6: 178 (1977). ISOTYPE. United States: Texas, Kerr Co., Guadalupe River Canyon, Hwy. 39, 22.4 km SW of Ingram, on road to Leakay. On vertical faces of limestone cliffs. W.A. Weber, 29 April 1974 (CANL 52198). = Xanthopsorella texana (W. Weber) Kalb & Hafellner Psoroma internectens Lamb, Farlowia 4: 427 (1955). ISOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Menendez. On soil among mosses. O. Kiihnemann 4747, 1940 (CANL 3049). Psorotichia rimosa Henssen, Henssen: Lich. Cyanoph. Fungi Saxe SEXS -erasc. 2221 (19902 ISOTYPE. France: Languedoc, Ariége, Montagnes du Plantaurel, anstehender Kalk an der Strasse zwischen Mas d'Azile und Sabarat. A. Henssen 29172a, 5 June 1983 (CANL 102557). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 50. Pterygiopsis australiensis Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 9 (1990). ISOTYPE. Australia: Kalbarri National Park, Red Bluff. A. Henssen 31357, 29 August 1987 (CANL 102550). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 44. Pterygiopsis canariensis Henssen, Henssen: Lich. Cyanoph. Fungi Sax. Exs. Fasc. 2: 8 (1990). PARATYPE. Canary Islands: Fuerteventura, Sukkulentenhalbwiiste, auf basisch-kristallinem Gestein. A. Henssen 32050, 13-14 February 1988 (CANL 102548). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 42. Pterygiopsis convoluta Henssen, Henssen: Lich. Cyanoph. Fungi Sax. -Exs. Fasc. 2: 8 (1990). ISOTYPE. Australia: Northampton Gorge, Konglomerat aus quarzitischem Sandstein im oberen Teil der Schlucht auf Sickerwasserflachen. A. Henssen 31373a, 1 September 1987 (CANL 102549). Henssen: Lich. Cyanoph. Fungi Sax. Exs. No. 43. Pterygium petersii Nyl., Syn. Meth. Lich. 1: 93 (1858). ISOTYPE. United States: Alabama, Moulton. 7T.M. Peters, 1853 (CANL 68740). Reliq. Tuck. No. 61. = Placynthium petersii (Tuck.) Burnham Pyrenastrum gemmeum Tuck., Amer. J. Sci Arts Ser. 2, 25: 429 338 (1858). SYNTYPE. United States: South Carolina, Santee Canal. H.W. Ravenel, 1853 (CANL 68791). Reliq. Tuck. No. 129. = Parmentaria astroidea Fée Pyrenopsis sojakii Vézda, Folia Geobot. Phytotax., Praha 14: 205 (1979). ISOTYPE. Iran: "Persia australis", Bandar Lengeh. Ad lapides Calcareos locis apricis. J. Sojak, 25 April 1977 (CANL TIE17)e Pyrenula cinerella var. quadriloculata Fink, Minnesota Bot. Stud w' 2257217 6atG2899)% ISOTYPE. United States: Minnesota, Grand Portage Island. B. Fink 85, 24 June 1897 (CANL 1384). = ‘unknown sp. (fide R- C. Harris, "Michigan “BOC. 7 1227949 (19°73) Pyrenula leucoplaca var. pluriloculata Fink, Minnesota Bot. Stud. 2: 709 (1902). SYNTYPE. United States: Minnesota, Thief River Falls. On trees. B. Fink 829, 19 July 1900 (CANL 1335). = Eopyrenula intermedia Coppins in Aptroot (1991). Pyrenula macounii R. Harris, Michigan Bot. 12: 45 (1973). HOLOTYPE. Canada: Ontario, Belleville. On trees. J. Macoun, April 1874 ‘(CANL 1350). Pyrenula neglecta R. Harris, Michigan Bot. 12: 45 (1973). ISOTYPE. United States: New Hampshire, Plymouth. On beech trees. C.E. Cummings, 26 August 1893 (CANL 19561). Cummings: Dec. N. Amer. Lich. No. 150. = Pyrenula pseudobufonia (Rehm) R. Harris Pyrenula neglecta subsp. occidentalis R. Harris, Michigan Bots §12:°'51°(1973)5 ISOTYPE. United States: Washington, Gate. On vine maple. A.S. Foster, 30 March 1912 (CANL 20502). Merrill: Lich. Exs. No. 5 Pale = P. occidentalis (R. Harris) R. Harris Pyrenula nitidella var. maculata R. Harris, Weber: Lich. Exs. Colo. Fasc. +120 eNow'4437(1975).. ISOTYPE. United States: Texas, McLennan Co., Canyon of the Middle Bosque River SW of Valley Mills and W of Waco. On saplings of a smooth-barked tree. W.A. Weber, 27 May 1974 (CANL 52193). Weber: Lich. Exs. Colo. No. 443. = P. maculata (R. Harris) R. Harris Pyrenula shirabeicola Kurok. & S. Nakan., Mem. Natl. Sci. Mus. TOKYO?4 : +67 VLSI ISOTYPE. Japan: Honshu, Prov. Hida, Mt. Ontake. On trunk of Abies veitchii. S. Kurokawa 64172, 13 August 1964 (CANL 45336). Kurokawa: Lich. Rar. Crit. Exs. No. 188. Pyrenula thelomorpha Tuck., Gen. Lich. p. 275 (1872). 339 ISOTYPE. United States: South Carolina, Santee Canal. On Ilex decidua. H.W. Ravenel 468 (CANL 68797). Reliq. Tuck. No. 135. = Anthracothecium thelomorphum (Tuck.) Zahlbr. Pyxine daedalea Krog & R. Sant., Publ. Herb. Univ. Uppsala, Thunbergia 2: 7 (1986). ISOTYPE. Costa Rica: Cartago Prov., 13 km SE of Cartago, 2.5 km SE of Orosi near the bridge over Rio Grande de Orosi. On a large boulder. H. Krog & R. Santesson 29074, 10 January 1979 (CANL 95093). Moberg: Lich. Sel. Exs. Ups. No. 17. Ramalina americana Hale, Bryologist 81: 599 (1978). ISOTYPES. United States: New Hampshire, Plymouth. On yellow birch. C.E. Cummings, 28 May 1891 (CANL 19451, 68913). Cummings: Dec. N. Amer. Lich. No. 43 & Lich. Boreali-Amer. No. 5. Ramalina atlantica Culb., Brittonia 19: 350 (1967). ISOTYPE. Scotland: Argyllshire, Ardnamurchan Peninsula, Portaick. On immense boulder. W.L. Culberson 12254, July 1964 (CANL 22578). Ramalina bicolor Mull. Arg., Flora 60: 476 (1877). ISOTYPE. Brazil: Bahia, Blanchet (CANL 42339). Ramalina cactacearum Follm., Nova Hedwigia 14: 256 (1967). ISOTYPE. Chile: Prov. Antofagasta, zerstreut im Dornenwerk von Sdaulenkakteen im Tornabenietum intricatae. I.A. Follmann- Schrag & G. Follmann, September 1965 (CANL 33001). Follmann: Pach mExs: Sell. No. #51. Ramalina euxini Vézda, Folia Geobot. Phytotax., Praha 14: 205 (1979). ISOTYPE. Bulgaria: Distr. Burgas, in litore Euxini Ponti, 5 km ad meridiem versus ab oppido Sozopol. Ad scopulos andesiticos. A, Vézda, 20 September 1979 (1978 in protologue) (CANL 73831). Vézda: Lich. Sel. Exs. No. 1668. Ramalina sideriza Zahlbr., Ann. K. K. Naturhist. Hofmus. 25: 246' (1911). ISOSYNTYPE. United States: Hawaii, Island of Kauai. On trunk of trees. A.A. Heller, 28 June 1895 (CANL 20370). Merrill: PLC uxs. NO-1e120. Ramalina superfraxinea Follm. & Sanch.-Pinto, Philippia 4: 192 (1980). ISOTYPE. Canary Islands: Tenerife, Vogelnestartig im halbschattigen Unterkronenteil solitdrer Nadelbadume auf jungen Lavastrémen im Ramalinetum subgeniculatae. G. Follmann & L. Sanchez-Pinto, April 1978 (CANL 75275). Follmann: Lich. EXS* wel No: 314" Ramalina tenuis Fries & Tuck. in Tuck., Amer. J. Sci. Arts Serie 2)) 2520-423 4(1858)™ ISOLECTOTYPE. United States: Texas, Mexican Bountary. Thickets of the Blanco. C. Wright, 1849 (CANL 68759). Reliq. 340 Tuck. No. 84. LECTOPARATYPE. United States: South Carolina, Santee Canal, H.W. Ravenel, 1869 (CANL 68760). Reliq. Tuck. No. 85. Ramalina tigrina Follm., Willdenowia 4: 227 (1966). ISOTYPE. Chile: Prov. Antofagasta, gesellig auf Erosionsrippen. I.A. Follmann-Schrag & G. Follmann, September 1965 (CANL 32975). Follmann: Lich. Exs. Sel. No. 10. Ramonia intermedia Kalb, Kalb: Lich. Neotr. Fasc. 6: 14 (1983). ISOTYPE. Brazil: Minas Gerais, Serra da Mantiqueira, Oberhalb von Vila Monte Verde, etwa 30 km 6stlich von Camanducaia. In einem Gebirgsregenwald. K. Kalb & H. Sipman, 2 July 1979 (CANL 84791). Kalb: Lich. Neotr. No. 242. Ramonia kandlerii Kalb, Kalb: Lich. Neotr. Fasc. 6: 14 (1983). | ISOTYPE. Brazil: Mato Grosso do Sul. Zwischen Rio Verde do Mato Grosso und Coxim. In einem dichten Cerradado in einer Bachschlucht. K. Kalb, 28 June 1980 (CANL 84792). Kalb: Lich. Neotr. No. 243. Rhizocarpon compositum Lamb, Lilloa 14: 222 (1948). ISOTYPE. Argentina: Prov. Tucuman, Quebrada de Lules. On rock outcrop. I.M. Lamb 5132, 5 October 1947 (CANL 18207). = Buellia excellens H. Magn. Rhizocarpon alaxensis Thomson, Nova Hedwigia 14: 471 (1967). ISOTYPE. United States: Alaska, Franklin Bluffs on the Sagavanirktok River. On rocks. J.W. Thomson, S. Shushan & J. Koranda 10859, 30 July 1958 (CANL 44991). Rhizocarpon atlanticum Lamb, J. Bot. 78: 132 (1940). PARATYPE. England: Cornwall, Rocky Valley near Boscastle. I.M. Lamb 877, 1939 (CANL 6111). = R. constrictum Malme Rhizocarpon barilochense Rds&nen, Arch. Soc. Zool.-Bot. Fenn. "Vanamo" 6(2): 85 (1952). ISOTYPE. Argentina: Patagonia, Prov. Rio Negro, Cerro Catedral near Bariloche. On rocks. I.M. Lamb 5958, 10 February 1950 (CANL 27348). Rhizocarpon cumulatum Thomson, Nova Hedwigia 14: 461 (1967). ISOTYPE. United States: Alaska, Pitmegea River, 15 miles upstream from Cape Sabine. On shaded layers at top of bluff. J.W. Thomson 10571, 12 July 1958 (CANL 45003). Rhizocarpon hensseniae Brodo, Biblioth. Lichenol. 38: 32 (1990). HOLOTYPE. Canada: British Columbia, Queen Charlotte Islands, Graham Island, Seal Inlet in Rennell's Sound. On exposed stone. I.M. Brodo 10264, 23 June 1967 (CANL 98871). Rhizocarpon inimicum Poelt & Vézda, Herzogia 6: 471 (1984). 341 ISOTYPE. Spain: Catalonia, montes Montseny, Matagalls. Ad saxa schistosa, in thallo Lecanorae rupicolae. J. Poelt & A. Vézda, 28 May 1983 (CANL 89186). Vézda: Lich. Sel. Exs. No. 1980 (1970 in protologue). Rhizocarpon renneri Poelt, Planta 51: 306 (1958). ISOTYPE. Austria: Samnaungruppe, Tirol. Steilflache des grossen Blockes am K6lner Haus. J. Poelt, July 1958 (CANL 52817). Poelt: Lich. Alpium No. 100. Ricasolia beckettii Stirton, Trans. & Proc. New Zealand Inst. 3.075 °392°°(1897)), 2 ISOTYPE. New Zealand: Mount Bossu, Banks Peninsula. On Podocarpus totara. T.W.N. Beckett L. 3, 1893 (CANL 3387). = Pseudocyphellaria coriacea (T. D. Hook. & Taylor) D. Galloway & P. James ? Rimularia fuscosora Muhr & T@nsberg, Nord. J. Bot. 8: 649 (1989). ISOTYPE. Sweden: Vdsterbotten Prov., Skelleftea par., ca. 3 km NE of Mt Stora Blabergsliden, by the small stream Djupgravbacken. On Alnus incana. L.-E. Muhr 7540, 8 August 1984 (CANL 102833). Moberg: Lich. Sel. Exs. Ups. No 94. Rinodina afghanica M. Steiner & Poelt, Pl. Syst. Evol. 155: 139 (1987). ISOTYPE. Afghanistan: Prov. Samangan, 1 km W des Passes Kotal-Mirza Atbili. M. Steiner Ste 50/3, 6 June 1970 (CANL 104080). Steiner & Poelt: Lich. Afghanica No. 46. Rinodina ascociscana Tuck., Gen. Lich. p. 124 (1872). ISOSYNTYPE. United States: New Hampshire, White Mountains. E. Tuckerman (CANL 23267). Reliq. Tuck. No. 86. Rinodina calculiformis W. Weber, Bryologist 67: 473 (1964). ISOTYPE. Mexico: Baja California, Isla Guadalupe, Melpomene Cove. On volcanic scoria. W.A. Weber & C.J. McCoy, 21 April 1963 (CANL 20883). Weber: Lich. Exs. Colo. No. 94. Rinodina dakotensis H. Magn., Bot. Not. 1947: 40 (1947). ISOTYPES. United States: South Dakota, Brookings. On plum trees. 1T.A. Williams, 11 May 1895 (CANL 19580, 69031). Cummings: Dec. N. Amer. Lich. No. 169b & Lich. Boreali- Amer. No. 163b. Rinodina humilis H. Magn., Bot. Not. 1939: 132 (1939). ISOTYPE. Sweden: Vastergdétland, Bjdérketorp, Stenbacka. On moist stones. A.H. Magnusson, 26 March 1934 (CANL 20147). Magnusson: Lich. Sel. Scand. Exs. No. 324. Rinodina milliaria f. obscura H. Magn., Bot. Not. 1947: 19 (1947). ISOTYPE. United States: Maine, Rockland. On bark of young trees. G.K. Merrill 163, 21 April 1910 (CANL 20406). Merrill: Lich. .Exs< No. 163) = R. dakotensis H. Magn. 342 Rinodina oregana H. Magn., Bot. Not. 1947: 49 (1947). ISOTYPES. United States: Oregon, Forest Grove. On Acer macrophyllum. A.R. Sweetser, December 1899 (CANL 19739, 69098). Cummings: Dec. N. Amer. Lich. No. 330 & Lich. Boreali-Amer. No. 262. Rinodina pallida H. Magn., Bot. Not. 1939: 313 (1939). ISOTYPE. Sweden: Vdstergétland, Broddetorp, Fjdllakra. On a bridge of sandstone. A.H. Magnusson, 10 June 1938 (CANL 20148). Magnusson: Lich. Sel. Scand. Exs. No. 325. Rinodina sabulosa Tuck., Lich. Calif. p. 21 (1866). ISOTYPE. United States: California. H.N. Bolander, 1865 (CANL 68761) Reliq.w2uck. eNO. S7: = R. conradii Kdérber Roccella galapagoensis Follm., Nova Hedwigia 15: 337 (1969). ISOTYPE. Ecuador: Galapagos Islands, Isla Santa Cruz. On seaward vertical sides of bouders. W.A. Weber, 13 February 1964 (CANL 20901) Weber: Lich. Exs. Colo. No. 112. Roccellina luteola Follm., Nova Hedwigia 14: 239 (1967). ISOTYPE. Chile: Prov. Antofagasta, koloniebildend an Stirnfldchen und Uberhdngen von Diorithéckern im Roccelletum portentosae. G. Follmann, September 1965 (CANL 33004). Follmann: Lich. Exs. Sel. No. 54. Roccellina olivacea Follm., Philippia 4: 111 (1979). ISOTYPE. Chile: Prov. Coquimbo, gesellig an schattseitigen, durch Gischtstaub beeinflussten Stirnfldchen und Uberhdngen von Dioritfelsen im Roccelletum portentosae. Ktistenformation nérdlich der Limarimtindung. G. Follmann, September 1965 (CANL 71806). Follmann: Lich. Exs. Sel. No. 296. Sagenidiopsis merrotsii R. W. Rogers & Hafellner, Lichenologist 19: 402 (1987). ISOTYPE. Australia: New South Wales, Border Ranges National Park, Brindle Creek. Ad truncum Nothofagi moorei. J. Hafellner, P. Merrotsy & R. Rogers, 30 August 1986 (CANL 97328). Vézda: Lich. Sel. Exs. No. 2226. Sarrameana tasmanica Vézda & Kantv., Lichenologist 20: 179 (1988). ISOTYPE. Australia: Tasmania, Little Fisher River. Ad corticem arboris. G. Kantvilas & J. Jarman 63/87, 15 May 1987 (CANL 97302). Vézda: Lich. Sel. Exs. No. 2200. Schistophoron variabile Tibell, Lichenologist 14: 242 (1982). ISOTYPE. Costa Rica: Lim6én, 1 km NW of Pto. Lim6én. On trunk of Cocos nucifera. L. Tibell 8514, 9 January 1979 (CANL 87469). Tibell: Caliciales Exs. No. 96. Schizopelte californica Th. Fr., Flora 58: 143 (1875). ISOTYPE. United States: California. G. Fisen, 1874 (CANL 2122). = Combea californica (Th. Fr.) Follm. & Geyer 343 Setaria trichodes Michaux, Fl. Boreal.-Amer. 2: 331 (1803). ISOTYPE. Canada: Ad ramulos putridos (CANL 34981). = Bryoria trichodes (Michaux) Brodo & D. Hawksw. subsp. trichodes. Skyttea tephromelarum Kalb & Hafellner, Kalb: Lich. Neotr. Frascw 103 "15 (1988)': ISOTYPE. Kenya: Nanyuki District zwischen Naro Moru und Nanyuki. K. Kalb & A. Schrégl, 18 August 1985 (CANL 102737). Kalb: Lich. Neotr. No. 449. Sphaerophorus notatus Tibell, Publ. Herb. Univ. Uppsala 10: 9 (1982). ISOTYPE. New Zealand: Otago, Mount Aspiring National Park, 10 km NNE of Makaroa. On trunk of Nothofagus menziesii. L. Tibell 10604, 23 January 1981 (CANL 82051). Tibell: Caliciales Exs. No. 73. Sphaerophorus ramulifer Lamb, Farlowia 4: 426 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Rio Negro, Lago Frias. On trunk of Fitzroya near the lake. I.M. Lamb 5977, 11 February 1950 (CANL 1751). Squamaria configurata f. diffracta Oxner, Ukrayins'k. Bot. Zhurns.4353) (1928) ISOTYPE. Ukraine: Ditio Kirowogradensis. Ad safa granitica ad flum. Subakleji in viciniis opp. Kirowograd. A. Oxner, 22 May 1926 (CANL 11909). = Lecanora laatokkaensis (R&sdnen) Poelt Stephanocyclos henssenianus Hertel, Hertel: Lecideaceae Exs. Fasc. 5, No. 96 (1983). ISOTYPE. Subantarctic Islands: Marion Island, Nellie Humps, zwischen Junior's Kop und Meteorologischer Station. Niedrige, bizarre Lava-Felsen im moorigen Geldnde. H. Hertel, 13 April 1982 (CANL 85425). Hertel: Lecideaceae Exs. No. 96. Stereocaulon alpinum var. erectum Frey, Rabenh. Kryptog. Fl. 9 Abt. 4, 1. Halfte: 164 (1932). ISOLECTOTYPES. Switzerland: Graubiinden, Engadin, Samaden, Kies- und Sandalluvionen des Inn nahe beim Bahnhof. E. Frey 1636, 1931 (CANL 77611, 63390). Poelt: Pl. Graecenses No. 232% Stereocaulon arbuscula var. aberrans Asah., J. Jap. Bot. 19: 282 (1943). ISOLECTOTYPE. Japan: Honshu, Prov. Musashi, Nippara. I. Sasaki, 1940 (CANL 48276). = Leprocaulon arbuscula (Nyl.) Nyl. (Ch. strain II.) Stereocaulon argodes Nyl., Compt. Rend. Hebd. Séances Acad. SGPIES3"87 61876) ¢ ISOTYPE. New Zealand: Campbell Island. Filhol, 1874 (CANL 67862)" = S. argus J. D. Hook. & Taylor 344 Stereocaulon argus var. stenospermum Lamb, Farlowia 4: 460 (1955). HOLOTYPE. Argentina: Prov. Rio Negro, near foot of Frias Glacier S. of Lago Frias. On dry exposed rocks. I.M. Lamb 6088, 1950 (CANL 63410). ISOTYPE. (CANL 77621). Poelt: Pl. Graecenses No. 242. = S. stenospermum (Lamb) Lamb Stereocaulon azulense Yoshimura & W. Weber, Mycotaxon 27: 494 (1986). ISOTYPE. Ecuador: Galapagos Islands, Isla Isabela, SW coast, Cerro Azul. On boulders. W.A. Weber & H. Beck, 17 January 1984 (CANL 97944). Weber: Lich. Exs. Colo. No. 645. Stereocaulon botryosum f. depressum Frey, Rabenhorsts Kryptog "Flos 9 (4) Di 26 1Clo3 2). ISOLECTOTYPE. Norway: Nordland, Umbukten. On sunny places. A.H. Magnusson 9040, 1924 (CANL 70674). = S. depressum (Frey) Lamb Stereocaulon botryosum f. pygmaeum H. Magn., Ark. Bot. 33A: 91 (1946). ISOTYPE. Sweden: Lycksele Lappmark, Tarna, Brakkffallet. On irrigated rock. A.H. Magnusson 8084, 1924 (CANL 75018). = S. spathuliferum Vainio f. pygmaeum (H. Magn.) Lamb Stereocaulon capense Lamb, Lilloa 26: 416 (1953). HOLOTYPE. South Africa: Western Cape Province, Tulbagh Division, Great Winterhoek. On rock face. E. Esterhuysen 19862 p.p., 1951 (CANL 63391). = S. corticatulum Nyl. var. capense (Lamb) Lamb Stereocaulon colensoi var. reagens R&dsdnen, Ann. Bot. Soc. Z00l.=Bot.. Fenn .."Vanamot 22624 2(1932)r, ISOSYNTYPE? Chile: Fuegia occ., Fjordo Finlandia, supra rupium. H. Roivainen, 1929 (CANL 70923). = S. glabrum (MUll. Arg.) Vainio Stereocaulon condensatum var. sorediatum Harm., Bull. Acad. Int. .Géoqr.) Bot. 22: 5157161913) ' ISOTYPE. France: Hérault, La Salvetat. Sur des rochers schisteux, terreux. F. Marc (CANL 71016). Claud. & Harm.: Lich. Gall. Praecip. Exs. No. 474. Stereocaulon coralloides Fr., Fr.: Lich. Suec. Exs. No. 118 (1817). ISOTYPE. Sweden: no locality or date designated (CANL 70733). = S. dactylophyllum Floérke Stereocaulon coralloides var. flabellatum Frey, Rabenhorsis Kryptog. ‘Fl. 9 (4,1)3,196 (1932). ISOLECTOTYPE. Germany: Bayen, Bohmerwald, Bayrisch- Eisenskein, Hochbergplatte, schatlig. J. Hillmann, 2 October 1930 (CANL 70680) = S. dactylophyllum Flérke var. flabellatum (Frey) Grumn. 345 Stereocaulon coralloides var. occidentale H. Magn., Géteborgs Kungl. Vetensk. Samhdlles Handl. Ser. 4, 30: 27 (1926). ISOLECTOTYPE. Sweden: Bohuslan, Dragsmark. A.H. Magnusson, 1915 (CANL 70696). = S. dactylophyllum Flérke var. occidentale (H. Magn.) Grumn. Stereocaulon cornutum var. corallizans Lamb, J. Hattori Bot. Babe 43082 5270-0197 7)2 ISOTYPE. Costa Rica: Prov. Heredia, Cerro Central de Zurqui. C.W. Dodge, J. Valerio R., W.S. Thomas & R. Valerio 6119, 1929 (CANL 63389). Stereocaulon corticatulum var. complanatum Lamb, J. Hattori Bots? Labs 4327 289¢ (1977). ISOTYPE? New Zealand: Stidinsel, Otago, Taieri Mouth, Kuistenfelsen. J. Murray 9742 (1421 in protologue), 1958. (CANL 77612). Poelt: Pl. Graecenses No. 233. Stereocaulon corticatulum var. procerum Lamb, Farlowia 4: 462 (1955). HOLOTYPE. Argentina: Prov. Rio Negro, between Lago Frias and Frias Glacier. On rocks. I.M. Lamb 6086, 1950 (CANL 63400). ISOTYPE. (CANL 77619). = S. melanopotamicum Lamb Stereocaulon crambidiocephalum Lamb, J. Hattori Bot. Lab. 43: 290 (1977). ISOTYPE. Peru: Dept. Cuzco, Prov. Paucartambo, Llulluchayocc. On rock. C. Vargas 4301, 1944 (CANL 63392). Stereocaulon dendroides Asah., J. Jap. Bot. 36: 231 (1961). ISOTYPE. Japan: Shikoku, Mt. Kenzan. F. Fujikawa, 1934 (CANL 63393). Stereocaulon denudatum var. umbricola ("umbricolum") Frey, Rabenhorsts Kryptog. Fl. 9 (4,1): 134 (1933). ISOTYPE. Switzerland: Berner Oberland, Grimsel, way towards Nagelisgraéatli. On granite rock. E. Frey 1429, 1930 (CANL 63414). = S. vesuvianum Pers. var. nodulosum (Wallr.) Lamb f. umbricola (Frey) Lamb Stereocaulon dusenii Lamb, J. Hattori Bot. Lab. 43: 292 (1977). ISOTYPE. Chile: Magallanes, Isla Desolaci6én, Puerto Angosto. On rocks. P. Dusén 186, 1896 (CANL 63394). Stereocaulon esterhuysenae Lamb, Lilloa 26: 417 (1953). HOLOTYPE. South Africa: Western Cape Province, Tulbagh Division, Great Winterhoek. On a rock face. E. Esterhuysen 19862 p.p., 1951 (CANL 63395). Stereocaulon evolutoides var. paschaleoides Hav., Arbok. Univ. Bergen, Naturvitensk. Rekke 12: 1720 (1954). ISOTYPE. Norway: Granvin, Gamladaemmo, near Skalsaete. On gneiss boulders. J. Havaas (CANL 63409). 346 = S. saxatile H. Magn. f. paschaleoides (Hav.) Lamb Stereocaulon exutum Nyl., Lich. Japon. p. 18 (1890). ISOTYPE. Japan: Fusijama. E. Almquist, 1879 (Vega Exped., 1878-1880) (CANL 70910). Stereocaulon fastigiatum f. confluens H. Magn., Gdteborgs Kungl. Vetensk. Samhalles Handl., Ser. 4, 30: 35 (1926). ISOTYPE. Sweden: Torne Lappmark, par. Jukkasjarvi, Kopparasen. A.H. Magnusson 6073, 1921 (CANL 70726). = S. depressum (Frey) Lamb Stereocaulon fastigiatum var. dissolutum H. Magn., Gdéteborgs Kungl. Vetensk. Samhalles Handl. Ser. 4, 30: 36 (1926). LECTOPARATYPE. Norway: Nordland, Umbukten, Krabbfjdall. A.H. Magnusson 9040, 1924 (CANL 75616). = S. spathuliferum Vainio f. congestum (H. Magn.) Lamb Stereocaulon glareosum var. brachyphylloides Lamb, J. Wash. ACAG siSCive 4 LOG m Loo Ljts ISOTYPES. United States: Alaska, Brooks Range, Anaktuvuk Pass. On soil. G.A. Llano & N. Weber 527, 1949 (CANL 63397, 77614). Poelt: Pl. Graecenses No. 235. Stereocaulon halei Lamb, J. Hattori Bot. Lab. 43: 294 (1977). ISOTYPE. Malaysia: Pahang, unterhalb Ganong Brinchang. M.E. Hale, 1965 (CANL 77615). Poelt: Pl. Graecenses No. 236. Stereocaulon intermedium f. compactum Lamb, J. Hattori Bot. Vaberssceccol oy). ISOTYPES. United States: Alaska, Aleutian Islands, Rat Island, Amchitka, near Consantine Harbor. G.A. Llano 1277, 1949 (CANL 63385, 77617). Poelt: Pl. Graecenses No. 238. Stereocaulon japonicum var. subfastigiatum Asah., J. Jap. Botte 35:1289) (196077 ISOTYPE. Japan: Honshu, Prov. Owari, Kiso River, Inuyama. Y. Asahina 1201, 1938 (CANL 63398). Stereocaulon japonicum var. tokioense Lamb, J. Jap. Bot. 44: 265 (1969). HOLOTYPE. Japan: Honshu, Tokyo. On stone wall of the University. Y. Asahina 650, 1943 (CANL 63399). Stereocaulon leprocauloides Lamb, Bryologist 74: 193 (1971). ISOTYPES. Papua New Guinea: Morobe District, Mt. Kaindi near Wau. W.A. Weber & D. McVean, 19 June 1968 (CANL 33463, 77618) .. .Weber:* ) Lichy/ -Exs.°"Colov’ =NovF “299% ~Poelt:” “Pi: Graecenses No. 239. Stereocaulon melanopotamicum Lamb, J. Hattori Bot. Lab. 43: 2961977 )r. (see Stereocaulon corticatulum var. procerum Lamb). Stereocaulon meyeri f. tucumanum Lamb, J. Hattori Bot. Lab. As AO BT. hee 347 ISOTYPE. Argentina: Prov. Tucuman, Dept. Chicligasta, between La Cascada and Las Cuevas. On rock. T. Meyer, 1949 (CANL 63401). Stereocaulon montagneanum Lamb, J. Jap. Bot. 40: 272 (1965). ISOTYPE. Malaysia: Cameron Highland, On exposed rocks. M. Togashi 62236, 1-3 February 1962 (CANL 34168). Kurokawa: Lich. tRar. (CritstExss) Now142: Stereocaulon myriocarpum var. altaicum Lamb, J. Hattori Bot. Lab. 43: 225 (1977). ISOTYPE. Kazakhstan: Siberia, Gub. Semipalatinsk, Altai Mts., Katon-Karagai. P. Krylov & L. Sergievskaya 8, 1928 (CANL 74958). Stereocaulon nanodes f. schadeanum Lamb, J. Hattori Bot. Lab. Ape e242) (197.7) i ISOTYPE. Germany: Sachsen, Freiberg. A. Schade 1767, 1927 (CANL 63402). Stereocaulon novoarbuscula Asah., J. Jap. Bot. 19: 283 (1943). . ISOLECTOTYPE. Japan: Shikoku, Prov. Tosa, Mt. Otoyama. F. Fujikawa, 1931 (CANL 48290). = Leprocaulon pseudoarbuscula (Asah.) Lamb & Ward Stereocaulon octomerelloides Asah., J. Jap. Bot. 45: 68 (1970). ISOTYPE. Japan: Honshu, Prov. Etigo, Sasagamine Pasture. Y. Asahina 1521, 1949 (CANL 63403). = S. octomerellum Miill. Arg. Stereocaulon papuanum Lamb, J. Hattori Bot. Lab. 43: 225 (197%) & ISOTYPE. Papua New Guinea: Central Division, Mt. Albert Edward. L.J. Brass 4394a, 1933 (CANL 63404). Stereocaulon paschale var. alpinum f. flabellans Lamb, Farlowia 4: 457 (1955). HOLOTYPE. Argentina: Prov. Rio Negro, Cerro Rigi near Lago Frias. On the ground between rocks and stones. I.M. Lamb 5787, 1950 (CANL 63396). ISOTYPE. (CANL 77613). Poelt: Pl. Graecenses No. 234. = S. glabrum (MUtill. Arg.) Vainio f. flabellans (Lamb) Lamb Stereocaulon paschale f. colligatum Lamb, Rep. (Annual) Natl. Mus. Canada, 1952-53, Bull. No. 132: 273 (1954). HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton Island, Ingonish, Middle Head. On granitic rocks. I.M. Lamb 6946, 29 July 1952 (CANL 10301). = S. paschale (L.) Hoffm. f. paschale Stereocaulon paschale subsp. evolutoides f. laxatum Lamb, Rep. (Annual) Natl. Mus. Canada, 1952-53, Bull. No. 132: 275 (1954). ISOTYPE. Sweden: Varmland, Sillerud, Jarnsj6n. A.H. 348 Magnusson, 1912 (CANL 71061). Stereocaulon paschale v. evolutoides f. sorediatum H. Magn., G6teborgs Kungl. Vetensk. Samhalles Handl. Ser. 4, 30: 36 (1926). ISOTYPE. Sweden: Vdarmland, par. Sillernd, Jarnsj6n. A.H. Magnusson, 1912 (CANL 71078). = S. saxatile H. Magn. f. sorediatum (H. Magn.) Lamb Stereocaulon paschale var. serpens Th. Fr., De Stereoc. et Pilophor. Comment p. 33 (1857). ISOTYPE. Sweden: Uppsala, Vitulfsberg. Th. Fries, 1852 (CANL 70990). = S. paschale (L.) Hoffm. Stereocaulon patagonicum Lamb, Farlowia 4: 454 (1955). HOLOTYPE. Argentina: Prov. Chubut, Lago Menéndez, W. end. On sandy gravelly shore of lake. I.M. Lamb 5790, 1950 (CANL 63405). ISOTYPE. (CANL 77616). Poelt: Pl. Graecenses No. 237. = S. glabrum (MUll. Arg.) Vainio Stereocaulon patagonicum f. subirregulare Lamb, Farlowia 4: 456 (1955). HOLOTYPE. Argentina: Prov. Rio Negro, Lago Frias, Cerro Rigi, near summit. In cleft between stones. I.M. Lamb 5789, 1950 (CANL 63406). = S. glabrum (MUll. Arg.) Vainio Stereocaulon philippinense R&sd&dnen, Arch. Soc. Zo0ol.-Bot. Fenn. "Vanamo" 3: 79 (1949). ISOTYPE. The Philippines: Negros, Canlaon Volcano. Terricola. E.D. Merrill 6874, April 1910 (CANL 71010). Stereocaulon pileatum f. macrum H. Magn., Géteborgs Kungl. Vetensk. Samhdlles Handl. Ser. 4, 30: 70 (1926). ISOTYPE. Sweden: Ljung, Kolbengtsered. On rock. A.H. Magnusson 9513, 13 August 1925 (CANL 71041). = S. pileatum Ach. Stereocaulon pileatum var. nipponicum Lamb, J. Hattori Bot. LAD Ina Ss ees iy HOLOTYPE. Japan: Jiushiu, Prov. Higo, Aida. K. Maebara, 1927 (CANL 63407). Stereocaulon proximum var. gracilius Miill. Arg., Rev. Mycol. (Toulouse) 1: 164 (1879). ISOTYPE? Colombia "Nova Granata": La Horqueta prope Dolores. André (coll. no. = 2813 p.p. in holotype), 1876 (CANL 77638). = S. ramulosum (Sw.) Rausch. var. gracilius (Miill. Arg.) Lamb Stereocaulon pseudoarbuscula Asah., J. Jap. Bot. 19: 282 (1943). ISOLECTOTYPE. Japan: Honshu, Prov. Musashi, Titibu. Y. Asahina 1249, 1933 (CANL 48292). = Leprocaulon pseudoarbuscula (Asah.) Lamb 349 Stereocaulon pseudomassartianum Lamb, Bot. Jahrb. Syst. 86: 246 (1967). ISOSYNTYPES. The Philippines: Luzon, Mountain Prov., Ifugao Subprov. On trail between Banaue and Mt. Polis. On earth. A.W. Herre, 1923 (CANL 71040, 77620). Poelt: Pl. Graecenses No. 241. Stereocaulon pygmaeum Vainio, Voyage S. Y. Belgica, Bot. p. 15 (1903). ISOTYPE. Antarctica: Graham Land (Palmer Peninsula), Cape Anna Osterrieth. Exped. Antarctica Belge 1897-99, No. 201. (CANL 12807). = Lecania brialmontii (Vainio) Zahlbr. Stereocaulon ramulosum var. pulvinare f. crebratum Lamb, J. Hatcoriebots Labosd43cee85in(1977))% ISOTYPE. New Zealand: South Island, Port Chalmers near Dunedin. On rock. J.S. Thomson D.15, T. 696 (CANL 63408). Stereocaulon speciosum Lamb, Farlowia 4: 458 (1955). HOLOTYPE. Argentina: Prov. Chubut, Lago Menendez. On mossy gravelly soil. I.M. Lamb 6069, 1950 (CANL 63387). ISOTYPE. (CANL 77623). Poelt: Pl. Graecenses No. 244. = §. verruciferum Nyl. Stereocaulon speciosum var. surreptans Lamb, Farlowia 4: 459 G1955)% HOLOTYPE. Argentina: Prov. Rio Negro, Cerro Rigi near Lago Frias, near summit. On detritus. I.M. Lamb 6098, 1950 (CANL 63388). ISOTYPE. (CANL 77624). Poelt: Pl. Graecenses No. 245. = S. verruciferum Nyl. var. surreptans (Lamb) Lamb Stereocaulon subalbicans Lamb, Bryologist 60: 220 (1957). ISOTYPE. Chile: Prov. Coquimbo, La Serena, Cerro Los Loros. R. Santesson 2530, 1940 (CANL 48278). = Leprocaulon subabicans (Lamb) Lamb & Ward Stereocaulon subcoralloides f. sorediascens Lamb, J. Hattori Bot. Lab. 43: 235 (1977). HOLOTYPE. Russia: Karelia ladogensis, Kurkijoki, Kuuppala, Jaavuori. On rock. V. R&s&nen, 1931 (CANL 63411). Stereocaulon tennesseense H. Magn., Ark. Bot. 30A: 48 (1941). ISOTYPE. United States: Tennessee, Great Smoky Mountains, near Alum Cave. On moist rocks. G. Degelius, 1939 (CANL 75021). Stereocaulon tennesseense var. nigrofastigiatum Lamb, J. MaAGlLori sSOG lab. b4oc 20 Om CLO Ia): HOLOTYPE. United States: New York State, Adirondacks, Mt. Colden near Lake Placid. On rocks. J.L. Lowe, 1952 (CANL 63412). ISOTYPE. (CANL 77622). Poelt: Pl. Graecenses No. 243. Stereocaulon tomentosum var. compactum Frey, Rabenhorsts 350 Krypts RL. -9°7°(471)2"187,"(1932)". ISOTYPE? Germany: Sachsen, Brambach im Vogtland Sohle in altem Steinbruch, Granit. A. Spindler, 1908 (CANL 74962). Stereocaulon tomentosum var. capitatum Lamb, J. Hattori Bot. Lab. 43:3) '239) (1977 )% HOLOTYPE. Argentina: Prov. Tucuman, Valle de Tafi, W. slope of Cumbre Potrerillo. I.M. Lamb 5309, 1947 (CANL 63386). Stereocaulon verruculigerum var. formosanum Asah., J. Jap. Bot.23557).295> (1950)* ISOLECTOTYPE. Taiwan: Rengechi. Y. Asahina 222, 1925 (CANL 63413). = §. verruculigerum Hue var. verruculigerum Stereocaulon vesuvianum f. verrucosum Lamb, J. Hattori Bot. Labs! 43.59247°.(1577) & ISOTYPE. United States: Alaska, Aleutian Islands, Attu. G. A. Llano 1463 p.p., 1949 (CANL 77625). Poelt: Pl. Graecenses No. 246. Stereocaulon vimineum Th. Fr., De Stereoc. & Pilophor. Comment p. 13 (1857). ISOTYPE? Mexico: Tiuzutlan. Liebmann 73, 1841 (CANL 75624). = S. ramulosum (Sw.) Rausch. Stereocaulon weberi Lamb, J. Hattori Bot. Lab. 43: 258 C1977 3% ISOTYPES. Ecuador: Galapagos Islands, Isla Santa Cruz, summit of Mount Crocker. W.A. Weber & J. Lanier, 15 April 1976 (CANL 58209, 77626). Weber: Lich. Exs. Colo. No. 494; Poelt: Pl. Graecenses No. 247. Sticta oregana Tuck., Bull. Torrey Bot. Club 5: 20 (1874). ISOTYPE. United States: Oregon. E. Hall, 1871 (CANL 68763). Reliq. Tuck. No. 90. = Lobaria oregana (Tuck.) MUll. Arg. Strangospora senecionis Lambinon & Vézda, Folia Geobot. Phytotax., Praha 14: ' 206 (1979). ISOTYPE. Zaire: Prov. Kivu, Birunga montes, in planitie Rukumi montis ignivomi Karisimbi. Ad corticem et lignum truncorum vetustorum. J. Lambinon 72/230, 22 January 1972 (CANL 73820). Vézda: Lich. Sel. Exs. No. 1657. Sulcaria badia Brodo & D. Hawksw., Opera Bot. 42: 146 (1977). ISOTYPE. United States: Oregon, Philomath. On old apple trees. F.P. Sipe 669 (CANL 29671). Note: This specimen is a portion of the same number designated by Gyelnik as the type of Alectoria pseudofuscescens Gyelnik. The holotype in BP, however, is quite different. (fide Brodo & Hawksworth 1977. Opera Bot. Aas 47) s Sulcaria isidiifera Brodo, Mycotaxon 27: 115 (1986). HOLOTYPE. United States: California, San Luis Obispo Co., Los 351 Osos Oaks State Reserve, Los Osos Valley Road. C. Bratt & J. Larson 3871, 8 January 1984 (CANL 88639). PARATYPE. United States: California. M.E. Hale 57816 (CANL 80150). Tephromela elixii Kalb, Kalb: Lich. Neotr. Fasc. 12: 15 (1991). ISOTYPE. Venezuela: Merida, Distr. Rangel, Paramo zwischen der Laguna Mucubaji und dem Pico Mucufiuque, etwa 15 km SE von Apertaderos. K. & A. Kalb, 16 August 1989 (CANL 103793). Kal bsvLicn. aNeOtr..iNO. 522). Tephromela muscicola Kalb, Kalb: Lich. Neotr. Fasc. 10: 15 (1991). ISOTYPE. Ecuador: Tungurahua, Siidhang des Mt. Tungurahua, nérdlich von Bafios. Uber Moosen in liickiger Vegetation. K. & A. Kalb, 16 August 1987 (CANL 102736). Kalb: Lich. Neotr. No. 447. Tephromela nashii Kalb, Kalb: Lich. Neotr. Fasc. 12: 16 (1991). ISOTYPE. Mexico: Baja California, 10 km N El Rosario in Cafion del Rosario. On small pebbles. K. & A. Kalb & T. Nash, 28 December 1990 (CANL 103794). Kalb: Lich. Neotr. No. 523. Tephromela septentrionalis Hertel & Rambold, Bot. Jahrb. Syst. 107: 495 (1985). ISOTYPE. Greenland: W-Grénland, Gemeinde Umanak. Uber eisenreichem Silikatgestein. J. Poelt & H. Ullrich, August 1983 (CANL 95897). Hertel: Lecideaceae Exs. No. 178. Tephromela tropica Kalb, Kalb: Lich. Neotr. Fasc. 10: 15 (1988). ISOTYPE. Kenya: Central Province, Nanyuki District, zwischen Naco Moru und Nanyuki. K. Kalb & A. Schrégl, 18 August 1985 (CANL 102737). Kalb: Lich. Neotr. No. 448. Tephromela velloziae Kalb, Kalb: Lich. Neotr. Fasc. 8: 16 (1984). ISOTYPE. Brazil: Bahiz, Chapada Diamantina, Serra do Tombador, etwa 1 km vor der Stadt Morro do ChapeG. K. Kalb, 18-20 July 1980 (CANL 89464). Kalb: Lich. Neotr. No. 349. Thelotrema brasiliana Hale in Kalb, Kalb: Lich. Neotr. Fasc. Py Sat 983 \,. ISOPARATYPE. Brazil: Sao Paulo, Ilha de Sao Sebastido, etwa 130 km S6stlich von Sado Paulo. Westhang des Morro das Tacas. In einem sehr feuchten und dunklen Regenwald an einem Fluss. K. Kalb, 6 July 1979 (CANL 85936). Kalb: Lich. Neotr. No. 298. Toninia arctica Timdal, Opera Bot. 110: 37 (1991). HOLOTYPE. Canada: Northwest Territories, Franklin District, Banks Island. G.W. Scotter 30146, 21-24 July 1979 (CANL 78540). ISOTYPE. (CANL 78541). 352 PARATYPE. Canada: Yukon. G.W. Scotter 19501 (CANL 54403). Toninia ruginosa var. andicola Lamb, Rhodora 56: 147 (1954). HOLOTYPE. Argentina: Prov. Catamarca, Nevados de Anconquija, Quebrada de los Cazadores. On soil. I.M. Lamb 5599, 24 November 1948 (CANL 6086). = T. bullata (Meyen & Flotow) Zahlbr. Toninia ruginosa subsp. pacifica Timdal, Opera Bot. 110: 90 (1991). ISOTYPE. United States: California, Santa Barbara Co., Channel Islands, Santa Cruz Island, West end of the island. On the rim of the "Trailer Barranca", in rock crevices. W.A. Weber & C. Bratt, 8 January 1986. (CANL 97962). Weber: Lich. Exs. Colo. No. 663. Toninia tristis subsp. canadensis Timdal, Opera Bot. 110: 113 (1991). HOLOTYPE. Canada: Northwest Territories, District of Mackenzie, Dolomite Lake. G.W. Scotter 6097, 6 August 1965 (CANL 41669). PARATYPES. Canada: British Columbia. I.M. Brodo 21333 (CANL 86793); Northwest Territories. G.W. Scotter 8283 & 8807 (CANL 43173 & 41670). Toninia tristis subsp. pseudotabacina Timdal, Opera Bot. 110: 1a (1991) PARATYPES. Cyprus: Larnaca. A. Vézda (CANL 99203). Vézda: Lich. Sel. Exs. No. 2262; Spain: Prov. Larida. J. Poelt & A. Vézda (CANL 89203). Trapeliopsis hainanensis Hertel, Herzogia 5 (3-4): 460 (1981). ISOTYPE. China: Prov. Guangdong, Insel Hainan. Bergwald- Schutzgebiet im Gebirge Jian Fung Ling im Kreise Ledong. H. Hertel, 23 May 1980 (CANL 77212). Hertel: Lecideaceae Exs. Now -59:. Trapeliopsis pseudogranulosa Coppins & P. James, Lichenologist 16: 259 (1984). ISOTYPE. Canada: British Columbia, Vancouver’ Island, MacMillan Provincial Park on Rt. No. 4, 8 miles E of Alberni. On Pseudotsuga. I.M. Brodo 7947, 5 June 1966 (CANL 53995). Brodo: Lich. Canad. Exs. No 129. Umbilicaria lambii Imsh., Bryologist 60: 232 (1957). HOLOTYPE. Canada: British Columbia, Mt. Assiniboine Provincial Park, Sunburst Lake. On a large rock. I.M. Lamb 6584, 3 August 1951 (CANL 66543). Umbilicaria phaea Tuck., Lich. Calif. p. 15 (1866). ISOTYPE. United States: California. On rocks of Pacific Coast. H.N. Bolander 11, 1866 (CANL 68767). Reliq. Tuck. No. 94. Usnea acromelana Stirton, Trans. & Proc. New Zealand Inst. 353 3035388 (1897)* ISOTYPE. New Zealand: Christchurch, Canterbury, Selwyn Gorge. "On trees" (saxicolous!). T.W.N. Beckett L.11, May 1894 (CANL 16943). Usnea durietzii Mot., Lich. Gen. Usnea Stud. Monogr., Pars Systems: 11s) 503 1(193 7) ISOTYPE. Chile: Regio Magellanica, Insula Elisabetha. Ad saxa erratica granitica. Lechler (CANL 17098). Usnea eulychniae Follm., Nova Hedwigia 14: 261 (1967) ISOTYPE. Chile: Prov. Antofagasta, zerstreut im Dornenwerk von Sdaulenkakteen im Tornabenietum intricatae. I.A. Follmann- Schrag & G. Follmann, September 1965 (CANL 64294). Follmann: Lich. Exs7ysel. = No.=60: Usnea kuehnemanrii Mot. ex Lamb, Farlowia 4: 467 (1955). ISOTYPE. Argentina: Patagonia, Prov. Neuquén, Parque Nacional Lanin, Lago Quillen. O. Kuhnemann 796, 1943 (CANL 104552). Usnea neoguineensis Asah., J. Jap. Bot. 43: 496 (1968). ISOTYPE. Papua New Guinea: Morobe District, Middle Creek logging area, Bullolo. S. Kurokawa 5787, 11-12 November 1965 (CANL 34214). Kurokawa: Lich. Rar. Crit. Exs. No. 149. Usnea neuropogonoides Mot., Lich. Gen. Usnea Stud. Monogr., pars System. I: 73° (1936). ISOTYPE. Argentina: Rio Fosiles, in rupibus. P. Dusén, April 1905 (CANL 17188). Usnea nidularis Asah., J. Jap. Bot. 44: 3 (1969). ISOTYPE. Japan: Honshu, Prov. Iwaki, Matsukawa-ura, Sohma- gun. On Pinus thunbergii. S. Kurokawa 58098-b, 18 July 1958 (CANL 65983). Kurokawa: Lich. Rar. Crit. Exs. No. 346. Usnea pseudoceratina Mot., Folia Geobot. Phytotax. 10: 326 (1975). ISOTYPE. Tanzania: Southern Highlands, Poroto Mountains, NW end of Kitulo/Elton/Plateau. T. Pécs 6750/G, 10 August 1972 (CANL 57344). Usnea tanzanica Mot., Folia Geobot. Phytotax., Praha 10: 327 (1975). ISOTYPE. Tanzania: Prov. Mbeya, in monte ignivomo Rungwe. Ad ramulos arborum in regione subalpina. T. Pécs 6507, 21 January 1972 (CANL 57292). Vézda: Lich. Sel. Exs. No. 1322. Verrucaria bagliettoi Servit, Ann. Mus. Civico Storia Nat. Giacomo Doria 64: 53 (1950). ISOTYPE. Italy: Genova, Val Bisagno, Rupicola loco Fullo. Sbarbaro, November 1950 (CANL 692). Verrucaria bisagnoensis Servit, Sborn. Nar. Mus. v Praze, Rada B, Pfrir. Védy. 9: 16 (1949). ISOTYPE. Italy: Genova, Val Bisagno, Prato. Sbarbaro, April 1949 (CANL 693). 354 Verrucaria boccana Servit, Sborn. Nar. Mus. v Praze, Rada B, Prir. Védy. 9: 17 (1949). ISOTYPE. Italy: Dalmatia meridionalis, Hercegnovi 40 nm, Silic. M. Servit, 1929 (CANL 694). Verrucaria contardinis Servit, Ann. Mus. Civico Storia Nat. Giacomo Doria 66: 236 (1953). ISOTYPE. Italy: Friuli, Zomeais, prope Tarcento. A. Contardo, 25 December 1951 (CANL 711). Verrucaria diplotommoides Servit, Preslia 24: 355 (1952). ISOTYPE. Italy: Genova, Staglieno. Sbarbaro, 30 March 1951 (CANL 718). Verrucaria durietzii Lamb, Lilloa 14: 205 (1948). HOLOTYPE. New Zealand: Auckland Islands, Port Ross, innermost small peninsula in Laurie Harbour. G.E. Du RietzZ 2225 b:1, 27 March 1927 (CANL 721). PARATYPE. New Zealand. G.E. & G. Du Rietz 2070:2 (CANL 724). Verrucaria durietzii f. rhabdota Lamb, Lilloa 14: 206 (1948). HOLOTYPE. New Zealand: E. Falkland Islands, Berkeley Sound, Port Louis. On seashore rocks. I.M. Lamb 2939, 9 February 1946 (CANL 720). PARATYPE. New Zealand. I.M. Lamb 2877, 29 January 1946 (CANL 723 Verrucaria fuscella f. omblensis Servit, Stud. Bot. Cech. 7: 59 (1946). ISOTYPE. Italy: Dalmatia, Ragusa, Ombla, Gionchello. Latzel 104, 1908 (CANL 802). = V. omblensis (Servit) Servit Verrucaria gorzegnoensis Servit, Webbia 8: 413 (1952). ISOTYPE. Italy: Cuneo, Gorzehno, le Langhe. Sbarbaro, September 1951 (CANL 744). Verrucaria haeyrenii ("hayrénii") Erichsen, Memoranda Soc. Faunavhl. Fenn .l2ice Se (1937 ) i. ISOTYPE. Finland: Nyland, Ekends, Tvdrminne, Langskar. E. Hayrén & C.F.E. Erichsen, 29 June 1933 (CANL 747). Verrucaria imperfecta Servit, Preslia 24: 359 (1952). ISOTYPE. Italy: Genova, Val Risagno, Ligorna. Sbarbaro, 21 September 1950 (CANL 749). Verrucaria incompta Sersit, Ann. Mus. Civico Storia Nat. Giacomo Doria 66: 237 (1953). ISOTYPE. Italy: Savona, Spotorno, Rupicola loco Lajolo. Sbarbarc, January 1952 (CANL 752). Verrucaria lacustris Lamb, Farlowia 4: 424 (1955). HOLOTYPE. Argentina: Patagonia, Prov. Chubut, Lago Verde near Futalaufquen. On stone. I.M. Lamb 5884, 1 February 1950 (CANL 760). 355 Verrucaria langhensis Servit, Ann. Mus. Civico Storia Nat. Giacomo Doria 64: 48 (1950). ISOTYPE. Italy: Gorzegno, Le Langhe, Piemonte. Sbarbaro, September 1947 (CANL 764). Verrucaria latebrosoides Servit, Sborn. Nar. Mus. v Praze, Rada B, Piir. Védy. 9: 31 (1949). SYNTYPE. Italy: Genova, Val Bisagno, Ligorna. Sbarbaro, 23 March 1950 (CANL 766). Verrucaria ornata Servit, Stud. Bot. Cech. 11: 119 (1950). ISOTYPE. Italy: Genova, Mt. Fasce. Sbarbaro, 1947 (CANL 803). Verrucaria pseudomacrostoma Servit, Webbia 8: 415 (1952). ISOTYPE. Italy: Cuneo, Langhe, Gorregno, 400 mt. Cortemilia. Sbarbaro, September 1951 (CANL 809). Verrucaria putnae Servit, Preslia 24: 374 (1952). ISOTYPE. Romania: Putna, Sovejanka. Cretzoiu, 1935 (CANL 613). Verrucaria rapallensis Servit, Stud. Bot. Cech. 11: 121 (1950). ISOTYPE. Italy: Genova, Rapalleo, Montallegro 500 mt. rupicola. Sbarbaro, October 1949 (CANL 814). Verrucaria santensis Nyl., Acta Soc. Sci. Fenn. 7: 489 (1863). ISOLECTOTYPE. United States: South Carolina, Santee Canal. H.W. Ravenel, 1850 (CANL 23286). = Pyrenula santensis (Nyl.) Miill. Arg. Verrucaria savonensis Servit, Webbia 8: 416 (1952). ISOTYPE. Italy: Savona, Albisola marina, loco Seitun. Sbarbaro, August 1951 (CANL 822). vézdaea rheocarpa Poelt & Débbeler, Bot. Jahrb. Syst. 96: 347 (1975). ISOTYPE. Austria: sitidliches Burgenland. Uber verschiedenen Moosen. J. Poelt & P. Ddébbeler, 17 March 1974 (CANL 63545). Poelt: Lich. Alpium No. 293. Wawea fruticulosa Henssen & Kantvilas, Lichenologist 17: 86 (1985). ISOTYPE. Australia: Tasmania, Cradle Mountain National Park. On burned trunk of Nothofagus cunninghamii. W.A. Weber & D. McVean, 22 February 1968 (CANL 52202). Weber: Lich. Exs. COLO. INO. 452). Xanthoparmelia cylindriloba Knox, J. South African Bot. 49: 147 (1983). ISOTYPE. Kenya: Regio montis Kenya, in parte superiore vallis Teleki. Ad terram glacie destructam. O. Hedberg, 27 July 1948 (CANL 93769). Vézda: Lich. Sel. Exs. No. 2063. Xanthoparmelia lipochlorochroa Hale & Elix, Mycotaxon 34: 551 356 (1989). ISOTYPE. United States: Wyoming, Lincoln Co., 15 mi. S of Kemmerer, Red Desert area. On sandy soil. B.J. Buckingham S 22155, 26 August 1959 (CANL 20819). Weber: Lich. Exs. Colo. NO. ‘29. Xanthoparmelia mollis Hale, Mycotaxon 27: 585 (1986). ISOTYPE. South Africa: Natal. Port Shepstone District, Oribi Gorge Nature Reserve. On exposed sandstone outcrops. M.E. Hale 74043, 3 February 1986 (CANL 99574). Almborn: Lich. African, iNos 1237 Xanthopeltis rupicola R. Sant., Svensk Bot. Tidskr. 43: 560 (1949). ISOTYPE. Chile: Prov. Santiago, San José. On rocks. R. Santesson 2420, 12 June 1940 (CANL 18087). Xanthoria filsonii Elix, Elix: Lich. Austral. Exs. Fasc. 7 Nove 17491988): ISOTYPE. Australia: Victoria, Hopetoun, Patchewollock Road, 5 km NE of Hopetoun. On Rhagodia. J.A. Curnow & H. Lepp 1422, 14:-March 1987. * (GANL (98903),. Elix: Lich: Austral. Exs. No. aes Xanthoria polycarpa var. maritima Lamb, Rep. (Annual) Natl. Musi; Canada,, 1992-54. Bull 213238 306 mo 54 Fr HOLOTYPE. Canada: Nova Scotia, Victoria Co., Cape Breton Island, Ingonish, Middle Head. On granite rocks. I.M. Lamb 6952, 29 July 1952 (CANL 18084). Zahlbrucknerella patagonica Henssen, Lichenologist 9: 41 (1977). ISOTYPE. Argentina: Prov. Santa Cruz, Lago Argentino, Calafate, Cuevas de Hualichu. Sickerwasserflachen auf Schieferfels in Uferndhe. A. Henssen & G. Vobis 24525a, 18 December 1973 (CANL 102535). Henssen: Lich. Cyanoph. Fungi Sax see NOs) 6 Us ISOPARATYPE. Argentina. I.M. Lamb 5858 p.p. (CANL 27339). ACKNOWLEDGEMENTS To thank (Drow. Mee Brodo and Drow rk. Kya relandm.or valuable suggestions and comments and for corrections to the manuscript, Dr. T. Ahti for critical review of this paper. I also thank Miss L. Ley for checking the abbreviations of periodicals. 357 REFERENCES Bridson, ei) ey 399019, Botanico-Periodicum- Huntianum/Supplementum. Carnegie-Mellon University, Pittsburgh. Egan, R.S., 1987. A fifth checklist of the lichen-forming, lichenicolous and allied fungi of the Continental United Stated and Canada. Bryologist 90: 77-173. Egan, R.S., 1989. Changes to the "Fifth Checklist of the Lichen-Forming, Lichenicolous and Allied Fungi of the Continental United States and Canada." Edition I. Bryologist 92: 68-72. Egan, R.S., 1990. Changes to the "Fifth Checklist of the Lichen-Forming, Lichenicolous and Allied Fungi of the Continental United Stated and Canada." Edition II. Bryologist 93: 211-219. Egan, R.S., 1991. Changes to the "Fifth Checklist of the Lichen-Forming, Lichenicolous and Allied Fungi of the Continental United States and Canada." Edition III. Bryologist 94: 396-400. Re cue, iow! CWE Mute ; wav 4 . ; iF Pe ade m4 ay: bly’ *? Pele hss Pe a te ws ck De eee Pi Lie j id ; §, t ) relate 5 VN ae Op eehl ak OE : J¥ va e a) % 7 Na Ws CT aA eo i Rt att i mat (i i, uM a ay ahi. Pet WAN ie. BAG ne! ; oh iv ‘ ! ak ys Ni bee Naas sha tt HEA ane abet ¥ Abas Pag ks vee it a, oe ly wai ay lite ' a Ayes ftp ' ; YN ‘ae S O NSN fy ‘alps ML an as is | a oka mpnate wt aA seh BT a man IER ay Mat ee; , Ht en Bie hen hh ti ; * i? . b Pe ae biga) | th Me ne ny hy oe Laivarly ate) vs f A wh GAR aes as eto he eh f CE COC Sa Reta byt veu ie j : ay alt A ‘ ‘ Ay As) | wiih 1 an rw | 4 PS fl e ; wu ‘ ; ih en als ui yi ‘ip RAR GOR? ite Ue ve ty \" Ane gt aren m) aia brat i Ae (Weta ical ba eis CAUAMAMS Sisal Aaa" ts va We : an 5 Me Ck tat SA ip! y ' el ee i A Ye } ’ heral a) 5 ne iv { ANN \ A Hy Bow a ny ty Wy PA Mast, aay he, M me awe vat ey ified pe ai Me act ie ant #0 wh isu Ni Ms pai 1 oo ie i vn eh in rae Y a ivr Mi iy Pal We va a \ hs pe tal Ais y ty \y bi rh : y US Viet geal mn a: gre bd herons a aed ok ia , A \ ‘ = WI ¥ 3 4 ; - ve 7 ‘, ‘ . ‘a ,? aa ry Peay yh ae ey i ; j i La en ' ‘ 1 A A ivy nay Not ie i 7. ies 7 | bal rr eat i iy ay wi 1" > a a fae 1 Lie tn Aen! te aon han, tans a anes Lvhd wi ink ‘a aa nde iiaee vite te oneal GER ant Ar Magi Ne ane end Rayent ia uth wh are i edged yaa ‘ he 9 Hel Nid aaa wit Ay ah Sana nee “! Llane i y oe a +, 7 2 “had J 1 y ‘ i PPA iaie ug bat ees lied “PY iy xl | ied i r Re ie | ' Me Pia OR VN Y Amun ite: ty i) My AH i Ca, Al, A } vu | Whaat vast wf, a i ie re . ieee LOA Ay ian on swhioh a ae ny CIN A ACM MT OF ie: ih 1 Bt ne Lie, Hs (Waa \ s Mer ‘ ‘ {a3 " en! ; 1 / j ’ i ita yt | ¥ | uh ie, a ‘1 yee i y " A “he eter: Mi b th . mer - ' ; n * * #* Ni AG Me 4 ae, i Be remanent) cote 4 ; : wee ad ui ri ' ’ o ed hyd a rl ‘ ’ ta BAM Mi if hi WE i sek VR) + es a :. a iar i , , s| F | y ‘ ; ; ty , iy } 4 Mv y ANG) Beak hh. Fil ba y ie i yt ahd “ld tte i] Me Aes) * if yal os j i i? a 4 vo Neh s rs it -a4) wi 5 aa : 3 On fi i ‘ vy A ‘ -»# A fi, ; nen ie ay om bie) eK ' Ay bude ae Qi Pissivares a “oy ies sR ’ ¥ wisi : ] TH i ad - i yay vA Mes th y es Aide Jehan i mi oy ; 7 ee " ee alts ah ca : Mp nwt ' ety sae Pea Se rie alspta Oy : ; ' ty A ; ne 7 1 nae, 1p Tey 4 MY COTAXON Volume XLVI, pp. 359-366 April-June 1993 NEOSARTORYA PRIMULINA, A NEW SPECIES OF FOOD- BORNE ASCOMYCETES Shun-ichi UDAGAWA!, Noritsuna TOYAZAKI¢ and Haruo TSUBOUCHI> 1 Nodai Research Institute, Tokyo University of Agriculture, 1-1-1, Sakuragaoka, Setagaya- ku, Tokyo 156, Japan 2 Public Health Research Institute of Kobe City, 4-6, Minatojimanaka-machi, Chuo-ku, Kobe 650, Japan | 3 Nagoya City Health Research Institute, 1-11, Hagiyama-cho, Mizuho-ku, Nagoya 467, Japan ABSTRACT A new species of Neosartorya, N. primulina Udagawa, Toyazaki et Tsubouchi, isolated from a canned oolong tea beverage in Japan, is de- scribed and illustrated. This fungus is char- acterized by its restricted growth on Czapek agar, chalky-buff ascomata, and lenticular ascospores with a very irregular ornamentation composed of several narrow crests and verrucose hemispheres. Recently some widely-distributed species of Neosarto- rya have been incriminated as spoilage agents in commer- cially processed fruit products (Kavanagh et al., 1963; Beuchat, 1986; Conner and Beuchat, 1987: Scott and Bernard, 1987; Gomez et al., 1989; Nielsen et al., 1989; Samson, 1989; Splittstoesser and Churey, 1989; Samson et al., 1992). In an earlier paper (Udagawa et al., 1991), Neosartorya hiratsukae Udagawa, ITsubouchi et Horie was reported as the new causal agent of spoilage of pasteur- ized aloe beverage in Japan. A variety of commercial pas-— teurized beverages: which were undergoing spoilage were 360 selectively surveyed for further isolation of heat resist- ants. Pasteurized beverages (packaged in cans) by com- mercial processors for fungal isolation. Fungal mycelia were removed from the beverage by straining through a sterile membrane filter by aseptically transferred to chloramphenicol (100 ug/1)-potato dextrose agar. During the course of this survey, several heat re- sistant fungi were encountered. Among them one isolate representing a hitherto undescribed species of Neosartorya (Malloch and Cain, 1972; Kozakiewicz, 1989; Peterson, 1992) was found which is herein described and illustrated, Neosartorya primulina Udagawa, Toyazaki et Tsubouchi, Sp. nov. GiGicge hee St. Anam. Aspergillus primulinus Udagawa, Toyazaki et Tsubouchi, anam. sp. nov. Coloniae in agaro Czapekii paulo restrictae, aliquan- tum floccosae, plus minusve radiatim sulcatae, laxae, flavo-albae; conidiogenesis sparsa; reversum incoloratum vel pallide flavum. Coloniae in "Czapek yeast extract agar (CYA)" celeriter crescentes, velutinae, conspicue sulcatae, saepe zonatae, ex coacto mycelio basali tenuiter constantes, numerosis ascomatibus formantes, granulares, flavo-albae vel pallide flavae vel primulinae; conidio- genesis varians, viridi-grisea vel avellanea; reversum pallide flavum vel rubro-brunneum vel brunneo-aurantiacum. Coloniae in agaro maltoso effusae, floccosae, parum zon- atae, tenues, laxae, abundantibus ascomatibus formantes, pallide flavae; conidiogenesis inconspicua; reversum incoloratum vel pallide flavum. Ascomata non-ostiolata, superficialia, dispersa, cretacea vel bubalina, globosa vel subglobosa, 100-250 um diam, hyphis brunneis laxe intricatis circumdata, lente maturescentia; peridium membranaceum, ‘textura epidermoi- dea’ et "textura angularis'; stratum externum ex cellulis brunneis, irregularibus, incrassatis, 5-/.5 um latis com- positum. Asci octospori, brevi-catenati, subglobosi vel plus minusve pyriformes, 12-15(-17.5) x 11-13(-15) um, evanescentes. Ascosporae hyalinae, lenticulares, sine cristis 4.5-5(-5.5) x 3.8-5 um, cristis numerosis angusta- tis irregulatim ornatae. Capitula conidica parva, brunneo-grisea, laxe radian- tia; conidiophora ex mycelio basali vel interdum hyphis aeriis ascendentes; stipites brunneoli, 200-600 x 2.5-5 (-6) um, leves et incrassati, saepe septati; vesiculae brunneae, ampulliformes, 7.5-16 um diam. Aspergilla uni- serialia; phialides cylindricae, 5-7.5 x 2-2.5 um, in sum- 361 V2) (i ied + © — “= oO +r CO ee ae = (VO) Cane oO ees QU oO felt = ty (2) mM O 3} ie &<— as | K, 8 SET © SE a eae ~ =O ee ee (ej (Os t= wv & O WOO Vy oO Y We @) jy eu ORO = wn Cat © e Link ace == 63) = ‘- Mm iS “= GQ OrOr ©: nN Nn u << << 362 ma 1/2-2/3 vesicula insidentes. Conidia hyalina, globosa vel subglobosa vel parum ovoidea, 2-3(-4.5) x 2-2.5 um, levia. Holotypus SUM 3014, colonia exsiccata in cultura ex aqua decocta thea, Okinawa in Japonia, 22.vii.1991, iso- lata et ea in collectione fungorum Musei et Instituti Historiae Naturalis Chiba (CBM) conservata. Etymology: from Latin, primulinus=primrose, referring to thevcorvonveco tan: Colonies on Czapek agar growing rather restrictedly, attaining a diameter of 27 mm within 7 days at 25°C, somewhat floccose, more or less furrowed in a radial pat- tern, loose-textured, yellowish white (M. 4A2 after Kornerup and Wanscher, 1978); conidiogenesis sparse, in— conspicuous; reverse uncolored to pale yellow (M. 4A3) or Straw (Rayner, 1970). Colonies on CYA growing rapidly, attaining a diameter of 46 mm within 7 days at 25°C, or 85 mm within 7 days at 37°C, velvety, deeply furrowed, often zonate, consisting of a thin basal felt intermixed with numerous ascomata in a granular appearance at 25°C, yellowish white (M. 4A2) to pale yellow (M. 3A3-4A3) or Primrose (Rayner); conidiogenesis limited or profuse at 37°C, in the latter case greenish gray to brownish orange (M. 29D2-6C4) or Hazel (Rayner); reverse light yellow to reddish brown (M. 4A4-8E5) or Dark Brick (Rayner). Colo- nies on malt extract agar (MEA) spreading broadly, attain- ing a diameter of 54-56 mm within 7 days at 25°C, or 85 mm within 7 days at 37°C, floccose, plane or slightly zon- ate, thin, loose-textured, characterized by abundant asco- mata in granular appearance, pale yellow (M. 4A3) or Buff (Rayner); conidiogenesis moderate at 37°C but not suffi- ciently produced to influence the colony appearance; re- verse uncolored to pale yellow (M. 4A2-3) or Pale Luteous (Rayner). Ascomata non-ostiolate, superficial, scattered, chalky to buff, globose to subglobose, 100-250 um in diam, covered loosely with brownish encrusted hyphae, maturing within 21 days at 25°C; peridium 10-12.5 um thick, membra- naceous, textura epidermoidea, two-layered; outer layer composed of brownish, irregular-shaped, thick-walled cells measuring 5-/.5 wm wide; inner layer of hyaline, thin, Figure 2. Neosartorya primulina, SUM 3014. A. Ascosporesu( SEM) ox co00eambe Aseria oo: C. Conidia, x 1300. D, E. Aspergilla, x 1000 and x 1300, respectively. 363 364 Ascomatal initials developing as short branches of hyphae with large curled tips measuring ca. 12.5 um in diam. Asci 8-spored, produced in short chains, subglobose to more or less pyriform, 12-15(-17.5) x 11-13(-15) um, evanescent at maturity. Ascospores hyaline, lenticular, spore body 4, 5- 5(-5.5) x 3.8-5 um, provided with several narrow crests measuring ca. 0.5 um wide and with irregularly verrucose hemispheres. Conidial heads intermixed with ascomata, small, brownish gray, loosely radiate; conidiophores aris-— ing from the basal mycelium or sometimes from the aerial hyphae, stipes brownish, 200-600 x 2.5-5(-6) um, smooth and thick-walled, often several times septate; vesicles brownish, flask-shaped, 7.5-16 um in diam. Aspergilla uni- seriate, phialides cylindric, 5-/.5 x 2-2.5 um, covering: the upper half to two-thirds of the vesicle. Conidia hya- line, globose to subglobose or slightly ovoidal, 2-3(-4.5) x 2-2.5 um, smooth-walled. Maximum growth temperature: 47°C. Pasteurization» atyee.C for 15 min. 1s effective. Specimen examined: SUM 3014 (holotype), a dried cul- ture of an isolate from a canned oolong tea beverage, Oki- nawa-Pref., Japan, 22.vii.1991. The specimen studied and living culture derived from the type are preserved in the Natural History Museum and Institute, Chiba (CBM), Japan. Neosartorya primulina can be readily recognized by its chalky-buff colored ascomata, short-stipitate initials of the ascomata, and morphology of the ascospores. The scann- ing electron micrograph (SEM) of the ascospores provided a much clearer differentiation between N. primulina and all other members of Neosartorya (Samson et al., 1990, 1992); the ascospore ornamentation in this species is character- ized by several narrow crests in the equatorial area and irregularly verrucose hemispheres (Fig. 2). Neosartorya hiratsukae somewhat resembles this species in the restrict- ed growth on Czapek agar and in the formation of pale yel- low ascomata, but differs in having ascospores with convex walls bearing numerous closely anastomosing ridges arrang- ed in a fine reticulate ornamentation(Udagawa et al.,1991). Acknowledgement: We thank Prof. D. Malloch of the Univer- sity of Toronto for reviewing this paper. Literature cited Beuchat, L. R. 1986. Extraordinary heat resistance of Talaromyces flavus and Neosartorya fischeri asco- SHOP esmenicru le prROGUCUSiwm Us Food! SGT, canine LoUG- 15708 365 Conner, D. E. and Beuchat, L. R. 1987. Heat resistance of ascospores of Neosartorya fischeri as affected by sporulation and heating medium. Intl. J. Food Microbiol., 4: 303-312. Gonezeaen Mae DUStagereure sander lug, imu.) 1989s Sef fect of the post-dry heat treatment temperature on the recovery of ascospores of Neosartorya fischeri. Lett. Appl. Microbiol.,.8: 59-62. Kavanagh, J., Larchet, N. and Stuart, M. 1963. Occurrence of a heat resistant species of Aspergillus in canned strawberries. Nature (Lond. ), 198: 1322. Kornerup, A. and Wanscher, J. H. 1978. Methuen Handbook of Colour. 3rd ed. Eyre Methuen, London. 252p. Kozakiewicz, Z. 1989. Aspergillus species on stored pro- ducts. CMI Mycological Papers No. 161: 1-188. Malloch, D. and Cain, R. F. 1972. The Trichocomataceae: Ascomycetes with Aspergillus, Paecilomyces, and Peni- cillium imperfect states. Can. J. Bot., 50: 2613- cocae i ciliscntmeny Seubeuchaues ark mranGeaprisvVadscds"U, 919097 Growth and fumitremorgin production by Neosartorya fischeri as affected by food preservatives and organic acids. J. Appl. Bacteriol., 66: 197-207. Peterson, S. W. 1992. Neosartorya pseudofischeri sp. nov. and its relationship to other species in Aspergiilus section Fumigati. Mycol. Res., 96: 547-554, Rayner, R. W. 1970. A Mycological Colour Chart. Common- wealth Mycological Institute, Kew and the British Mycological Society. Samson, R. A. 1989. Filamentous fungi in food and feed. Je App ls Bacteriol). Symp. supp ls .t/07s 275-355. Samson, R. A., Nielsen, P. V. and Frisvad, J. C. 1990. The genus Neosartorya: Differentiation by scanning elect- ron microscopy and mycotoxin profiles. In: Modern Concepts in Penicillium and Aspergillus Classifica- Ci OncO mrSamscOnmenk een emander tbCend sete ee 405-401, Plenum Press, New York. SANSOM mane MA em HOCKING ee AU sence Camu eau, MONGEN UNC aiee ns (ed. ) 1992. Modern Methods in Food Mycology. Elsevier, Amsterdam. 388p. Scott, V. N. and Bernard, D. T. 1987. Heat resistance of Talaromyces flavus and Neosartorya fischeri isolated from commercial fruit juices. J. Food Prot., 50: 18- (A>. Solittstoesser, 0: Fovand Churey,- J. JU, .1989." Effect of low concentration of sorbic acid on the heat resist- ance and viable recovery of Neosartorya fischeri asco- 366 spores. J. Food Prot., 52: 821-822. Udagawa, S., Tsubouchi, H. and Horie, Y. 1991. Neosarto- rya hiratsukae, a new species of food-borne Asco- mycetes. Trans. Mycol. Soce-Wapany. ocimes-49. MY COTAXON Volume XLVI, pp. 367-377 April-June 1993 STUDIES ON BOLETUS SECTION LURIDI Re. TREU Virginia Polytechnic Institute and State University, Department of Biology, Blacksburg, Virginia, 24061 SUMMARY Four species of Boletus section Luridi described by Smith and Thiers have been studied: Boletus pseudo-olivaceus, B. roseobadius, B. rufocinnamomeus, and B. subluridellus. B. pseudo-olivaceus can be easily differentiated from the other species by its weakly but distinctly amyloid trama in stipe and pileus. There are no major microscopical differ- ences between B. subluridellus, B. rufocinnamomeus, and B. roseobadius. INTRODUCTION The section Luridi Fr. is a relatively well defined group in the genus Boletus Fr. Singer (1986) characterized the section by its Boletus habit, the narrow pores, which are often discolorous and red, and by the occasional presence of toxins in the fruitbodies. The sporocarps turn immediately blue on bruising or cutting. Smith and Thiers (1971), who interpreted the group as a subsection within the section Boletus, have increased our knowledge of the group considerably; they also described several new species from Michigan. A recently found collection from Virginia revealed however, that species delimitation within this group is still problematic; a number of characters considered to be important for species delimitation showed variation within the collection. Smith and Thiers (1971) described a "Spatter pattern" for the distribution of different characters within this group and suspected a gene exchange between some taxa, such as B. subvelutipes Peck and B. pseudo-olivaceus Smith and Thiers, two species growing in the same habitat. Therefore, a reevaluation of characters seems to be necessary. As a first step the collection from Virginia and specimens of four different species which are closely related have been studied here. 368 MATERIALS AND METHODS Microscopical observations were made in KOH (10%) unless stated otherwise, but the cuticle was observed in Melzer's reagent. A drawing tube was used for the illustra- tions. At least 15 spores per collection were measured and the mean dimensions D" for the spores were calculated; the length/width ratio (Q) was obtained for each of the spore measurements and a mean value (Q,) was calculated for each of the species. Detailed descriptions of the macroscopic characters of each taxon were presented by Smith and Thiers (1971). Color descriptions follow Kornerup & Wanscher (1967). Official colors are in quotes. The terminology of cystidia has been adopted from Singer (1986) and Wolfe (1991). RESULTS BoLeeus wR 316 Figs 1521-25 Pileus 4-14 cm, convex, surface tomentose to almost velvety, »color.darks"olive’ brown" (tou"raw umber"). "soot brown", or "coffee", (4 F4-8; 5 F5-8), a few fruitbodies with rose-red areas. Tubes adnexed, up to 1 cm thick, yellow, but rapidly staining blue when cut. Pores round, small, “lake red", "Pompeian red", "dark red" until old age (9 C7-8, 10 C7-8), staining greenish blue when injured. Spore print "bronze", "mustard brown", "linoleum brown" (5 E5-7). Stipe clavate or equal, solid, 2.5-9 cm long, 1.5-3.5 cm thick, orange-yellow, dark red toward the base, bruising instantly blue. In some specimens a fine red reticulation was observed on the very tip of the stipe. Context of the whole fruitbody after cutting instantly changing to dark blue. Chemical reactions of (blued) pileus context: KOH orange; NH,OH yellow, FeSO, yellow. Chemical reactions on pileus supiace:s KOH (—)/7)NH,OH i>) FeSO .a(—)e Spores 10:8-13.5) x 3%6-5/4 pm; D" = 12.1% 4.4. pm780 = 92.353 537 0 J= 2.8 ;etusoid,, yellow to, brownishsyellowiin KOH; spores from deposit 11.7-14.4 x 3.6-5 um, pale yellow- brown in KOH; spores in Melzer's reagent 10.4-12.6 x 3.6- 4.5 um, occasionally thick-walled, pale yellow-brown. Basidia 26-41 x 8-10 um, 4-spored, occasionally 2-spored, no basal clamps, hyaline in KOH and Melzer's reagent, a few with a yellow, plasmatic pigment. Pleurocystidia 24-52 x 4.7-8 wm, fusiform. Pseudocystidia 19-31 x 3.7-5.8 wun, fusiform. All septa without clamps. Cuticle a trichoderm or occasionally a cutis, 180-190 um thick; trichodermial end cells connected at their tips and forming fascicles. Trichodermial hyphae 3.5-4.1 pm 369 thick, with deep red walls and occasionally brown contents in Melzer's reagent; yellowish, pigment plasmatic, some- times with granular contents in KOH. Context below tricho- derm a tangle of hyaline hyphae, 3.5-5.9 wm thick, with granular, yellow-brown debris between hyphae; occasionally some hyphae have peculiar wall thickenings up to 1.5 um. Oleiferous hyphae can be found occasionally (diam. 5 pum). Stipe dermatocystidia of two types: first type 21-26 x 4.2-7.4 um clavate, similar to probasidia. Second type (less frequent) 19-110 x 4.1-8 wm, fusiform. Scattered basidia can be found all along the stipe cortex, 23-35 x 8- 10 um, 2- or 4-spored, with yellow content in KOH, yellow- brown to dark brown in Melzer's reagent. Tramal hyphae inamyloid. Comments. This collection seems to be close to B. subluridellus although the Q value for the spores is some- what higher for this collection (2.8 here versus 2.6 in the holotype of B. subluridellus). A number of characters shows high variation within the collection: The long and setiform caulocystidia which were regarded by Smith and Thiers (1971) as an important character to delimit the group around B. subluridellus could be found only sporadically in some but not all fruitbodies of this collection. Therefore their presence seems to vary within at least this species. The structure of the cuticle is also variable: usually it consists of conical fascicles of trichodermial hyphae but in the same collection it can be a cutis. Collection examined: Virginia, Montgomery Co., near Pandapas Pond, under oak and white pine, 9 Aug 1991, R. Treu (TR 1316), Massey Herbarium (VPI). Boletus pseudo-olivaceus Smith & Thiers. 1971. The boletes OLPMa Chigan awe 5s: Firgss [=5 Spores 12.2-15.3 x 5.4-6.3" pm; DY = '141.%'°5.9 pm?'OQ =2 3726.07, Or = 2.45) fusoid, often thick-walled}(0.5=-1 um), in KOH yellow; spores in Melzer's reagent 9.9-15.3 x 5-5.9 um, thick-walled (0.5-1 wm), pale brownish yellow. In Melzer's reagent many spores show a minute, indistinct germ pore. Basidia 31-39 x 9-13 wm, four-spored or two-spored, hyaline or yellow. Pleurocystidia 37-41 x 6-7 um, fusiform. Pseudocystidia 26-29 x 6-7 um, clavate, with golden yellow pigmentation (plasmatic). All septa without clamps. Cuticle a cutis, 350-450 um thick, with many oleifer- ous hyphae; hyphae 2.3-6.3 um thick, hyphal walls not red in Melzer's reagent; oleiferous hyphae with yellow, homoge- neous content. Stipe dermatocystidia 29-63 x 12-14 um, fusiform or 370 -etptysAoojeurep edtas g ‘etptysAooiznetd 7 ‘eTptsed ~ ‘sezods T ‘*“SneoeATTO-opnes “w7#O0T = saeq eTeos -etptysAoopnesd € SnjeTOd G-T °*shTa 371 clavate, yellow or brownish yellow. Occasionally yellow caulobasidia can be found among the cystidia. Tramal hyphae of stipe as well as tramal hyphae of pileus slightly amyloid, purple to grayish in Melzer's reagent. Yellow- orange tomentum at the base of the stipe consisting of bundles of parallel hyphae, 1.8-3.6 wm thick, yellowish in KOH, brownish and some hyphae with dark brown contents in Melzer's reagent. Comments. B. pseudo-olivaceus is easily distinguished from the other species by a weak, but distinct amyloid reaction of the hyphae in both the pileus and the stipe trama. This reaction is persistent and should not be confused with the "fleeting-amyloid" reaction described by Smith and Thiers (1971) for some boletes. The spores of B. pseudo-olivaceus also have a lower Q, value (2.4) than the other species (2.6-2.7). Collections examined: Michigan, Emmet Co., Mackinaw City Hardwoods, scat- tered at edge of hardwood forest, 12 Jul 1967, A.H. Smith 74510*- holotype, MICH. Michigan; Charlevoix) Co.) Eastidordan; ,12%Jul) 1967, James Bennet (A.H. Smith 74511), MICH. Boletus roseobadius Smith & Thiers. 1971. The boletes of Michigan. Ps )354. Figs. 6-10 Spores 13.1-16.2 x 4.5-6.3 pm; DY = 14.5 x 5.5 pm; Q = 2.4-3; Q = 2.7; fusoid, sometimes thick-walled, yellow- ish-brown to yellow in KOH; spores in Melzer's reagent 12.2-16.7 x 4.5-6.3 wm, yellowish-brown to brown. Spores on the stipe cortex sometimes dextrinoid and broader than hymenial spores. Basidia 25-34 x 8-12 wm, four-spored or two-spored, hyaline or sometimes with yellow, homogenous or granular, plasmatic pigment. Pleurocystidia 24-62 x 6-10 um, fusiform, mostly with apical neck. Pseudocystidia 21-34 x 5-6 um, fusiform or clavate, with golden yellow pigmenta- tion (plasmatic), along the tube edges; occasionally a few pseudocystidia with size and shape of the pleurocystidia, but with yellow pigment, on the tube walls. All septa without clamps. Cuticle a cutis, 130-230 wm thick; hyphae 2.7-5.4 um thick, hyphal walls red in Melzer's reagent; some hyphae with yellow, granular content. Stipe dermatocystidia 24-80 x 5-11 wm, fusiform or cylindric sometimes with an apical neck, hyaline or brown. Occasionally 3- or 4-spored brown basidia can be found among the cystidia. Tramal hyphae inamyloid, in stipe base sometimes with red walls. Comments. It is almost impossible to distinguish this 372 10 i Pre Figs. 6-10 Boletus roseobadius. 6 Spores. 7 Basidia 8 Pleurocystidia. 9 Pseudocystidia. 10 Stipe dermatocystidia. Figs. 11-15 Boletus rufocinnamomeus. 11 Spores. 12 Basidia 13. Pleurocystidia. 14 Pseudocystidia. 15 Stipe dermato- cystidia. Scale bars = 10ysm. 373 species from B. subluridellus on microscopic characters. A very distinctive feature of these species is the deep red color of the walls of many cuticular hyphae in Melzer's reagent. Smith and Thiers (1971) characterized the species by the size of the pleurocystidia, the red (in Melzer's reagent) cuticular hyphae, and the brown cortex elements (in KOH). However, there is no differences in the size of the pleurocystidia (24-62 um in B. roseobadius, 34-50 um in B. subluridellus). Smith and Thiers (1971) described B. roseobadius as having a whitish or pallid flesh in compari- son the bright yellow flesh in B. subluridellus. The interpretation of this feature might be difficult because of the instant color change of the flesh to deep blue. Collections examined: Michigan, Washtenaw Co., solitary under low hardwoods, 20 Sep 1965, F. V. Hoseney (A.H. Smith 72668 - holo- type), MICH. Michigan, Washtenaw Co., Pinckney State Recreation Areas) 258 Auge 9 ish A. HeSmatn $8453:-7:, IMiCH: Boletus rufocinnamomeus Smith & Thiers. 1971. The boletes OTeMVchiganiiPses5 Gy Figss214-15 Spores 13.2-16.2 x 5.1-6.3 pm; DY = 14.4 x 5.4 pm; Q =e 4-319; 0.) = 92,75) Lusoid; amostily, thin-walled,, pale’to golden yellow; spores in Melzer's reagent 12.6-17.1 x 4.5- 6.3 um, yellowish brown. Basidia 30-34 x 10-12 um, four- spored, occasionally two-spored, hyaline. Pleurocystidia 24-40 x 5-9 um, fusiform with apical neck. Pseudocystidia 25-34 x 4-6 pum, fusiform, sometimes with a neck, with golden yellow pigmentation (plasmatic), along the tube edges, sometimes a few on the tube walls. All septa without clamps. Cuticle a trichodermium, 180-200 um thick, formed of conical fascicles of trichodermial hyphae with brown, often granular content in Melzer's reagent; occasionally some of the hyphae below the fascicles with weakly red walls in Melzer's reagent. Hyphae forming the fascicles 1.8-3.6 um thick, hyphae below 2.7-6.3 um. Stipe dermatocystidia 24-100 x 4-14 um, fusiform, with or without a neck, occasionally with a secondary septum, hyaline or golden yellow or brownish. Tramal hyphae inamyloid. Collections examined: Michigan, Cheboygan Co., Burt Lake, hardwoods, 11 Aug 1969, A.H. Smith 77831 - holotype, MICH. Michigan, Cheboygan Co., Burt Lake, Colonial Point Hardwoods, 7 Aug 1969, A.H. Smith 77746, MICH. 374 Boletus subluridellus Smith & Thiers. 1971. The boletes of Michigan. P. 349. Figs. 16-20 Spores 10.6-15.3 x 4.5-5.4 um; DY = 12.8 x 4.9 um; Q = 2.2-2.9; Q. = 2.6; fusoid, occasionally slightly thick- walled, brownish yellow in KOH; spores in Melzer's reagent 11.7-14.4 x 4.5-5.9 um, yellow-brown. Spores on stipe cortex frequently with golden yellow pigmentation. Basidia 22-36 x 11-12 um, 4-spored, hyaline, yellow-brown or brown. Pleurocystidia 34-50 x 6-8 um, fusiform with apical neck or clavate-mucronate. Pseudocystidia 32-33 x 4-6 um, fusiform and sometimes with apical neck or clavate, with golden yellow pigmentation (plasmatic), along the tube edges; occasionally a few pseudocystidia with size and shape of the pleurocystidia, but with yellow pigment, on the tube walls. All septa without clamps. Cuticle a trichodermium formed of conical fascicles of hyphae; thickness of cuticle 180 um; diameter of tricho- dermial hyphae 3.5-5 um, hyphal walls deep red in Melzer's reagent. Stipe dermatocystidia variable, fusiform, with or without a neck, or broadly clavate-mucronate; fusiform cystidia 26-66(89) x (3)5-13 um; clavate-mucronate cystidia 30-35 x 14-18 wm; hyaline or golden yellow in KOH, brown in Melzer's reagent. Tramal hyphae inamyloid. Comments: see B. roseobadius Collection examined: Michigan, Washtenaw Co., Ypsilanti Golf Course, Ypsilanti, in grassy oak woods, 3 Sep 1961, A.H. Smith 64046 - holotype, MICH. DISCUSSION There is not much variation in the spore size of the different taxa studied here. However, the Q and Q values are increasingly used as a taxonomic character (Breitenbach and Kranzlin 1991, Wolfe 1991). B. pseudo-olivaceus with a Q, of 2.4 differs from the other species with a Q, between 2eGoand) 258% The hymenial cystidia range in shape from usually fusiform to almost clavate. Fusiform cystidia often have an apical neck. Following the terms used by Singer (1986) and Wolfe (1991) for hymenial cystidia, most species have fusiform or clavate, yellow pseudocystidia along the tube edges and hyaline pleurocystidia located on the tube walls and larger in size. Occasionally a few pseudocystidia can be found on the tube wall, and they differ from the pleuro- cystidia only in their yellow pigment. In contrast to the genus Tylopilus Karsten, where hymenial cystidia play an 375 Figs. 16-20 Boletus subluridellus. 16 Spores. 17 Basidia. 18 Pseudocystidia. 19 Pleurocystidia. 20 Stipe dermato- cystidia. Figs. 21-25 Boletus (TR 1316). 21 Spores. 22 Basidia. 23 Pleurocystidia. 24 Pseudocystidia. 25 Stipe dermatocystidia. Scale bars = 10m. 376 important role for the delimitation of species (e.g. Wolfe 1986, Wolfe 1991), the cystidia in the section Luridi do not seem to be important taxonomic characters. Stipe dermatocystidia have been used by Smith and Thiers (1971) to delimit species in the section Luridi (B. subluridellus, B. roseobadius, and B. rufocinnamomeus with long, often setiform caulocystidia). Measurements taken here show however, that they can range widely within one collection and are often difficult to find, especially in small collections. Some sporocarps of collection TR 1316 had very long stipe dermatocystidia but these were scat- tered, difficult to detect and only present in some fruitbodies of the collection. The shape of the stipe dermatocystidia ranges from clavate to rostrate to fusiform and their contents often stain brown in Melzer's reagent. Basidia do not show variation between the species. They are usually hyaline, but sometimes also pigmented. Their basal septum is simple, without clamps. The hymenophoral trama in all the species studied is divergent, sometimes only slightly so, with a central strand of parallel hyphae which stains brownish in Melzer's reagent. The cuticle can range from a trichoderm to a cutis. Larger collections (TR 1316) show that there is variation within species. The content of cuticle hyphae of B. rufocinnamomeus turns brownish in Melzer's reagent. The same character could occasionally be observed in the colrectiuon. TRI 2a In most cases the tips of trichodermial hyphae are glued together to form conical fascicles. A very conspicuous character of both B. subluridellus and B. roseobadius are the cuticular hyphae many of which have deep red walls in Melzer's reagent. However, this constitutes no dextrinoidity, because the pigment can be observed in water mounts as well. It can not be detected in KOH, where it apparently changes its color or dissolves. Rarely those pigmented hyphae could also be detected in the cuticle of B. rufocinnamomeus. The pigment is often indicated macroscopically by a red color of parts of the fruitbody (e.g. pileus, stipe trama). A peculiar character of B. pseudo-olivaceus is the amyloid trama of stipe and pileus. Although very distinc- tive, the intensity of the staining is not very strong and quite homogeneous throughout the trama. A stronger amyloid reaction in the genus Boletus can be found in cross walls of tramal hyphae of Boletus calopus Fr. (Miller and Watling 1968) and in B. piedmontensis Grand and Smith in the form of amyloid particles in the cuticular hyphae (Grand and Smith 1971). .According “to “Smith and =Thiers, (1971)'«B. pseudo-olivaceus is closely related to B. queletii Schulz. from Europe, a view supported by Singer's (1977) observa- tion of distinctly amyloid hyphae in the stipe of B. queletii. None of the species shows clamps in any parts of the O77, fruitbodies. Themcollectson pik i316, wwhich has’ been included in this study, seems to be close to or identical with B. subluridellus. The presence of many red-walled hyphae in the cutis and the spore measurements indicate a high Similarity between the material from Virginia and collec- tions of B. subluridellus. At the same time however, many characters, like the pigmentation of the cuticular hyphae, seem to vary even within collections of one species and may depend on the age of sporocarps. Larger collections may reveal the variability of some of the characters which are traditionally used. Therefore a further study of a larger number of other taxa in this group is obligatory and would necessarily include the European taxa of this group. ACKNOWLEDGEMENTS I wish to thank Dr. C.B. Wolfe, Pennsylvania State University, for helpful comments and a critical revision of SHewmanuscCh pl uiMmeIndebGeneto Dre O.KeeMiller, 0%. for hasenospitality ands lab! space. =f thank Dr. R.L. Shaffer, curator of the Herbarium of the University of Michigan, for providing loans. The "Deutsche Forschungsgemeinschaft" (DFG) is acknowledged for providing a research grant. LITERATURE Breitenbach, J. and F. Kranzlin. 1991. Fungi of Switzer- land. Vol. 3. Boletes and agarics. lst part. Edition Mykologia, Lucerne, Switzerland. Grand, L.F. and A.H. Smith. 1971. A new species of Boletus, section Luridi, from North Carolina. Mycologia 63: 884-888 Kornerup, A. and J.H. Wanscher. 1967. Methuen Handbook of Colour. Methuen and Co. Ltd., London, Great Britain. Mebier, O-K., Jr.)and oR. | Watlinga FPl968.. The status . of Boletus calopus Fr. in North America. Notes from the Royal Botanic Garden Edinburgh 28: 317-325. Singer, R. 1977. Keys for the identification of the species of Agaricales I. Sydowia 30: 192-279. Singer, R. 1986. The Agaricales in modern taxonomy. 4th ed. Koeltz Scientific Books, Koenigstein, Germany. Smith, A.H. and H.D. Thiers. 1971. The boletes of Michigan. The University of Michigan Press, Ann Arbor, Michigan. WOPlepeC=b. wi Loloashype studies tin Tylopilus-vltl Taxa described by Walter H. Snell, Esther A. Dick, and co- workers. Mycologias/s 2: 22-317 Wolfe, C.B., Jr. 1991. Type studies in Tylopilus (Boleta- ceae). V. Taxa described by Alexander H. Smith, Harry D. Thiers and Samuel J. Mazzer. Can. J. Bot. 69: 1833- LOS. Or , é ? Wek ae , wn ia a cm ‘ ~ a | Pa) z ae “at At 6» aot nee i A $ ae BS a a ad “-t, bed Bah 7" 7c: aa ¢ is Oe Sa eet iain Ad . Ne ‘sf ns ; \ ate ‘e ive sD dares: ee yt ie x bY Poker ct» Be ee eee See ae, Pr. ns ned Ley Peas | ata eee CLL eae roe i ‘it ie ine My ha } ‘ t a . si is Pd oe det tak va ‘ he 22 MYCOTAXON Volume XLVI, pp. 379-385 April-June 1993 TWO NEW CRINIPELLINAE (TRICHOLOMATACEAE: MARASMIAE) FROM SOUTH AMERICA Roy E. HALLING Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126 ABSTRACT Crinipellis ticoi from lowland Bolivia and Chaetocalathus magnus from Andean Colombia are newly described from the Neotropics. Both taxa possess dextrinoid, long, thick-walled, setoid hairs on the pileus that distinguish these lamellate genera in the Crinipellinae. Key Words: Agaricales, Crinipellinae, Crinipellis, Chaetocalathus, Bolivia, Colombia Study of specimens gathered in South America over a four year period (1986- 1990) has resulted in the discovery of many new taxa of agarics and boletes (eg., Halling & Ovrebo, 1987; Halling 1989a, 1989b, Singer et al., 1990; Horak & Hall- ing, 1991; Tulloss et al., 1992; Baroni & Halling, 1992; Halling, 1992). Further ob- servations on additional material has yielded a previously unknown species of Crinipellis with bright orange pigmentation and the largest species of Chaetocala- thus known to date. Fifty years ago, Singer (1942) provided a worldwide mono- graph of these two genera, and more recently, published an updated monograph for the Neotropical taxa (Singer, 1976). Two additional Neotropical species (one from each genus) have since been recognized (Singer, 1989). Color designations in parentheses (e.g., SA6) are taken from Kornerup and Wanscher (1983). Other color terms (e.g. brick red) are general approximations of those designations. Letter abbreviations describing spore sizes are: n = number of spores measured, x = mean length x mean width, Q = mean length/width ratio. Crinipellis ticoi Halling, sp. nov. Figs. 1-4 TYPE: BOLIVIA. Dpto. Beni: Prov. Iturralde, S of Rurrenabaque, Rio Tuichi near junction with Rio Beni, “Laguna del Tigre,” +67°30’W, 14°25’S, 14 Apr 1990, Hailing 6433 (Holotype: LPB; Isotype: NY). Pileus aurantius sulcatus tomentosus vel subtomentosus, epilosus. Lamellae adnexae subdistantes crassae latae aurantiae. Stipes centralis strictus +aequalis tenax tortilis pubescens vel fibrillosus vel 380 fibrilloso-pruinosus, pubescentis aurantiis, superfacies flava. Basidiosporae in cumulo albae, 12.1-14.3 x 5-7.1 um, inamyloideae. Hymenium pigmento aurantio in solutionibus alkalicis dissolventibus. Pleurocystidia nulla. Cheilocystidia dispersa, hyalina, parietibus tenuibus et glabris. Setae pileipellis sparsae dextrinoideae, parietibus crassis, interdum 2-3 septatae, apicibus obtusis vel subacutis vel sub- capitatis. Setae stipitipellis abundantes dextrinoideae, parietibus tenuibus vel crassis, apicibus obtusis, aliter setae pileipellis similis. Fibulae praesentes. Etymology: In honor of Sr. Abellardo “Tico” Tudela R., of Rurrenabaque, whose logistical support aided in the discovery of this species. Pileus 1—3(-—4) cm broad, convex to plane, dry or moist and greasy then fading when drying but not truly hygrophanous, orange (7A8—7B8) fading to a paler orange (6A7-6B7 to 6A8-6B8 to near 5A6), sulcate to subsulcate to the disc, even on disc, tomentose to matted subtomentose on disc, faintly so toward margin, more yellowish on ridges with age and remaining orange in sulcae; margin even. Flesh pale orange (5A2) in pileus, pale yellowish in stipe apex and below, up to 2 mm thick. Odor mild. Taste styptic but not unpleasant. Lamellae adnexed to sub- free, subdistant, thick and broad (up to 7 mm), orange (7A8 to 6A8) but slightly paler toward the even, nonmarginate edges; with lamellulae in 1-2 tiers. Stipe (1—)2.5-7 cm long, (1—)3-—4 mm broad, insititious, strict, central, twisted, tough, +equal but often broadest at apex and base, sometimes with a bulb at base; sur- face pubescent to fibrillose to fibrillose-pruinose overall, denser toward the base, with orange (color range as cited above) fibrils on yellowish ground above and be- low when young, brown toward base with age, sometimes cream-colored slightly at base; interior stuffed, not hollow. Fig. 1. Crinipellis ticoi. (Halling 6393). x1. 381 5 um Figs. 2-4. Microscopic features of Crinipellis ticoi (Halling 6433). 2. Basidiospores. 3. Cheilocystidia. 4. Setae from pileipellis. Basidiospores white in deposit, (10—)12.1—14.3 x 5—7.1 wm (n = 20, x = 13 x 5.8 um, Q = 2.3), ellipsoid to suballantoid and inequilateral in profile, ellipsoid to subcylindric in abaxial view, inamyloid and acyanophilous, smooth, thin-walled. Hymenium embedded in an amorphous golden orange pigment that dissolves in alkali. Basidia 28-35 x 7-8.4 um, clavate, 4-sterigmate, not siderophilous. Pleurocystidia absent. Cheilocystidia scattered, inconspicuous, hyaline, thin-walled, smooth, subfusoid to subcylindric, 38-49 x 6-8 wm. Lamellar trama interwoven, | inamyloid; hyphae smooth, thin-walled, 3.5-12.6 um diam. Pileus trama radially oriented, inamyloid; hyphae smooth, thin-walled, S—10(-14) um diam. Setae sparsely distributed on pileus surface especially toward the margin, more abundant on the disc, 70-210 x 5—9 um, thick-walled, dextrinoid, occasionally 2—3 septate, with obtuse to subacute to subcapitate apices, arising from a pileipellis of radially oriented, repent, thin-walled, hyaline hyphae; cells 7—14(-17.5) um in diam, sometimes lightly encrusted and slightly inflated. Setae abundant on stipe surface, Strict or curved, dextrinoid, thin- to thick-walled, with obtuse apices, otherwise resembling setae in pileipellis, arising from a stipitipellis whose elements are verti- cally oriented, dextrinoid, smooth, and thin-walled, 3.5-—6 um diam. Clamp con- nections present. On rotten wood in lowland Amazonian rainforest. 382 Additional material examined: BOLIVIA. Dpto. Beni: Prov. Iturralde, S of Rurrenabaque, along Rio Beni between town of Rurrenabaque and junction of Rio Tuichi, +67°30’W, 14°25’S, 4 Apr 1990, Halling 6393 (LPB, NY). Prov. Gral. José Ballivian, +1 km E of Rurrenabaque, along road to air- port, +67°30’W, 14°20’S, 29 Mar 1990, Halling 6372 (LPB, NY); S of Rurrenabaque, along Rio Beni between town of Rurrenabaque and junction of Rio Tuichi, +67°30’W, 14°25’S, 3 Apr 1990, Halling 6383 (LPB, NY), 7 Apr 1990, Halling 6416 (LPB, NY). Crinipellis ticoi seems best accommodated in sect. Jopodinae (Singer, 1976) characterized by the brightly pigmented pileus and tissues not greening or graying in alkali. In this section, C. insignis Sing., C. purpurea Sing., and C. hygrocybioides (Henn.) Sing. appear most similar. Crinipellis insignis differs in the purple to pur- plish pink pileus, white lamellae, and short, eccentric stipe. Crinipellis purpurea is distinguished by the dark purple red pileus, white lamellae, and smaller spores (6.5-10.3 x 3-4.7 um). Also, both of these species are described as having abundant setae giving the pileus a pilose aspect. Crinipellis ticoi has a tomentose pileus with scattered setae. Comparison of C. ticoi to C. hygrocybioides is more problematical. When Singer (1989) transferred Marasmius hygrocybioides Henn. to Crinipellis, he did not provide further data or documentation other than to indicate that the type is a species of sect. Ionopodinae [sic]. Henning’s (1902) original des- cription tersely describes a yellow orange, small, marasmioid fungus. Two isotypes (Kamerun. Bipindi, Mai 1899, G. Zenker 2039, PC!; K!) have been compared. The Kew material consists of one basidiome which was not examined microscopically. The Paris specimen contains some 14 complete basidomata and additional stipes lacking pilei. Abundant dextrinoid setae are present on the pileus and stipe in- dicating Crinipellis, but spores and cystidia were not observed. Contrary to C. ticoi, C. hygrocybioides is a smaller fungus (pileus 6-11 mm broad, stipe 10-20 x 0.5-0.8 mm) and has an umbilicate to papillate pileus which is pilose at the margin. Chaetocalathus magnus Halling, sp. nov. Figs. 5-8 TYPE: COLOMBIA. Dpto. Antioquia: Mpio. Guarne, near Centro Experimental Piedras Blancas, 14 km E of Medellin, 75°29’W, 6°15’N, 2350 m, 28 Nov 1988, Halling 6159 (Holotype: HUA; Isotype: NY). Basidiomata estipitata. Pileus siccus pigmento brunneo, primo planus et minime granulosus vel velutinus tum rugulosus vel subsulcatus. Lamellae primo confertae et albae tum subdistantes et aurantio-albae. Basdiosporae 9.3-12.1 x 6.4-8.6 um, dextrinoideae et cyanophilae. Pleurocystidia rarissima, inamyloidea, aliter cheilocystidia simili. Cheilocystidia inferme dextrinoidea, nonincrustata, parietibus crassis vel tenuibus, appendiculis apicalibus. Setae pileipellis dextrinoideae, parietibus crassis, apicibus obtusis. Fibulae praesentes. Etymology: Latin magnus, great or large. Basidiomata nonstipitate. Pileus (0.7-)1-—2.2(-—2.7) cm broad, pendant- ungulate to barely laterally attached, dry, even and matted velutinous to minutely granulose and burnt sienna to brick red (7D8-—7) when young, soon rugulose to subsulcate and fading to near grayish orange (6BS5), white to tan near attachment point. Flesh thin, pallid. Odor and taste mild. Lamellae close and white when young with unequal widths and barely subfimbriate edges, soon subdistant and orange white, afimbriate, subintervenose. 383 Fig. 5. Chaetocalathus magnus (Halling 6159). x2. Figs. 6-8. Microscopic features of Chaetocalathus magnus (Halling 6159). 6. Basidiospores. 7. Cheilocystidia. 8. Setae from pileipellis. 384 Basidiospores 9.3-12.1 x 6.4-8.6 um (n = 20,x = 10.4 x 7 um, Q = 1.5) broadly ellipsoid, sometimes subamygdaliform, weakly dextrinoid when observed- singly, more strongly so in mass, cyanophilous, hyaline in KOH, smooth, thin- wailed, with a slightly thickened wall in age, occasionally transversely septate. Basidia 28-35 x 8-9.5 um, clavate, hyaline, thin-walled, not siderophilous, 4- sterigmate, Pleurocystidia inconspicuous and extremely rare, thick-walled, in- amyloid, otherwise resembling cheilocystidia. Cheilocystidia 25-46 x 10-15 um, thin-walled or thick-walled, with 2—4 apical appendages, typically subcylindric, not encrusted, usually inamyloid or very rarely weakly dextrinoid, and then the reac- tion extremely localized on a given cystidium. Lamellar and pileal trama inter- woven, hyaline, inamyloid, not gelatinous; hyphae 5-7 um broad, filamentous, thick-walled (up to 3 wm). Pileipellis an interwoven layer of repent, hyaline, in- amyloid hyphae, 4—8 um in diam, lightly encrusted with pigment, thin-walled to progressively thicker-walled as they give rise to setoid hairs; setae 100-200 um x 3.5-7 ym, thick-walled, sometimes entirely lacking a lumen, dextrinoid, smooth, highly refractive to pale golden in KOH, with obtuse apices. Clamp connections present. On stems of Vaccinium (Ericaceae) in Quercus humboldtii forest. Additional material examined: COLOMBIA. Dpto. Antioquia: Mpio. Guarne, near Centro Ex- perimental Piedras Blancas, 14 km E of Medellin, 75°29’W, 6°15’N, 2350 m, 10 Nov 1988, Desjardin 4763 (HUA, SFSU). This is the eighth species of Chaetocalathus known from the Neotropics, and the second known from Colombia [after the widespread Ch. liliputianus (Mont.) Sing.]. Chaetocalathus magnus fits best in sect. Holocystis Sing. by virtue of the dextrinoid (rarely), thick-walled, apically appendaged cystidia. However, it can be distinguished easily by its large size, brown pigmentation of the non-pilose pileus, non-encrusted cystidia, and lack of entire and simple pleurocystidia. However, when pleurocystidia are present, they are extremely rare and resemble the in- amyloid cheilocystidia. ACKNOWLEDGMENTS Field work in Bolivia and Colombia was supported by National Science Foun- dation Grant #BSR — 8600424. I am indebted to the staff and students at Univer- sidad de Antioquia (HUA) for aid and guidance in Colombia; for assistance in Bolivia, I thank Emilia Garcia (LPB), the late David Smith (MO), David and Sandy Williams (USDA), and Sr. Abellardo “Tico” Tudela R. in Rurrenabaque. Gregory Mueller (F) is thanked for his review of this paper. Further, I am grateful to Dennis E. Desjardin (SFSU) and Ana E. Franco-M. (NY) who provided out- standing field assistance. LITERATURE CITED Baroni, T. J. & R. E. Halling. 1992. New species of Rhodocybe from South America with a key to species. Mycologia 84: 411-421. 385 Halling, R. E. 1989a. A synopsis of Colombian Boletes. Mycotaxon 34: 93-113. 1989b (1990). Notes on Collybia III. Three neotropical species of subg. Rhodocollybia. Mycologia 81: 870-875. . 1992. A new species of Boletus section Luridi from Colombia. Brittonia 44: 108-113. , & C. L. Ovrebo. 1987. A new species of Rozites from oak forests of Colombia with notes on biogeography. Mycologia 79: 674-678. Hennings, P. 1902. Fungi camerunensis novi. III. Bot. Jahrb. Syst. 30: 39-57. Horak, E. & R. E. Halling. 1991. New records of Phaeocollybia from Colombia. Mycologia 83: 464- 472. Kornerup, A. & J. H. Wanscher. 1983. Methuen Handbook of Colour. Ed. 3. Eyre Methuen Ltd., London. Singer, R. 1942. A monographic study of the genera Crinipellis and Chaetocalathus. Lilloa 8: 441-534. . 1976. Marasmieae (Basidiomycetes —Tricholomataceae). Flora Neotrop. Monogr. 17: 1-347. . 1989. New taxa and new combinations of Agaricales (Diagnoses fungorum novorum Agari- calium IV). Fieldiana, Bot., n.s. 21: 1-133. , C. L. Ovrebo, & R. E. Halling. 1990. A new species of Phylloporus and a new species of Tricholomopsis from Colombia with notes on Phylloporus boletinoides. Mycologia 82: 452-459. Tulloss, R. E., C. L. Ovrebo, & R. E. Halling. 1992. Studies on Amanita (Amanitaceae) from Andean Colombia. Mem. New York Bot. Gard. 66: 1-46. ra Sar. i ee | MM 3 : 4 gh be x an ‘, ' ‘ Onl F) 9 } . ; “5 ae, © Ad ' 7 AN ost calf Arie Gi ae , ey fae Ap ‘ ih Blin Oy ait ee, hie, te Lek Tih Reet V. oy ee bs) . A” Ve! Wolves ie LW OM PR il ~ ty fy t sre ¥ PET boat Vege PCL, : a wa pay OY ) . 4 , ry " in ‘ ie os vty th, Ve " ay) pais ul rk a wae iat so ael Cun AG ads SP i? ‘ } TUS" el Eee ij vw peed p ee ve . ahs say nt phi? ‘ bi a. a : by | i, ine A f 7 Saas wi +. . or , ay ie aS iv Re nee sata a ean! tiie Rome rl He Pahang eee by ellie 1 fs ell hates ve aa Loe a = ads ane rere ae angel die ave Jt 5 ae A gy Nien SN . ' Abad, - ih pers Ant je Wessha ny il eae Pet iy iret cL: ay ay a Ms. nr ty rn ae rey? neces ALi 1 ath y Aha) a, . a A y Ane | : } ie , (ah LA hes » ean oe ane Gert | Sale ete near rene hg ny ce Toa e 4 big’ . ive Wh v } 5 : ; ie se ba — ink , ew | ; ; my ; ay x « a * A. Dy: b 4 ° , al inn: ae Ms wi ae Ly a ny e ri - Poe e a = ; ae ar ewe He. Wy hie ef See 7) eal is pti ; bial talk ni on this ise il ive mete tenn | i ‘re } a ast tek “ee shies ay, pine ek ieee at atex agi nha nee aie, . i Aaa Deiat P ye ee ; i ; Aa : copa We, vs i yi! ae f : ¥ 4 + uf ¥i wu] ei, i af et > ht ~~ | . Be by a! sie Nain earns ep “Nolthh Av oe Cole a wore neat iar Abad rat as UAL oh 6 Ait ine ca i a son sgctal vie Ney hel ohh rt a is ey WRC a 0 cae ta ae oe an ‘ " ny, ‘ oe bee 4 is a i< i hale a! et b. eve? a ie alae rm — ce us wale toy yi inh aT oy mT at ie ae » ya yea} yi apn: Ue. A es ow | : e FY { e' , y q Me ee Py. oe rr pita i ae : T ; } H ' si co : oe nes MY COTAXON Volume XLVI, pp.387-388 April-June 1993 SEPTORIA THYMI SP. NOV. FROM BULGARIA Ekaterina F. Sameva and Ganka G. Bakalova Institute of Botany, 1113 Sofia, Bulgaria ABSTRACT. A new species of the genus Septoria Sacc. (Coelomycetes), Septoria thymi Sameva et Bakalova sp. nov. from Bulgaria is described and illustrated. During inventory investigations of the genus Septoria Sacc. (Sphaeropsidales, Coelomycetes) in Bulgaria, a new species belonging to this genus is established. Septoria thymi Sameva et Bakalova, sp. nov. Maculae 0.1-0.3 cm in diam, orbiculares vel irregularis, solitariae, albidis, ochraceae vel pallide-brunneae indistincto atropurpureo marginate. Pycnides epiphylae, globosae vel depresso- globosae tenuiter tunicatae, pallido-brunneae, 140-180 um in diam, poro orbiculari 10-20 um in diam, cellulis parvis obscurioribus cincto. Conidia cyindrica, elongato-fusiformia, clavata, utrinque rotundata, recta vel leniter curvata, 3-5(7) septata, hyalina, 30.0-(X =38.94; S =4.31)-54.0 x 2.5-(X =3.41; S=0.43)-5.0 um, N=100 (Figs. 1, 2). | In foliis Thymi pulegioidi L., Bulgaria, mons Vitosha, supra pagum Bistrica, 13.07.1980, E. F. Sameva, G. G. Bakalova, SOM 20763 M, holotypus. In speciebus Thymi Septoria primum observatur. Leaf spots 0.1-0.3 cm in diam, rounded or irregular, single, whitish, ochraceus or pale brown, surrounded by a brown or purple border. Pycnidia epiphyllous, spheric or flattened, with thin walls, pale brown, 140-180 um in diam, with circular ostiole, surrounded by darker cells, 10-20 wm in diam. Conidia cylindric, fusiform or clavate, rounded at both ends, straight or slightly curved, 3-5 (rarely 7) septate, hyaline, 30.0-(X-38.94; S=4.31)-54.0 x 2.5-(X =3.41; $=0.43)-5.0 um. N=100 (Figs. 1, 2). 388 On leaves of Thymus pulegioides L., Bulgaria, Mount Vitosha, above the village of Bistritsa, 13.07.1980, E. F. Sameva, G. G. Bakalova, SOM 20763 M (holotype). First description of a species of Septoria Sacc. on Thymus L. Figs. 1, 2. Pycnidium and conidia of Septoria Thymi MY COTAXON Volume XLVII, pp. 389-394 April-June 1993 THREE NEW SPECIES AND A NEW VARIETY OF PLUTEUS FROM THE UNITED STATES PARTHA BANERJEE! AND WALTER J. SUNDBERG Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901, U.S.A. ‘Present Address: Biological Survey, Cultural Education Center, New York State Museum, Albany, NY 12230, U.S.A. ABSTRACT: Three previously unreported taxa, Pluteus cinerascens, P. heterocystis, P. laricinus, and P. salicinus var. americanus were discovered during field and herbarium studies of Pluteus section Pluteus (Pluteaceae, Agaricales) in the United States. They are described below as new. KEY WORDS: Agaricales, Pluteus, United States. The genus Pluteus Fr. (Pluteaceae, Agaricales) is a large group of presumably non- mycorrhizal mushrooms found on all continents excepting Antarctica (Singer 1986). The genus has three sections: Pluteus Fr., Hispidoderma Fayod and Celluloderma Fayod. In section Pluteus, the pileipellis is a filamentous cutis with cylindric or cylindric-fusoid end- cells; the hyphae are appressed with the end-cells sometimes erect or suberect in fascicles. The pleurocystidia are metuloid. Metuloid cystidia are thick-walled, veniricose (i. e., swollen in the middle), and usually with apical projections. In the following descriptions, color names within quotes are from Ridgway (1912, R); Kelly and Judd (1976, = NBS); and the Royal Botanic Garden Color Chart (1969, RBG). Color names without quotes are author-generated. All anatomical observations were made in 3% KOH. The following are definitions used to describe pleurocystidia: "cervinus-type" refers to pleurocystidia with apical ornamentations while "magnus-type" indicates pleurocystidia without apical ornamentations (modified after Singer 1956). Unless otherwise noted, herbarium abbreviations (Holmgren et al. 1990) for specimens examined are given in parentheses after the specimen number. Standard two-lettered abbreviations are used for American states. 390 Pluteus cinerascens P. Banerjee et W. J. Sundberg, sp. nov. Fig. 1. Speciebus aliis sectionis Plutei, pileo 2-7 cm lato, margine cinerasci ubi vulnerato (injurio vel contuso); cervini-pleurocystidiis metuloideis ventricosis, tunica #2.2 Lm crassa, apice acuto vel truncato, appendiculo, cornibus 2-4 plerumque fortibus longis acutis; epicutis pilei fibulis destituta. Holotypus: A. H. Smith 57433! (MICH). Solitary on a very rotten log, Proud Lake Recreation Area, Oakland Co., Michigan, U.S.A. Pileus 2-7 cm broad, broadly convex with a slightly flattened disc, surface with small fascicles of appressed, cinnamon-buff fibrils on a white ground color over the disc, white and glabrous elsewhere, a dingy orange flush developing in some areas overnight, injured and bruised places along margin becoming "cinereous" (R) to "clay buff" (RBG); context white, soft; odor and taste raphanoid. Lamellae free and remote from stipe, close, broad, white becoming dingy pale "vinaceous" (R) to "vinaceous buff" (RBG), edges serrulate, concolorous with lamellae, beaded with drops when fresh; lamellulae 1-2-tiered. Stipe 6.5- 10 cm long, 0.2-1 cm thick at apex, base cylindrical to clavate and up to 1.5 cm broad, naked above, lower third may or may not have appressed fibrils, fibrils white overall but staining or discoloring gray to fuliginous in injured places and with dingy "ochraceous" (R) streaks over the base; solid but fragile. Basidiospores 6-7.4 x 4.4-5.6 jim, oval-ellipsoid, wail smooth, moderately thick, rarely collapsing, with hyaline cytoplasmic content, some with a large lipid droplet; hilar appendix inconspicuous. Basidia 20-28 x 7-8.4 um, clavate; four-spored. Pleurocystidia very abundant, (30-)42-85 x 12-22 um, ventricose, thick-walled (1.4-2.0 um), almost all hyaline ("content pale lilac in KOH when fresh", Smith, unpublished); of cervinus-type, apex acute to truncate and with (1-)2-4(-5) well-developed, mostly long and sharp apical hooks, some pleurocystidia with short to blunt apical hooks, lateral spinules occasionally present, occasional sphaeropedunculate pleurocystidia interspersed with those of the cervinus-type; pleurocystidia near lamellar edge mostly of cervinus-type interspersed with a few magnus-type. Cheilocystidia not abundant to abundant, in fascicles when abundant; 30-65 x 12-22 um, sphaeropedunculate, hyaline; with a tendency to gelatinize when remoistened. Pileipellis hyphae 4-12 tm in diameter, ends rounded, most hyaline, some with a homogeneous dark to light brown content; pellicle in cross section easily separates from the context; clamp connections absent on pileipellis hyphae. Other collections examined: Klavins s. n., Pomona Natural Bridge, Jackson Co., IL (SIU). Observations: The morphological and anatomical characters are significant enough to describe it as a new species. A. H. Smith (unpublished) considered P. cinerascens close to P. magnus McClatchie, but the presence of very strongly developed long, sharp pleurocystidial hooks indicate closer affinities to P. cervinus (Schaeff.: Fr.) Kumm. The cinerous Stains and the strongly developed hooks on many of the pleurocystidia distinguish it. A solitary liginicolous basidiocarp of the type specimen was collected from Michigan by A. H. Smith, and another solitary terrestrial basidiocarp has been found recently in Illinois. Figs. 1-4. Anatomical characters of Pluteus species. 1. Pluteus cinerascens sp. nov.; 2. P. heterocystis sp. nov.; 3. P. laricinus sp. nov.; 4. P. salicinus var. americanus var. nov. (P: Pleurocystidia; C: Cheilocystidia; L: Pileipellis; B: Basidia; S: Basidiospores. Scale lines: 10 um for basidiospores; 20 um for other structures). 392 The morphological description as well as notes on its habit, habitat and distribution of the type specimen are adapted from data obtained by A. H. Smith (unpublished). Pluteus heterocystis P. Banerjee et W. J. Sundberg, sp. nov. Fig. 2. Speciebus aliis sectionis Plutei, pileo 5 cm lato, disco umbrino et fusco fibrillulo in Strias versus marginem radiatim; lamellis crassis marginibus cariosis; pleurocyStidiis metuloideis clavatis late ventricosis, apice obtuso vel rotundato, appendiculo cornibus brevibus vel nullis; epicutis pilei fibulis destituta. Holotypus: A. H. Smith 13616! (MICH). Solitary on soil, Hoh River, Olympic National Park, Washington, U.S.A. Known only from the type collection. Pileus 5 cm broad, expanded-umbonate, umbo obtuse, moist to subviscid (as in Inocybe geophylla); disc "Saccardo's umber" (R) and with dark fibrils extending as streaks toward the margin; marginal area watery gray but opaque; odor and taste not recorded. Lamellae free, remote from the stipe, close, moderately broad, somewhat thick, white becoming dingy pink; edges eroded and pallid; lamellulae 1-2-tiered. Stipe 7 cm long, 1 cm thick at apex, equal to a slightly bulbous base, glabrous over most of the surface, some "umber" (RBG) fibrillose streaks present on lower portion; solid; context pallid. Basidiospores 6.6-8(-9) x 5.6-6.4(-7) um, broadly ovate to ellipsoid, smooth, wall slightly thick, some hyaline and with a large lipid droplet, some with cytoplasmic content; hilar appendix visible. Basidia 24-32 x 8-10 um, clavate, usually 4-spored, occasionally 2-spored, hyaline. Pleurocystidia abundant and dimorphic: one form 50-80 x 10-25 um, clavate to broadly fusoid-ventricose, apices more or less truncate, many with several short horns at apex and others merely with aborted apical ornamentation; the second form with apex rounded and lacking horns, wall uniformly 1.2-1.8 um thick, no lateral ornamentation seen; all pleurocystidia hyaline., Cheilocystidia 35-45(-54) x 8-12.6 um, clavate-pedicellate to slightly ventricose, thin-walled, hyaline; apices obtuse to rounded, smooth. Pileipellis of radial hyphae 6-10 tm in diameter; end cells often subfusoid and 9-14 tm in diameter; content hyaline or rarely dingy yellow-brown, rarely a few hyaline pilocystidia (15-25 um diameter) visible; clamp connections not seen on pileipellis hyphae. Observations: The thick gills with eroded edges, and the presence of two types of pleurocystidia with one form having truncated apices with short ornamentations are distinctive. A. H. Smith (unpublished) found clamp connections in the pileipellis, but our repeated attempts failed to demonstrate them. The morphological description as well as data on its habit, habitat and distribution are modified from notes of A. H. Smith (unpublished). Pluteus laricinus P. Banerjee et W. J. Sundberg, sp. nov. Fig. 3. Speciebus aliis sectionis Plutei, pileo 1.5 cm lato, fusci-umbrino, squamuloso in disco; lamellis albis ad marginem pilei griseolis; epicute pilei fibulis infrequentibus; stipite gracili, 0.2 cm in diametro ad apicem. Holotypus: A. H. Smith 38888! (MICH). Solitary on a rotten larch log. Proud Lake Recreation Area, Oakland Co., Michigan, U.S.A. Known only from the type collection. 393 Pileus 1.5 cm broad, convex-umbonate, squamulose on the disc, elsewhere appressed- fibrillose, dark "wood brown" (R); margin slightly wavy but not striate. Lamellae broad, remotely free, close, white, becoming pale pink and edges grayish near cap margin; lamellulae 1-2-tiered. Stipe 3 cm long, 2 cm thick at apex, only slightly enlarged downward, whitish and not darkening, with faint "avellaneous" (R) pruina near base. Basidiospores (4.4-)5-6.4 x 4.6-5 1m, oval-ellipsoid, smooth, wall only slightly thick, hyaline or nearly so; hilar appendix visible. Basidia 22-25 x 6-7.4 tm, clavate, four- spored, hyaline. Pleurocystidia 35-75 x 8-20 tm, fusoid-ventricose, hyaline, wall of upper part thicker (1.2-1.5 jum thick) than lower part; most pleurocystidia near the gill edge are of magnus-type (i.e., without hooks), others with 2-3 hooks at the apex, rarely with 1 lateral hook. Cheilocystidia 28-48 x 10-14.6 um, clavate-pedicellate to broadly fusoid, smooth and apices rounded, walls thin, content hyaline to dingy brownish. Pileipellis hyphae 3-5 tm in diameter, with rounded to subacute apices, occasionally hyphal end cells fusoid-ventricose. Pilocystidia 85-150 x 8-14 um, deeply embedded in the pileal trama, content of all pileipellis hyphae hyaline or dull yellow brown; clamp connections on pileipellis hyphae present but not frequent. Observations: The small spores, the grayish gill margin, and slender stipe distinguish the species. A. H. Smith (unpublished) found abundant clamp connections on pileipellis hyphae, but after repeated attempts, we found only a few. The morphological description as well as notes on its habit, habitat and distribution are adapted from notes by A. H. Smith (unpublished). Pluteus salicinus var. americanus P. Banerjee et W. J. Sundberg, var. nov. Fig. 4. Varietatibus aliis Plutei salicini, pileo 5-6 cm lato, glabro hygrophano, ad marginem translucidi-striato; pleurocystidiis acutis vel truncatis, appendiculis cornibus apicis complexis; epicute pilei fibuligera. Holotypus: A. H. Smith 57841! (MICH). Solitary on aspen wood. Pellston Hills, west of Pellston, Michigan, U.S.A. Pileus 5-6 cm broad, broadly convex expanding to plane with no umbo, color between "drab" (R) and "hair brown" (R), surface moist and hygrophanous, sometimes turning greenish-gray upon handling or drying, opaque when faded; lacking any spinules or squamules over disc; margin translucent-striate fading to an ashy gray; context watery grayish fading to pallid; odor of crushed context disagreeable, taste very disagreeable: somewhat metallic. Lamellae fairly broad, remote, close, whitish pallid, "pink" (NBS) in age; edges concolorous; lamellulae 1-2-tiered. Stipe 4-6 cm long, 0.5-0.6 cm thick, silvery whitish to grayish and base soon staining greenish gray when handled or dried; naked; hollow, terete or compressed. Basidiospores 8-10(-11) x 6.4-7.4 um, ellipsoid to ovoid, smooth, hyaline, wall slightly thick; hilar appendix conspicuous. Basidia 28-34 x 7-8.8 tm, clavate, both 2- and four-spored. Pleurocystidia abundant, 50-85 x 12-28 1m, ventricose to very broadly fusoid with apex more or less truncate, some with simple sharp and pointed projections, some others with short (2-5 zm) truncate projections arising around the neck and these usually with a crown of 4-5 small but pronounced projections around their apices (i.e., primary and secondary ornamentations present), rarely some aborted ornamentations observed on the neck just below the apex; wall from ventricose portion downward to pedicel becoming thin and readily collapsing, 1.2-1.4 um thick near neck, +1 um thick near pedicel; all pleurocystidia hyaline. Cheilocystidia in fascicles, 34-46 x 12-20 um, 394 sphaeropedunculate to clavate-pedicellate, thin-walled, smooth, hyaline. Pileipellis a trichoderm formed of dimitic hyphae 3-6 1m in diameter, occasionally with cystidioid end- cells up to 16 tm in diameter, content pale brownish "fuscous" (R) to dingy brown; clamp connections present on pileipellis hyphae. Other collections examined: A. H. Smith 57846, MI (MICH); A. H. Smith 57099, MI (MICH); A. H. Smith 57787, MI (MICH)—no other data available on these collections; TN 14277, Indian Creek woods, Great Smoky Mountains National Park, Swain Co., NC (TENN). Observations: This taxon is in the Pluteus salicinus (Pers.: Fr.) Kumm. group, but is distinctive because of its hygrophanous pileus with translucent-striate margin when moist. Also, the presence of some pleurocystidia with compound apical ornamentations is distinctive. Among the other recognized American varieties of P. salicinus, P. salicinus var. salicinus is not hygrophanous, and P. salicinus var. achloes Sing. is occasionally hygrophanous. Moreover, these two varieties almost never have pleurocystidia with compound apical ornamentations. InP. salicinus var. americanus, the stipe (and sometimes pileus) turn dark bluish-fuscous upon handling or drying which is quite distinctive for a Pluteus but this character is present to some extent throughout the P. salicinus group. ACKNOWLEDGEMENTS We wish to thank Dr. Gregory M. Mueller of the Field Museum of Natural History for his pre-submission review of the manuscript. We express our gratitude to Dr. Charles Sheviak and Ms. Patricia Kernan of the New York State Museum Biological Survey for their help with the latin diagnoses and illustrations, respectively. The University of Michigan Herbarium is thanked for the loan of the Pluteus specimens. LITERATURE CITED Henderson, D. M., P. D. Orton, and R. Watling. 1969. Colour Identification Chart. British Fungus Flora. Agarics and Boleti: Introduction. Royal Botanic Garden, Edinburgh. 84 pl. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett (Eds.). 1990. Index Herbariorum. Part I. 8th Ed. New York Botanical Garden, Bronx. 693 p. Kelly, K. L., and D. B. Judd. 1976. Color. Universal Language and Dictionary of Names. Nat. Bur. Stand. Spec. Publ. 440. U.S. Gov't. Printing Office, — Washington, D.C. 158 p. Ridgway, R. 1912. Color Standards and Color Nomenclature. Publ. by the author. Washington, D.C. 43 p.+ 53 pl. Singer, R. 1956. Contributions toward a monograph of the genus Pluteus. Trans. Brit. Mycol. Soc. 39(2): 145-232. Singer, R. 1986. The Agaricales in Modern Taxonomy. 4th ed. Koeltz Scientific Books, Koenigstein, F.R.G. 981 pp. MY COTAXON Volume XLVI, pp. 395-403 April-June 1993 WORLD LIST OF CETRARIOID LICHENS TIINA RANDLANE & ANDRES SAAG Institute of Botany and Ecology, Tartu University Lai Street 38, Tartu, Estonia EE-2400 Abstract. The list sums up 120 cetrarioid lichen species. The present generic location and valid epithet is indicated for each of them. The generic location of 27 species (marked with *) is not acceptable but not yet converted. Information available about 9 species (marked with ?) is unsufficient. This is the first attempt after 40 years (Rassadina, 1950; Rdsdnen, 1952) to compile a total list of cetrarioid lichens. The terms "cetrarioid", "parmelioid" and "alectoroid lichens" are nowadays in wide use but not strictly defined. All those terms indicate the certain morphological trends in the fam. Parmeliaceae, "cetrarioid" being perhaps the most unlimited of the three. Classically foliose thallus with dorsiventral structure which is not closely adnate to the substrate is characteristic to this trend as well as dominating marginal position of apothecia and pycnidia. We have followed this classical treatment and comprised into the list of cetrarioid lichens all the species corresponding to those general characters. According to the contemporary taxonomy the cetrarioid lichen species may belong to different genera while most of them have earlier been included in the genus Cetraria. The latter was described in 1803 by Erik Acharius and consisted then of 8 species. The number of species in the genus has varied greatly being the biggest (76) in the treatment of the Russian lichenologist Ksenya Rassadina in 1950. At present the obvious trend in the systematics of the fam. Parmeliaceae is towards the delimitation of small and more or less homogeneous genera. During last decades 13 separate genera of cetrarioid lichens have been newly described or resurrected: Ahtiana (Goward, 1985), Allocetraria (Kurokawa & Lai, 1991), Asahinea (Culberson & Culberson, C.,1965), Cetrariopsis (Kurokawa, 1980), Cetrelia (Culberson & Culberson, C., 1968), Cetreliopsis (Lai, 1980), Esslingeriana (Hale & Lai in Lai, 1980), Masonhalea (Karnefelt, 1977), Nephromopsis (Miller Argoviensis, 1891; resurrected by Lai in 1980), Parmelaria (Awasthi, 1987), Platismatia (Culberson & Culberson, C., 1968), Tuckermannopsis (Gyelnik, 1933; resurrected by Hale in Egan, 1987) and Vulpicida (Mattsson & Lai, 1993). Among these units the genera Cetraria 396 and Tuckermannopsis are still the conglomerates of poorly connected taxa and in urgent need of further studies. Description of some more new cetrarioid genera are expected in the near future. In our list the specific epithets are presented in the alphabetical order. The present generic location and valid epithet is indicated for each of them. All together 120 species are accepted as separate. Their epithets are numbered. The epithets without numbers are treated as synonyms, or they represent the species of other, not cetrarioid genera. There is not much information available about 9 species which may therefore turn out to be either synonyms or belong even to quite different genera. These species are marked with ? in the list. The species which present generic location is not acceptable but not yet converted, are marked with * The synonymity of some species is still doubtful. On these occasions the marks =? are used. aculeata (Schreber) Fr. [Cetraria] = Coelocaulon aculeatum (Schreber) Link *1. agnata (Nyl.) Krist. [Cetraria] = Cetraria agnata (Nyl.) Krist. 2. alaskana Culb.& C.Culb. [Cetraria] = Cetrelia alaskana (Culb. & C.Culb.) Culb.& C.Culb. ?3. albopunctata Zahlbr. {Cetraria] aleurites (Ach.) Th.Fr. [Cetraria] = Imshaugia aleurites (Ach.) S.F.Meyer alvarensis (Wahlenb.) Lynge [Cetraria] = Vulpicida tubulosus (Schaerer) J.-E.Mattsson & Lai 4. ambigua Bab. [Cetraria] = Allocetraria ambigua (Bab.) Kurok. & Lai americana Sato [Cetraria| = Tuckermannopsis orbata (Nyl.) Lai 5. americana Sprengel [Nephroma] = Tuckermannopsis americana (Sprengel) Hale 6. andrejevii Oxner [Cetraria] = Cetraria andrejevii Oxner ?7. annae Oxner [Cetraria] 28. antarctica Zahlbr. aradensis Gyelnik [Pseudoparmelia| = Cetrelia olivetorum (Nyl.) Culb. & C.Culb. arborialis (G.K.Merr.) Howard [Cetraria] = Cetraria subalpina Imsh. arctica (Hook.) Tuck. [Cetraria] = Dactylina arctica (Hook.) Nyl. arctica Magnusson [Cetraria] = Cetraria nigricascens (Nyl.) Elenkin 9. arenaria Karnef. [Cetraria] = Cetraria arenaria Karnef. 10. asahinae Sato [Cetraria] = Nephromopsis asahinae (Sato) Raésdénen atlantica (Tuck.) Du Rietz [Cetraria] = Platismatia tuckermanii (Oakes) Culb. & C.Culb. *11. aurescens Tuck. [Cetraria] = Tuckermannopsis aurescens (Tuck.) Hale 12. australiensis W.Weber ex Karnef. [Cetraria] = Cetraria australiensis W.Weber ex Karnef. bavarica Krempelh. [Cetraria] = Tuckermannopsis oakesiana (Tuck.) Hale billardieri Mont. [Cetraria] = Hypogymnia billardieri (Mont.) Filson bohemica Anders [Cetraria] = Coelocaulon aculeatum (Schreber) Link 13. braunsiana (Mill. Arg.) Zahlbr. [Cetraria] = Cetrelia braunsiana (Miill. Arg.) Culb. & C.Culb. californica G.K.Merr. [Cetraria] = Tuckermannopsis merrillii (Du Rietz) Hale *14, 15. 16. 1; *18. Lo 20. ZA B22. Pao 24. 2). 397 californica Tuck. [Cetraria] = Cetraria californica Tuck. californica Gyelnik [Nephromopsis| = according to the description (Gyelnik, 1931) ought to be Tuckermannopsis orbata (Nyl.) Lai. This is confirmed by TLC of isotype (FH) which contains protolichesterinic acid. canadensis (Rasanen) Rasanen [Cetraria] = Vulpicida canadensis (Rasaénen) J.-E.Mattsson & Lai caperata sensu Vainio [Cetraria] = Vulpicida pinastri (Scop.) J.-E.Mattsson & Lai capitata Lynge [Cetraria] = Cetraria nigricans Nyl. cavernosus Menzies [Lichen] = Platismatia lacunosa (Ach.) Culb. & C.Culb. cetrarioides (Delise ex Duby) Nyl. [Parmelia] = Cetrelia cetrarioides (Delise ex Duby) Culb. & C.Culb. chicitae Culb. [Cetraria] = Cetrelia chicitae (Culb.) Culb. & C.Culb. chlorophylla (Willd.) Vainio [Cetraria] = Tuckermannopsis chlorophylla (Willd.) Hale chrysantha Tuck. [Cetraria] = Asahinea chrysantha (Tuck.) Culb. & C.Culb. ciliaris Ach. [Cetraria] = Tuckermannopsis ciliaris (Ach.) Gyelnik citrina Taylor [Cetraria] =? Nephromopsis pallescens (Schaerer) Park clarkti Tuck. [Cetraria] {nomen nudum} = Nephromopsis endocrocea Asah. f. clarktt Asah. collata (Nyl.) Miill.Arg. [Cetraria] = Cetrelia collata (Nyl.) Culb. & C.Culb. commixta (Nyl.) Th.Fr. [Cetraria] = Cetraria commixta (Nyl.) Th.Fr. complicata Laurer [Cetraria] = Nephromopsis laureri (Krempelh.) Kurok. coralligera (W.Weber) Hale [Cetraria] = Tuckermannopsis coralligera (W.Weber) W.Weber corallophora Mill.Arg. [Cetraria] =Punctelia rudecta (Ach.) Krog corniculata Egeling [Cetraria] = Coelocaulon aculeatum (Schreber) Link corrugis Fr. [Cetraria] = Parmotrema perforatum (Wulfen) Hale crispa (Ach.) Nyl. [Cetraria] = Cetraria ericetorum Opiz cucullata (Bellardi) Ach. [Cetraria| = Allocetraria cucullata (Bellardi) Randl. & Saag culbersonit Hale [Cetraria] = Cetraria culbersonii Hale culbersoniorum ‘Trass [Asahinea] = Asahinea chrysantha (Tuck.) Culb. & C.Culb. daibuensis Rasanen [Cetraria] = Nephromopsis laxa (Zahlbr.) Sato . davidiana Culb. & C.Culb. [Cetrelia] = Cetrelia davidiana Culb. & C.Culb. . delavayana Culb. & C.Culb. [Cetrelia] = Cetrelia delavayana Culb. & C.Culb. . delavayi (Hue) Sato [Cetraria] = Nephromopsis delavayi Hue . delisei (Bory ex Schaerer) Nyl. [Cetraria] = Cetraria delisei (Bory ex Schaerer) Nyl. . denticulata Hue [Cetraria] . dermatoidea Zahlbr. [Cetraria] diffusa (Weber) Lynge [Cetraria] = Imshaugia placorodia (Ach.) S.F.Meyer . ectocarpisma (Hue) Gyelnik [Nephromopsis] = Nephromopsis ectocarpisma (Hue) Gyelnik elenkinii Krog [Cetraria] = Cetraria nigricascens (Nyl. in Norrl.) Elenkin 398 S87 34. DD: 36. oe *38. 39. 40. 41. 42. *43. 44, *40: 46. *A7. 48. 49, 50. endocrocea (Asah.) Sato [Cetraria] = Nephromopsis endocrocea Asah. endoxantha Hue [Cetraria] = pro parte Nephromopsis endocrocea Asah., pro parte Nephromopsis ornata (Miill. Arg.) Hue endoxanthoides Awasthi [Cetraria] = Nephromposis endoxanthoides (Awasthi) Randl. & Saag epiphorella Nyl. in Crombie [Cetraria] = Coelopogon epiphorellum (Ny)l. in Crombie) Brusse & Karnef. ericetorum Opiz [Cetraria] = Cetraria ericetorum Opiz eriophylla (Knight) Zahlbr. [Cetraria] = Erioderma knighti Shirley erosa Culb. & C.Culb. [Platismatia] = Platismatia erosa Culb. & C.Culb. everniella (Nyl.) Krempelh. [Cetraria] = Allocetraria stracheyi (Bab.) Kurok. & Lai fahlunensis sensu Schaerer [Cetraria] = Cetraria hepatizon (Ach.) Vainio fahlunensis sensu Vainio [Cetraria] = Cetraria commixta (Nyl.) Th.Fr. fallax (Weber) Ach. [Cetraria] = Platismatia glauca (L.) Culb. & C.Culb.. fastigiata (Delise ex Nyl. in Norrl.) Karnef. [Cetraria] = Cetraria fastigiata (Delise ex Nyl. in Norrl.) Karnef. fendleri (Nyl.) Tuck. [Cetraria] = Tuckermannopsis fendleri (Ny1.) Hale formosana Zahlbr. [Cetraria] = Platismatia formosana (Zahlbr.) Culb. & C.Culb. frostii Tuck. [Parmelia] = Cetraria culbersonii Hale furcellata Fr. [Cetraria] = Bryoria furcellata (Fr.) Brodo & D.Hawksw. gilva Asah. [Cetraria| = Tuckermannopsis gilva (Asah.) Lai glauca (L.) Ach. [Cetraria] = Platismatia glauca (L.) Culb. & C.Culb. globulans (Nyl. ex Hue) Zahlbr. [Cetraria] = Nephromopsis globulans (Nyl. ex Hue) Lai gracilenta (Krempelh.) Vainio [Cetraria] = Coelopogon epiphorellum (Nyl. in Crombie) Brusse & K4rnef. halei Culb. & C.Culb. [Cetraria] = Tuckermannopsis americana (Sprengel) Hale hepatizon (Ach.) Kurok. [Tuckermannopsis] = Cetraria hepatizon (Ach.) Vainio. herrei Imsh. [Cetraria] = Platismatia herrei (Imsh.) Culb. & C.Culb. hiascens (Fr.) Th.Fr. [Cetraria] = Cetraria delisei (Bory ex Schaerer) Nyl. hypotrachyna Mill.Arg. [Cetraria] = Cetraria hypotrachyna Mill.Arg. iberica Crespo & Barreno [Cetraria] = Tuckermannopsis merrillii (Du Rietz) Hale idahoensis Essl. [Cetraria] = Esslingeriana idahoensis (Ess}.) Hale & Lai inermis (Nyl.) Krog [Cetraria] = Cetraria inermis (Nyl.) Krog inflata J.D.Hook. & Taylor [Cetraria] = Ramalina inflata (J.D.Hook. & Taylor) J.D.Hook. & Taylor interrupta Culb. & C.Culb. [Platismatia] = Platismatia interrupta Culb. & C.Culb. isidiata (Asah.) Culb. & C.Culb. [Cetrelia] = Cetrelia isidiata (Asah.) Culb. & C.Culb. isidiigera Kurok. & Lai [Allocetraria] = Allocetraria isidiigera Kurok. & Lai S79 51. isidioidea (Rasaénen) Awasthi [Cetraria} = Nephromopsis _ tsidioidea (Rasénen) Randl. & Saag 52. japonica Zahlbr. [Cetraria] = Cetrelia japonica (Zahlbr.) Culb. & C.Culb. 53. islandica (L.) Ach. [Cetraria] = Cetraria islandica (L.) Ach. 54. juniperina (L.) Ach. [Cetraria] = Vulpicida juniperinus (L.) J.-E.Mattsson & Lai 55. kamczatica Savicz [Cetraria] = Cetraria kamczatica Savicz 56. komarovii Elenkin [Cetraria] = Nephromopsis komarovii (Elenkin) Wei kurodakensis Asah. [Cetraria] = Asahinea scholanderi (Llano) Culb. & C.Culb. *57. kurokawae Shibuichi & Yoshida [Cetraria] = Nephromopsis kurokawae (Shibuichi & Yoshida) Kurok. lacera J.D.Hook. & Taylor [Cetraria] = Sticta lacera (J.D.Hook. & Taylor) Mill. Arg. 58. lacunosa Ach. [Cetraria] = Platismatia lacunosa (Ach.) Culb. & C.Culb. lacunosa Elenkin [Cetraria] = Asahinea chrysantha (Tuck.) Culb. & C.Culb. lacunosa Th.Fr. [Cetraria] = Platismatia norvegica (Lynge) Culb. & C.Culb. laeteflava Zahlbr. [Cetraria] =? Cetreliopsis rhytidocarpa (Zahlbr.) Lai 59. laevigata Rassad. [Cetraria] = Cetraria laevigata Rassad. lanata (Necker) Schaerer [Cetraria] = Pseudephebe pubescens (L.) M.Choisy 60. laureri Krempelh. [Cetraria] = Nephromopsis laurert (Krempelh.) Kurok. *61. laxa (Zahlbr.) Sato [Cetraria] = Nephromopsis laxa (Zahlbr.) Sato *62. leucostigma Lev. [Cetraria| = Cetraria leucostigma Lev. libertina Stuckenb. [Cetraria] = Cetraria ericetorum Opiz madreporiformis (Ach.) Miill.Arg. [Cetraria] = Dactylina madreporiformis (Ach.) Tuck. magnussonii Llano [Cetraria] = Cetraria nigricascens (Nyl. in Norrl.) Elenkin megaleia (Nyl.) Rasanen [Cetraria] = Cetrelia sanguinea (Schaerer) Culb. & C.Culb. *63. melaloma Krempelh. [Cetraria] = Cetraria melaloma Krempelh. *64. merrillii Du Rietz [Cetraria] = Tuckermannopsis merrillii (Du Rietz) Hale 265. microphylla Elenkin [Cetraria] 66. microphyllica Culb. & C.Culb. [Cetraria] = Tuckermannopsis microphyllica (Culb. & C.Culb.) Lai 67. monachorum Zahlbr. [Parmelia] = Cetrelia monachorum (Zahlbr.) Culb. & C.Culb. 68. morrisonicola Lai [Nephromopsis] = Nephromopsis morrisonicola Lai muricata (Ach.) Eckf.[Cetraria] = Coelocaulon muricatum (Ach.) Laundon 69. nepalensis Awasthi [Cetraria] = Cetraria nepalensis Awasthi nephromoides (Nyl.) Vainio [Cetraria] = Nephromopsis ectocarpisma (Hue) Gyelnik 70. nigricans Nyl. [Cetraria] = Cetraria nigricans Ny]. 71. nigricascens (Nyl. in Norrl.) Elenkin [Cetraria] = Cetraria nigricascens (Ny). in Norrl.) Elenkin nipharga (Ach.) R6ohling = Allocetraria cucullata (Bellardi) Randl. & Saag 72. nipponensis (Asah.) Culb. [Cetraria] = Nephromopsis nipponensis (Asah.) Lai 400 TS nivalis (L.) Ach. [Cetraria] = Allocetraria nivalis (L.) Randl. & Saag normoerica (Gunn.) Lynge [Cetraria] = Cornicularia normoerica (Gunn.) Du Rietz . norvegica (Lynge) Du Rietz [Cetraria] = Platismatia norvegica (Lynge) Culb. & C.Culb. nova-zelandiae Zahlbr. [Cetraria] . nuda (Hue) Culb. & C.Culb. [Cetrelia] = Cetrelia nuda (Hue) Culb. & C.Culb. . oakesiana Tuck. [Cetraria] = Tuckermannopsis oakesiana (Tuck.) Hale ochrocarpa (Eggerth) Lettau ([Cetraria} {nomen invalidum} = Tuckermannopsis oakesiana (Tuck.) Hale . odontella (Ach.) Ach. [Cetraria] = Cetraria odontella (Ach.) Ach. . olivetorum Nyl. [Parmelia] = Cetrelia olivetorum (Nyl.) Culb. & C.Culb. . orbata (Nyl.) Fink [Cetraria] = Tuckermannopsis orbata (Nyl.) Lai . orientalis Randl. & Saag [Cetrelia] = Cetrelia orientalis Randl. & Saag . omata Miill.Arg. [Cetraria] = Nephromopsis ornata (Miill.Arg.) Hue pachysperma (Hue) Zahlbr. [Cetraria] = Parmelaria thomsonii (Stirton) Awasthi . pallescens Schaerer [Cetraria] = Nephromopsis pallescens (Schaerer) Park pallida Awasthi [Cetraria] = Cetraria melaloma Krempelh. . pallidula Tuck. ex Riddle [Cetraria] = Tuckermannopsis pallidula (Tuck. ex Riddle) Hale perstraminea Zahlbr. [Cetraria] = Nephromopsis komarovii (Elenkin) Wei . pinastri (Scop.) Gray [Cetraria] = Vulpicida pinastri (Scop.) J.-E.Mattsson & Lai placorodia (Ach.) Tuck. [Cetraria] = Imshaugia placorodia (Ach.) S.F.Meyer platyna Ach. [Cetraria] = Cetraria islandica (L.) Ach. platyphylla Tuck. [Cetraria] = Tuckermannopsis platyphylla (Tuck.) Hale . platyphylloides (Asah.) Sato [Cetraria] = Tuckermannopsis platyphylloides (Asah.) Lai polyschiza (Nyl.) Jatta [Cetraria] = Cetraria hepatizon (Ach.) Vainio . potaninit Oxner [Cetraria| = Allocetraria potaninii (Oxner) Randl. & Saag . pseudocollata Randl. & Saag [Cetrelia] = Cetrelia pseudocollata Randl. & Saag . pseudocomplicata Asah. [Cetraria] = Nephromopsis pseudocomplicata (Asab.) Lai . pseudolivetorum Asah. [Parmelia| = Cetrelia pseudolivetorum (Asah.) Culb. & C.Culb. pubescens (L.) Desport. [Cetraria] = Pseudephebe pubescens (L.) M.Choisy ramulosa (Hook.) Tuck. [Cetraria] = Dactylina ramulosa (Hook.) Tuck. . rassadinae Makryi [Cetraria] = Cetraria rassadinae Makryi . regenerans Culb. & C.Culb. [Platismatia] = Platismatia regenerans Culb. & C.Culb. reticulata Krempelh. [Cetraria] {nomen nudum} = Platismatia erosa Culb. & C.Culb. rhizophora (Vainio) Rassadina [Cetraria] = Cetraria nigricascens (Nyl. in Norrl.) Elenkin *04. 95. 96. se 98. AMEE 100. 101. 102. 103. 104. CALEY *106. Elie *108. 109. 401 rhytidocarpa Mont. & v.d.Bosch [Cetraria] = Cetreliopsis rhytidocarpa (Mont. & v.d.Bosch) Lai richardsonii Hook. in Richardson [Cetraria] = Masonhalea richardsonii (Hook. in Richardson) Karnef. rubescens (Th.Fr.) Vainio [Parmelia] = Cetrelia olivetorum (Nyl.) Culb. & C.Culb. rugosa (Asah.) Sato [Cetraria] = Nephromopsis rugosa Asah. sanguinea Schaerer [Cetraria] = Cetrelia sanguinea (Schaerer) Culb. & C.Culb. saviczii Oxner & Rassad. [Cetraria] = Asahinea scholanderi (Llano) Culb. & C.Culb. scholanderi Liano [Cetraria] = Asahinea scholanderi (Llano) Culb. & C.Culb. scutata (Wulf.) Poetsch [Cetraria] = Tuckermannopsis sepincola (Ehrh.) Hale scutata auct. [Cetraria] = Tuckermannopsis chlorophylla (Willd.) Hale sepincola (Ehrh.) Ach. [Cetraria] = Tuckermannopsis sepincola (Ehrh.) Hale septentrionalis (Nyl.) Almq. [Cetraria] = Asahinea chrysantha (Tuck.) Culb. & C.Culb. sibirica Magnusson [Cetraria] = Cetraria nigricascens (Nyl. in Norr].) Elenkin sikkimensis Rasénen [Cetraria] = Cetraria leucostigma Lev. simmonsii Krog [Cetraria] = Cetraria andrejevii Oxner sinensis Culb. & C.Culb. [Cetrelia] = Cetrelia sinensis Culb. & C.Culb. sorediella (Lettau) Rico & Manrique [Cetraria] {nomen nudum} = Cetraria commicta f. sorediella Lettau sphaerosporella Miill.Arg. [Parmelia] = Ahtiana sphaerosporella (Miill.Arg.) Goward stenophylla (Tuck.) G.K.Merr. [Cetraria] = Platismatia stenophylla (Tuck.) Culb. & C.Culb. stracheyi Bab. [Evernia] = Allocetraria stracheyi (Bab.) Kurok. & Lai stracheyi Bab. [Cetraria] = Nephromopsis stracheyi (Bab.) Miuill.Arg. straminea Krempelh. [Cetraria] straminea Vainio [Cetraria] {nom. illegit} =? Cetreliopsis rhytidocarpa (Mont. & v.d.Bosch) Lai stuppea Flotow ex Sandst. = Coelocaulon muricatum (Ach.) Laundon stygia (L.) Schaerer [Cetraria] = Melanelia stygia (L.) Essl. subalpina Imsh. [Cetraria] = Cetraria subalpina Imsh. subfendleri Essl. [Cetraria] = Cetraria subfendleri Ess\. subperlatum Nyl. [Platysma] = Cetrelia sanguinea (Schaerer) Culb. & C.Culb. subscutata Lindsay [Cetraria] = Cetraria subscutata Lindsay subthomsoni Awasthi [Parmelaria] = Parmelaria subthomsonii Awasthi sulphurea Mont. [Cetraria] =? Nephromopsis pallescens (Schaerer) Park tenuifolia (Retz.) R.H.Howe [Cetraria] = Cetraria ericetorum Opiz tenuissima (L.) Vainio [Cetraria] = Coelocaulon aculeatum (Schreber) Link terrestris (Schaerer) Fink [Cetraria] = Vulpicida tilesti (Ach.) J.-E.Mattsson & Lai teysmanni Mont. & v.d.Bosch [Cetraria] =? Nephromopsis pallescens (Schaerer) Park 402 110. thomsonii (Stirton) Miill.Arg. [Cetraria] = Parmelaria thomsonii (Stirton) Awasthi thyreophora (Ach.) Rohl. [Cetraria] = Cetraria islandica (L.) Ach. 111. tilesii Ach. [Cetraria] = Vulpicida tilesii (Ach.) J.-E.Mattsson & Lai *112. togashii Asah. [Cetraria] = Cetraria togashit Asah. tristis (Weber) Fr. [Cetraria] = Cornicularia normoerica (Gunn.) Du Rietz 113. tubulosa (Schaerer) B.de Lesd. [Cetraria] = Vulpicida tubulosus (Schaerer) J.-E.Mattsson & Lai tuckermanii Herre [Cetraria] {nom. illegit.} = Platismatia herrei (Imsh.) Culb. & C.Culb. tuckermanii R.H.Howe [Cetraria] {nom. illegit.} = Tuckermannopsis mernillii (Du Rietz) Hale 114. tuckermanii Oakes [Cetraria] = Platismatia tuckermanii (Oakes ) Culb. & C.Culb. ulophylla (Ach.) Rebent. [Cetraria] = Tuckermannopsis chlorophylla (Willd.) Hale *115. ulophylloides Asah. [Cetraria] = Tuckermannopsis ulophylloides (Asah.) Lai urceolata (Eschw.) Mont. [Cetraria] = Parmotrema sp. 116. viridis Schwein. [Cetraria] = Vulpicida viridis (Schwein.) J.-E.Mattssom & Lai wallichiana (Taylor) Lai [Ahtia - nom. illegit.] = Cetrariopsis wallichiana (Taylor) Kurok. 117. wallichiana (Taylor) Miill.Arg. [Cetraria] = Cetrariopsis wallichiana (Taylor) Kurok. *118. weberi Essl. [Cetraria] = Cetraria weberi Essl. 2119. xizangensis Wei & Jiang [Cetraria] 120. yunnanensis Zahlbr. [Cetraria] = Nephromopsis yunnanensis (Zahlbr.) Rand. & Saag ACKNOWLEDGEMENTS We wish to express our gratitude to Dr.Ingvar Karnefelt, University of Lund, for general support and reviewing this paper. Professor Teuvo Ahti (University of Helsinki), Dr.Ming-Jou Lai (Dept. of Landscape Architecture, Fu-jen University, Taipei), Jan-Eric Mattsson and Arne Thell (University of Lund) are thanked for fruitful discussions. LITERATURE CITED Awasthi, D.D. 1987. A new position for Platysma thomsonii Stirton. Journ. Hattori Bot. Lab. 63: 367-372. Culberson,W.L. & Culberson, C.F. 1965. Asahinea, a new genus in the Parmeliaceae. Brittonia 17: 182-190. 403 Culberson, W.L. & Culberson, C.F. 1968. The lichen genera Cetrelia and Platismatia (Parmeliaceae). Contr. U.S. Natl.Hern.: 449-558. Egan, R.S. 1987. A fifth checklist of the lichen-forming, lichenocolous and allied fungi of the continental United States and Canada. Bryologist 90: 77-173. Goward, T. 1985. Ahtiana, a new lichen genus in the Parmeliaceae. Bryologist 88: 367-371. Gyelnik, V. 1931. Additamenta ad cognitionem lichenum extraeuropaeorum. Ann. Cryptog. Exot. 4: 166-174. Gyelnik, V. 1933. Lichenes varii novi critique. Acta Fauna et Flora Univ. 1: 3-10. Karnefelt, I. 1977. Masonhalea, a new lichen genus in the Parmeliaceae. Bot. Notiser 130: 101-107. Kurokawa, S. 1980. Cetrariopsis, a new genus in the Parmeliaceae, and its distribution. Mem.Natn. Sci. Mus. 13: 139-142. Kurokawa, S. & Lai, M.J. 1991. Allocetraria, a new lichen genus in the Parmeliacea. Bull. Natn. Sci. Mus. 17: 59-65. Lai, M.J. 1980. Studies on the cetrarioid lichens in Parmeliaceae of East Asia. Quart. Journ. Taiwan Museum 33: 215-229. Mattsson, J.-E. & Lai, M.-J. 1993. Vulpicida, a new genus in Parmeliaceae (lichenized Ascomycetes). Mycotaxon 46: 425-428. Miller Argoviensis, J. 1891. Lichenes Miyoshiani in Japonia a cl. Miyoshi lecti et a cl. Professore Yatabe communicati. Nuov. Giorn. Bot. Ital. 23: 120-131. Rassadina, K.A. 1950. Cetraria in the U.S.S.R. Plantae Cryptogamae, 5: 171-304 (in Russian). Rasanen, V. 1952. Studies on the species of the lichen genera Cornicularia, Cetraria and Nephromopsis. Kuopion Luonnon Ystavain Yhdistyksen Julkaisuja B 2: 1-53. ay a my - oh a | Ne OLA @ owt DI Loa i Py ie =f \ , pe) e JS ny eee eT ie he hy int Es ns a Pad _* Fat reid . > y eee, ; tr a} as peat, Ay te H A Ae ae hae Ws 4 1A athe ha 4 AOC URy ah x Rian “a Ruby tie Any Vi thy ey ; as SASS os i eaahte Nott Danii ac i ee Oe ir | | y Pie cna it, Me Gis y ype ry nr LOR iad i } aie way t Miley ie a bya} in | aN Ut Gl ott ee Ak ye ae } | | nee a ‘G liyh as eRE A Ne | | an ) a Eee nebape Ye ce wed Heche ns i | ' eae HRN He gh Barve te ke r ow \ ~ . f ’ : : i id papel ated Ba oe ssh | vi “s eh ’ re hile, a Ls Pipe ¥ aii bar va 1 A Wt) ( ven mi . ie pn ibm Sid in wb | i ht ai cont Di i r t. ; wh Atk iy WY eu , aes Pen % ay U ae: | ; byt wv f AA Ae | ean asides nom rat accel ih hie | a Liukin ate ieee Ni j ‘ 4 i ne ey sche ity at sith a Has ) { | hh in ; hs } tae (aw rit a if Ly at Pit . a fel lt Ne iy, ms Vs } ar yi sa ; hee ite ki abe ad ay . j As iisy UIE ae” cat Pale x i ok | a, adits , bi at Rarer ied: Ai an We sii mt ay ii pate R athe ae hicieron he ve 4 He : Scares ; s) att ths ve mens My ash mihi ad te 7 Oe evenly chbi ae a aes be ee 8 Vins yh espa ; 4 a av / oe ; che hat ‘ * 7 : rd = 7) = ; i A x 2 eh ae sy a y ’ f j ie y ra) iy a f B ie : iy ee oe bad My oes Amn mrae Matic ny hI clade be") )° Th : ; . * . p fi i a f we [ w a, ew wee aie? 2.1 | ; Li ay ea eae ARE Gee Pp eins ree Ne a TC ee ag tn. EM UN TAA PTE Wie PY CR eer aly tan ay ao Pia pany ah ak aye ce ae ge fu AUR EA Te aie By. Sibays co i ie aay Petrie oui; pane ae WS 1 ; i ‘a P ’ 5 oi ms ; iw laf , aoe q fi] y i cuit Moe ire eo eo pyre sate i aS ae sek i ni eat fish, nia Wie 3 tian an ‘a ie hi ANd RY pee ial Se Ti Bani fi Whe ae a hiss aR | a ib aelams gba peg: AVI Cot Ne & MYCOTAXON Volume XLVI, pp. 405-410 April-June 1993 TWO NEW SPECIES OF CRISTINIA (BASIDIOMYCOTINA, APHYLLOPHORALES) AND A SURVEY OF THE GENUS Kurt Hjortstam Malaregatan 12 S-441 35 Alingsas, Sweden and Helga Grosse-Brauckmann Weingartenstrasse 10 D-6104 Seeheim-Jugenheim, Germany Two new species of Cristinia Parm. are described viz. C. brevicellularis Hjortst. and C. rhenana Grosse-Brauckmann. A new combination Cristinia coprophila (Wakef.) Hjortst. is proposed. The genus Dacryobasidium Jiilich is considered as a synonym of Cristinia. A key to the species of Cristinia is offered. INTRODUCTION Cristinia Parm. with the generic type Hydnum helveticum Pets., is a genus well character- ized by arather loose basidiome, occurrence of hyphal strands and above all with cyanophi- lous granulation in the basidia and thick-walled, cyanophilous basidiospores. The genus was originally introduced with two species, the second being C. sasae Parm. which, however, was later synonymized with C. helvetica by Baici and Hjortstam (1984). CRISTINIA Parm. Consp. syst. corticiacearum p. 47 (1968). BASIDIOMES resupinate, effused,loosely attached to the substratum. HYMENOPHORES hypochnoid, mucedinioid, smooth to grandinioid, more rarely odontioid or raduloid, hyphal strands occur regularly in the periphery. - HYPHAL SYSTEM monomitic; hyphae thin-walled or with slight wall thickening, mostly short-celled at least in the subhymenial layer, with or without clamp-connexions or sometimes with scattered clamps on the subicular hyphae. CYSTIDIA absent. BASIDIA fairly short (15—25 tum), subclavate to cylindrical, slightly constricted, with four sterigmata and a distinct cyanophilous granulation in the protoplasm. SPORES fairly uniform, almost globose to subglobose and often slightly angular, 4-7 um diam., smooth, moderately thick-walled to distinctly thick-walled and cyanophilous, not amyloid or dextrinoid. KEY TO SPECIES OF CRISTINIA i. Clanip-Connexions absenigiegs ni tas send alte wy te eae PRN nu 2 1. Clamp-connexions present throughout, scattered or absent on the subicular hyphae 3 2. Hymenophore distinctly grandinioid, basal hypae 4—6 [um wide. Europe. Be So PIG) nen RR cy ep Oe Vat a ieee PAR ER nach LN fit OY C. artheniensis 406 2. Hymenophore smooth to slightly grandinioid, basal hyphae 8-10 tm wide. Brazil. ete Bl a gre eee rere ee mre te! oT ae eee bet NY Sete na C. brevicellularis 3. Hymenophore raduloid to irpicoid. An uncommon species. Holarctic .. C. gallica 32 Hymenophore smoothito erandinigid 92 eye, ee eee eee eee eee 4 4. Hymenophore grandinioid, all hyphae with clamp-connexions, spores 4.5—-5 um broad. A rather common species. Cosmopolitan (7?) ........... C. helvetica 4. Hymenophore for the most part smooth, basal hyphae either without or with scat- tered clamp=connexions, estat eta Pes AV cee sR eG oo aera ae ates 5 5. Hymenophore normally with a violet tint in KOH, basal hyphae with scattered clamp- connexions, spores 5—7 um diam. Known from Germany only ....... C. rhenana 5. Hymenophore without a violet reaction in KOH, basal hyphae without clamp-con- nexions, spores 4-5 um diam. England, additional records notknown . C. coprophila SPECIES 1. CRISTINIA ARTHENIENSIS Baici & Hjortst. in Sydowia, Annls Mycol. Ser.II, 37:11 (1984). 2. CRISTINIA BREVICELLULARIS Hjortst. nov. sp. Basidioma resupinatum, plus minus laxe adnatum, farinosum vel hypochnoideum. Hyme- nium fere leve vel leviter grandinioideum, pallide isabellinum. Margo simillimus. Systema hyphale monomiticum; totis hyphis efibulatis. Hyphae basales rectae, laxe intertextae, tenuitunicatae, plus minus hyalinae, 8-10 um latae; hyphae subhymeniales brevicellulares, 5-8 tm latae. Hypharum fila presentes; hyphis tenuitunicatis vel crassiusculis, pallide ochraceis, 5—8 tim latis. Cystidia nulla. Basidia subcylindracea, leviter constricta, 15-18 x 6-7 m, 4 sterigmatibus. Sporae subglobosae, leves, moderate crassitunicatae, circiter 5 x 4 m, neque amyloideae, neque dextrinoideae, distincte cyanophilae. Holotypus: Brazil, Sao Paulo, Santos, Cananeia, Ilha do Cardoso, on deciduous wood, 2-5 Feb. 1987, Hjortstam 16753 (K). Isotypus: (GB). BASIDIOME resupinate, loosely adnate, farinaceous to hypochnoid. HYMENOPHORE smooth to slightly grandinioid, pale isabelline. Hyphal strands present in the periphery, with individual hyphae 5—8 um wide, pale ochraceous. Margin indeterminate. HYPHAL SYSTEM monomitic; basal hyphae straight, branched at more or less right angles, thin-walled, hyaline, 8-10 um wide; subhymenial hyphae short-celled, almost isodiametric and with septa at 5-15 um intervals, 5-8 im wide; all hyphae without clamp-connexions. CYSTIDIA absent. BASIDIA subcylindric, with a distinct cyanophilous granulation, slightly constricted in the middle part, 15-18 x 6-7 m, with four sterigmata, lacking a basal clamp-connexion. SPORES subglobose, moderately thick-walled (KOH), smooth, about 5 x 4 m, inamyloid, indextrinoid, strongly cyanophilous. This is a typical species of Cristinia and is superficially similar to C. artheniensis Baici & Hjortst. which also lacks clamp-connexions. The latter species, however, has a grandinioid hymenophore and narrower hyphae. 407 3. CRISTINIA COPROPHILA (Wakef.) Hjortst. nov. comb. Basionym: Corticium coprophilum Wakef. in Trans. Br. mycol. Soc. 6:480 (1916). Holotype: England. On dung, beds near Temp. House, Kew, 5 July, 1912, Wakefield (K)! This species is characterized by a nearly smooth and pale yellow hymenophore with hyphal strands in the periphery. The basal hyphae are without clamp-connexions, but the sub- hymenial ones regularly bearing clamps. The basidia are about 20-25 x 5.5—6.5 m, more or less subclavate and with a distinct cyanophilous granulation. All basidia are provided with a basal clamp-connexion and the spores are 4—5 im diam., thick-walled and distinctly cyanophilous. The species was reported by Jiilich (1972 and 1984) to be distributed in England (type-locality), Germany, The Netherlands, Sweden and also in Canada and USA. Except for the type no other specimens have been examined. The specimen from Sweden (Smaland, Almesakra) represents Byssocorticium lutescens Erikss. & Ryv. and is the holotype of this species and preserved in TRTC! For illustration of Byssocorticium coprophilum and comments of B. lutescens see Eriksson and Ryvarden (1973). Jiilich (1981) described the genus Dacryobasidium, with Corticium coprophilum Wakef. as type besides Byssocorticium lutescens Erikss. & Ryv. The latter lacks the basidial cyanophilous granulation, and the morphology of the basidia as well as that of the spores is more reminiscent of Hypochniciellum Hjortst. & Ryv. Also Amphinema Karst. and Piloderma Jiilich should be considered, as the basidia are basally narrowed and more or less pedunculate. The species is, however, at present maintained in Byssocorticium. Con- sequently, the genus Dacryobasidium Jiilich is here considered as a synonym of Cristinia Parm. 4. CRISTINIA GALLICA (Pilat) Jiilich in Persoonia 8:298 (1975) Radulum gallicum Pilat in Mykologia 2:54 (1925); Cristinia mucida Erikss.& Ryv. nom.nov. Corticiaceae North Eur. 3:311 (1975). Radulum mucidum (Pers.) sensu Bourd. & Galz.in Bull. trimest. Soc. mycol. Fr. 30:247—248 (1914); non Hydnum mucidum Pers.:Fr. Syst. Mycol. 1:418 (1821) = Dentipellis fragilis (Pers.:Fr.) Donk; non Hydnum mucidum Pers. Mycol. Europ. 2:179 (1825) = Trechispora farinacea (Pers.:Fr.) Liberta. For illustration and description (sub Cristinia mucida) see Eriksson and Ryvarden (1975). See further Jiilich (op.cit.) for a nomenclatural discussion about Cristinia mucida Erikss.& Ryv. 5. CRISTINIA HELVETICA (Pers.) Parm. in Consp. syst. corticiacearum p. 48 (1968) Hydnum helveticum Pers. Mycol. Eur. 2:184 (1825). For description and illustration of the species see Eriksson and Ryvarden (op.cit.). 6. CRISTINIA RHENANA Grosse-Brauckmann nov. sp. fig. 1 Basidioma resupinatum, laxe adherens, cremeum. Hymenium leve; margine fibrilloso cum rhizomorphis subtilibus. Systema hyphale monomiticum, hyphae subhymeniales omnes fibulatae, tenuiter tunicatae, 3.5—-5 tm latitudine. Hyphae basales fibulatae vel efibulatae, latitudine 4-7(—9) m. Cystidia nulla. Basidia subclavata vel subcylindrata, (20—)25—35(—S0) x (6—)6.5—7.5 m, 4 sterigmatibus. Protoplasma basidiorum cum guttulis oleosis cyanophilis. 408 Fig. 1. Cristinia rhenana. a) section through part of the basidiome b) basidia c) young basidia with cyanophilous granulation d) spores. — From the holotype. 409 Sporae globosae vel obovatae, nonnumquam leviter angulatae, leves, cyanophilae, neque amyloideae, neque dextrinoideae, incrassate tunicatae, diametro (4—)5—6.5(—7) m. Holotypus: Germany, Hessen, Oberrheinebene, Nature Reserve “Kiihkopf’, on Populus nigra, 8 Jan. 1988, H. Grosse-Brauckmann 3696 (GB). Paratypi: Germany, Hessen, Oberrheinebene, Nature Reserve “Kiihkopf’, on Populus nigra, 31 Dec. 1988, H. Grosse-Brauckmann 4351; Rheinland-Pfalz, Oberrheinebene near Mann- heim, ““Maudacher Bruch”, on Acer, 28 Dec. 1987, leg. U. Sauter. Duplicate H. Grosse- Brauckmann 4371. All specimens in Grosse-Brauckmann’s priv. Herb. BASIDIOME resupinate, effused, loosely attached to the substratum, soft, when fresh whitish grey, in dried state cream-coloured, with thin hyphal strands in the subiculum and in the periphery, margin indistinctly fibrillose. HYMENOPHORE at first reticulate, then continuous and smooth, in KOH normally with a violet tint. HYPHAL SYSTEM monomitic; subhymenial hyphae indistinctly short-celled, thin-walled, with clamp-connexions at all septa, 3.5-5 um wide; subicular hyphae somewhat thick- walled and with scattered, mostly large clamp-connexions, 4—7(—9) um wide. CYSTIDIA absent. BASIDIA subclavate to subcylindrical, (20—)25—35(—50) x (6—)6.5—7.5 m, with oily contents in the protoplasm and normally with four sterigmata, with a basal clamp-connexion. Immature basidia with cyanophilous granulation. SPORES globose to ovoid, now and then somewhat angular, with a small, but distinct apiculus, smooth, thick-walled, inamyloid, indextrinoid, but distinctly cyanophilous, mostly uniguttulate, (4-)5—6.5(—7) um diam. The species is characterized by an almost smooth basidiome and scattered clamp-con- nexions on the subicular hyphae. Further, the violet tint with KOH is distinctive. SPECIES EXCLUDENDAE Cristinia filia (Bres.) Liberta in Can. Journ. Bot. 51:1891 (1973); Corticium filium Bres. in Annls mycol. Berlin, 6:43 (1908). Lectotype: France, Bourdot no. 6984 (P), designated by Liberta (op. cit.). This is not a species of Cristinia because of the acyanophilous basidial contents and the - dextrinoid spores. Nevertheless, we have no suggestion as to its generic position. Cristinia sonorae Nakas. & Gilb. in Mycologia 70:271—272 (1978). The holotype, (USA, Arizona, KKN 237), is mixed and comprises two different species, but neither belonging in Cristinia. One part is a species of Hyphodontia (subsectio Ellipsosporae Parm.). This portion of the holotype fits the diagnosis of C. sonorae by presence of cystidia and the size of spores, but neither hyphae nor spores are cyanophilous. The second part seems to be a species of Radulodon Ryv. and is somewhat reminiscent of R. americanus Ryv. ACKNOWLEDGEMENTS We wish to express our appreciation to Dr. Roy Watling, Royal Botanic Garden, Edinburgh who revised the English language and to Prof. Leif Ryvarden, University of Oslo for reviewing the manuscript. Additional thanks to Dr. Karl-Henrik Larsson, University of Géteborg for editing and formatting the final manuscript. 410 REFERENCES Baici, A. and Hjortstam, K. 1984. A new species of Cristinia (Aphyllophorales, Corticiaceae) from North Italy. Sydowia Ann. Mycol. 37:1 1-14. Eriksson, J. and Ryvarden, L. 1973. The Corticiaceae of North Europe, Vol. 2. Aleurodiscus — Confertobasidium. Fungiflora. Oslo. __ 1975. The Corticiaceae of North Europe, Vol. 3. Coronicium — Hyphoderma. Fungiflora. Osio. Jiilich, W. 1972. Monographie der Athelieae (Corticiaceae, Basidiomycetes). Willdenowia, Beih. 7:1-283. — 1981. Higher Taxa of Basidiomycetes. Bibliotheca Mycologica 85, 485 pp. __ 1984. Die Nichtblitterpilze, Gallertpilze und Bauchpilze (Aphyllophorales, Heterobasidiomycetes, Gastromycetes). Kleine Kryptogamenflora Band II b/1. Stuttgart New York. MY COTAXON Volume XLVI, pp. 411-413 April-June 1993 COMMENTS ON RECENT WORK ON OPHIOSTOMA AND ITS SYNNEMATOUS ANAMORPHS H. P. Upadhyay Departamento de Micologia, Universidade Federal de Pernambuco 50739 Recife, Pernambuco, Brasil The following is my reaction to two recent publications on the anamorphs of Ophiostoma: Wingfield, M. J., Kendrick, B. and van Wyck, P. S. "Analysis of conidium ontogeny in anamorphs of Ophiostoma." Mycological Research 95: 1328- 1333, 1991, and Mouton, M., Wingfield, M. J. and van Wyck, P. S. "Conidium development in the synnematous anamorphs of Ophiostoma." Mycotaxon 46: 371-379, 1993. As I understand, the objective of these contributions were to study the type species of Pesotum Crane & Sckoknecht and of Phialographium Upadhyay & Kendrick _ developmentally in order to compare their conidiogenesis with that which occurs in the type species of Graphium Corda. I am satisfied that the analysis that the authors carried out was appropriate to answer the specific question that the authors posed. However, I feel that this question itself is not the most important to raise regarding the taxonomic value of percurrently proliferating conidiogenous cells in three genera, since it is now apparent that the conidiogenous cell growth can no longer be viewed as compartmentalized (Upadhyay, 1981; Minter et a/., 1983; Minter, 1987). Although the authors entitled their paper "Analysis of conidium ontogeny in anamorphs of Ophiostoma: Pesotum and Phialographium are synonyms of Graphium," they delineated, with some justification only two species of Pesotum, i.e. P. ulmi (Schw.) Crane & Schoknecht and P. piceae Crane & Schoknecht as assigned by Crane and Schoknecht (1973) vs. Graphium, but failed to examine the type species of Phialographium. The rest of their discussions and decisions with respect to other anamorphs of Ophiostoma were apparently based on scrutiny of the literature without examination of types or authentic specimens. In my, A monograph of Ceratocystis and Ceratocystiopsis (Upadhyay, 1981, p. 61), I cited a number of cultures and specimens of the Phialographium state of Ceratocystis sagmatospora Wright & Cain (=P. sagmatosporae Upadhyay & Kendrick) which they claimed could not be obtained for EM studies. In my studies, these cultures of the type species of Phialographium produced anamorphic and teleomorphic states, including conidiogenous cells with distinct collarettes. I wish also to mention that the Hyalopesotum Upadhyay & Kendrick, Pesotum, Pachnodium Upadhyay & Kendrick and Verticicladiella Hughes anamorphs of Ceratocystis and Ceratocystiopsis, in my observations under phase contrast photomicroscope, were commonly accompanied by a _ Sporothrix-like morphogenesis in pure culture, either consistently or under certain conditions. Such morphogenesis was never observed in the species of the form-genera Graphilbwm 412 Upadhyay & Kendrick, Leptographium Lagerberg & Melin, Phialocephala Kendrick, or Phialographium that I examined. Ceratocystis ips (Rumb.) C. Moreau produced a Graphilbum anamorph accompanied with Bainieria-like morphogenesis in most of the cultures examined. On the other hand, synnematous and penicillately branched mononematous conidiophores often lose fertility in artificial cultures to such an extent that they may produce conidia only on simple conidiophores similar to those of Sporothrix Hektoen & Perkins (Pesotwm-like and Verticicladiella-like) and Bainieria Arnaud. These monomematous synanamorphs and their conidiation are important issues that also need to be considered relative to the value of anamorph-generic delimitation posed by the authors. Strangely enough, conidiogenous cells of Pesotum- like and Verticicladiella-like anamorphs only can be discerned as_ proliferating percurrently by the use of an electron microscope, which may not be feasible for every worker for quick identification of species of these fungi. Also, I would like to note that the first of these two articles contains two erroneous citations: On page 1331 paragraph 2, the authors state, "Upadhyay and Kendrick (1975) suggested that the conidiogenesis in the synnematous anamorph of Ophiostoma sparsum (Davidson) de Hoog & Scheffer was phialidic, and established the segregate anamorph-genus Graphilbum Upadhyay & Kendrick as the hyaline analogue of Phialographium'"; whereas, Upadhyay and Kendrick (1975) clearly characterized and delineated the genus Graphilbum as having percurrent, annellidic conidiogenous cells (Mycologia 67: 800) and noted that this genus is a hyaline analogue of Graphium (Mycologia 67: 799). Also, on page 1332, on the penultimate line, the authors miscited Ceratocystis davidsonii Olchowecki & Reid, Can. J. Bot. 52: 1698 (1974) as "Ophiostoma davidsonii Olchowecki & Reid." One reason that I feel that these issues need to be raised is that the use of conidial ontogeny in delimitation of anamorphic genera is under intense scrtuiny (Minter ef al. 1982, 1983, 1983, Minter, 1987). In light of present controversy surrounding the use of conidial ontogeny in taxonomy of the ophiostomataceous fungi and their anamorphic genera, the developmental study of types or authentic specimens, preferably by comparing a combination of features of all closely related anamorphs and teleomorphs, is necessary in order to understand and delineate the holomorph more broadly than narrower definitions favored by Wingfield et al. For more detail, see Upadhyay (1981, 1992) Dr. Richard A. Humber, USDA-ARS, Ithaca, NY, critically reviewed that manuscript, for which I am grateful. Dr. Leonor C. Maia of this Department kindly read the manuscript. LITERATURE CITED Crane, J. L. and J. D. Schoknecht. 1973. Conidiogenesis in Ceratocystis ulmi, Ceratocystis piceae, and Graphium penicilloides. Am. J. Bot. 60: 346-354. Minter, D. W., P. M. Kirk and B. C. Sutton. 1982. Holoblastic phialides. Trans. Br. Mycol. Soc. 79: 75-93. Minter, D. W., P. M. Kirk and B. C. Sutton. 1983. Thallic phialides. Trans. Br. Mycol. Soc. 80: 36-66. 413 Minter, D. W., B. C. Sutton and B. L. Brady. 1983. What are phialides anyway? Trans. Br. Mycol. Soc. 81: 109-120. Minter, D. W. 1987. The significance of conidiogenesis in pleoanamorphy. Pages 241-262 in: Pleomorphic fungi. J. Sugiyama, ed. Kodansha, Tokyo and Elsevier, Amsterdam. Upadhyay, H. P. and W. B. Kendrick. 1975. Prodromus for a revision of Ceratocystis (Microascales, Ascomycetes) and its conidial states. Mycologia 67: 798-805. Upadhyay, H. P. 1981. A monograph of Ceratocystis and Ceratocystiopsis. Univ. Georgia Press: Athens, GA. Upadhyay, H. P. 1993. Classification of the Ophiostomatalean Fungi. In: Ceratocystis and Ophiostoma: Biology, Taxonomy, Ecology and Pathology. M. J. Wingfield, K. A. Seifert and J. Weber, eds. American Phytopathological Society Press. (in press). 7 me > ™ bg > i a VW wily i i) ' ‘ wn t e r J | i phat i a are bias ; Rey dias at .3 Wain iy Garin ty \ nan ai twat <= SS he, RSS! Yate, y ES Wy) WD, y SS \\ J iG i OLE ONC 4, Cea Ny Y ) 0o Dae AN ————— AS) SY Xd a 09h SSG SEAS EEE FSS S > aay BE Oa ° QF A a “ — Sine IS Re soa | I eae ESEOSST SS Se” 421 Fig. 3. Herbampulla crassirostris. Longitudinal section of peri- thecium (Holotypus, lactophenol permanent slide). Bar SO um. 422 brown cells covered with dark brown hyphae, and a thicker, well distinguishable, + colourless inner layer of slightly larger, more flattened, guttulate cells. On the inside the cell walls are thin, towards the outside they gradually become thicker, and the lumen smaller. Ostiolar channel lined with hyaline, partly guttulate periphyses which are often forked at the base and embedded in a gelatinous matrix, diverging inward from a textura oblita in the wall of the neck. Paraphyses filiform, 2-5 um thick, + degenerated in mature perithecia. Asci eight-spored, fusiform, 80-129 x 10-12(-15) um, very numerous, apical ring ca. 2.8 x 2.8 um, staining deeply in a solution of Congo red. Croziers not seen. Ascospores fusiform, 30-44(-48) x 4-6(-7.5) um, with slightly rounded ends, 5-septate, yellowish brown with paler end cells, guttulate (one larger oil drop in each cell). At first, the monotypic genus Stioclettia DENNIS. (1975: 363, with St. luzulina DENNIS) seemed to be quite appropriate for this peculiar fungus. But a comparison of our taxon with the type material in K proved that there are some important differences: (1) The ascus bases do not seem to dissolve before maturity, which is also suggested by the narrowly cylindrical shape of the asci with an uniseriate arrangement of ascospores. (2) The apical apparatus as figured by DENNIS (1975) and visible in the permanent microscope slides in K has a very different shape which reminds of a large Phomatospora. (3) The ascospores are apparently colourless, not pigmented with paler end cells. (4) The stromatic structures around the perithecia are probably more prominent in Stioclettia, but the type material is rather poor and the permanent microscope slides of longitudinal sections of the peridium in K do not allow any other statement. It would not have been appropriate to assign our new fungus to the wood-inhabiting, stromatic genus Melogramma FR. either, although there is some similarity in spore structure, and probably also in the apical apparatus of the asci and the development of the hymenium. Therefore Herbampulla is tentatively placed in the Melogrammataceae. Herbampulla crassirostris and Cainia graminis (NIESSL) v. ARX & MULLER apparently prefer similar habitats and hosts. Both fungi occur on Carex firma and Sesleria varia in the Caricetum firmae, but also in the Seslerio-Semper- viretum on Sesleria varia, Cainia graminis also on Helictotrichon parlatorei (WOODS) PILGER. Compared to other broad-leaved Gramineae and Cyperaceae of the Alps, these plants have rather xeromorphic leaves and prefer relatively well-drained localities - which also seems to be true for the small pyreno- mycetes fruiting on their dead leaves in the early stages of decay. Fig. 4. Herbampulla crassirostris: (A) Diagrammatic habit sketch of perithecia in the substrate. (B) Asci and paraphyses. (C) Ascospores (Holotypus, lacto- phenol permanent slides). (D) Ascospores (08.VIII.1982, Ch. SCHEUER; herba- rium material in water). B-D: Bar = 20 um. 424 Acknowledgments Thanks are due to Dr. Dietmar KORES for drawing our attention to Herbampulla crassirostris, to Dr. Sergio CHIESA (Padova) for arranging the loan of the type material of Leptosphaeria pachycarpa, to Prof. Dr. Lennart HOLM and PD. Dr. Orlando PETRINI for acting as pre-submission reviewers. References BERLESE, A.N. (1894). Icones Fungorum omnium hucusque cognitorum. Vol. I. - Avellino. DENNIS, R.W.G. (1975). New or interesting British microfungi. III. - Kew Bulletin 30: 345-365. NOGRASEK, A. (1990). Ascomyceten auf GefaBpflanzen der Polsterseggen- rasen in den Ostalpen. - Bibliotheca Mycologica 133: 1-271 + 5 pl. - Berlin, Stuttgart: J. Cramer. SACCARDO, P.A. (1891). Sylloge Fungorum omnium hucusque cognitorum. IX. Supplementum universale. - Padova. SHOEMAKER, R.A. & C.E. BABCOCK (1989). Phaeosphaeria. - Canadian Journal of Botany 67: 1500-1599. MY COTAXON Volume XLVII, pp. 425-431 April-June 1993 NEW OR INTERESTING LICHENICOLOUS FUNGI. 3.* KARSTENIOMYCES LLIMONAE SP. NOV. AND SCLEROCOCCUM SERUSIAUXTI SP. NOV. (DEUTEROMYCOTINA) Montserrat BOQUERAS Dept. Biologia Vegetal (Botanica), Fac. Biologia Univ. Barcelona Diagonal 645, E-08071 Barcelona, Spain Paul DIEDERICH Musée national d'histoire naturelle Marché-aux-Poissons, L-2345 Luxembourg, G. D. Luxembourg Abstract: Karsteniomyces llimonae Boqueras & Diederich sp. nov. (Coelomycetes) is distinguished from related species by setose pycnidia, much longer conidiophores and a different host (Parmelina quercina); it is known only from the type locality in Spain (Catalonia). Sclerococcum serusiauxii Boqueras & Diederich sp. nov. (Hyphomycetes) is distinguished by very small sporodochia, well-delimited thickened and darker regions of the conidial wall, and the host (Parmelina pastillifera and P. tiliacea); it is known from Spain (Navarra and Catalonia). During an excursion in south Catalonia (Spain) in 1991, the authors collected an appearently undescribed species of Karsteniomyces growing on Parmelina quercina. A new species of Sclerococcum, growing also on species of Parmelina, has been discovered by the senior author in Catalonia, and by Dr E. Sérusiaux and Dr J. Etayo in Navarra (Spain). Both species are described below. * 2. In: Bull. Soc. Nat. luxemb. 93: 155-162 (1992). 426 Karsteniomyces llimonae Boqueras & Diederich sp. nov. Figs. (1-2). Fungus lichenicola. Conidiomata pycnidia, singularia, dispersa, subglobosa superficialia, ostiolata, straminea ad cinnamomea, 60-120 {zm diam., setis hyalinis, non-septatis, 17-27 x 4-8 tum. Conidiophora elongata, simplicia ad sparse ramosa, sparse septata, hyalina, 30-40 x 0.7-1 wm. Cellulae conidiogenae holoblasticae, monoblasticae. Conidia sicca, elongato-ellipsoidea, hyalina, 1-septata, levia, 13-17 x 2.5-3.5 um. Typus: Spain, Catalonia, Montsianés, La Sénia, Barranc del Retaule (Serra des Ports), BF6914-BF6915, 1000-1100 m, on Corylus avellana, on Parmelina quercina, 10 11 1991, M. Boqueras, P. Diederich 9845 & A. Gémez-Bolea (BCC-Lich. 5224 - holotypus; herb. Diederich - isotypus). Conidiomata pycnidial, arising singly, scattered or loosely aggregated, subglobose, superficial, with only the base entering the host tissues, ostiolate, pale yellow orange to dark red orange, 60-120 {1m diam., with setae near the ostiole; setae non-septate, hyaline, with a rounded apex, often constricted in the middle, 17-27 x 4- 8 um; pycnidial wall thin, composed of pseudosclerenchymatous cells, 15-25 tm thick. Conidiophores arising from the inner pycnidial wall, cylindrical, elongate, sometimes branched or septate, hyaline, 30-40 tum in length, 0.7-1 um thick. Conidiogenous cells arising terminally and laterally from the conidiophores, holoblastic, monoblastic, indistinguishable from the conidiophores except in the absence of septa. Conidia dry, elongate-ellipsoid, with rounded ends or slightly truncate at the base, hyaline, 1-septate, smooth-walled, with slightly unequal cells, 13-17 x 2.5-3.5 um. Etymology: The species is named in honour of Dr Xavier Llimona (Barcelona, Catalonia, Spain) for his enthusiasm and his contribution to the mycology (including lichenology) of Catalonia. Host: Parmelina quercina. The fungus acts as a parasite producing poorly delimited decolorized patches on the host thallus. Distribution: Known only from the type locality in south Catalonia (Spain). Observations: The new fungus is easily distinguished from the two previously known species of Karsteniomyces, K. peltigerae (P. Karsten) D. Hawksw. and K. tuberculosus Alstrup & D. Hawksw., by the presence of hyaline setae near the ostiole, by the much longer conidiophores, and by the different host. The two other species of Karsteniomyces grow on Peltigera and both represent the anamorph of a species of Scutula, S. miliaris (Wallr.) Trevis. and S. aggregata (Bagl. & Car.) Rehm respectively. 427 "9819S “D “eIpluod pue saloydorpruog ‘g ‘umIprusAd y3nonp uonseg ‘y :(sndAjost - youapaiq “qiay) apuowny sarkwouassuvy - “| “31 url QO] foe ae aE Sa TAD Hy iyiit —— > xy i ae fo His — i a f m ‘ e ny ie 2 pe) a : : pS és ‘ U Yon as Whe ? . yi on oh “* un Pe Y we = ila ' wg “ f day!’ ree | pi! ha) z hie vio 74 ie <. ; dares Rey hind ie er ny, cory hae Nees te hail i . an A Mi . 7 heey: uv . % ey me fi . Lea "ys Al x MY “pane aoe be oe aa Mae adh coe a ethhy oi i £) oN We ty GP: Wy tas an) Rig tne a 4 1 Ve ot ae 7 res: fy Be ee ily A th kad | ast rd ath tn ah Ow re nite ede WOE fis Wh af aio nj Histor: ut pet ines es Nel 10) om i fame ah Pat ha Po) oe ty We pean hs any iv ene 1A , iM) wm, 7 ny de a ca 8 fe | ARN et Ue alls et a ee ea esr ae i ar Ae 20 tae Faye Wy AL ate f ae < ee us Terie aan MY COTAXON Volume XLVI, pp. 433-437 April-June 1993 AMANITA REIDII - A NEW SPECIES FROM SOUTH AFRICA ALBERT EICKER, J.V. VAN GREUNING AND DEREK A. REID Department of Botany, University of Pretoria, Hillcrest, Pretoria 0002, South Africa SUMMARY Amanita reidii is described and compared with the closely related A. phal- loides (Fr.) Link Since the publication of the recent monograph of South African species of Amanita (Reid & Eicker, 1991) a further undescribed species has been found growing in open stands of Eucalyptus. This new taxon is recognized by its grey-brown cap, conspicuously streaked with darker radial markings and by its saccate volva. A description of the new taxon follows: Amanita reidii Eicker & Greuning, sp. nov. (Figs 1-8) Etym.: reidii - After Dr Derek A. Reid, for his significant contribution to South African mycology Pileus 3.0-7.2 cm diam., convexus vel applanatus, vel leviter campanulatus, fusco-griseo-brunneus, centrum alibique versus pallide maculatescens; marginem versus lineis conspicuis radialibus atrofuscis ornatus; superficies pilei glabra, sine reliquiis veli; margo pilei non striatus. Lamellae adnexae, fere liberae, albae. Stipes 3-7 cm altus, ad apicem 0.5-1.5 cm. diam., aequalis vel clavatus, basi usque 2.2 cm diam., albus vel sordide albus, annulo albo apicali membranaceo ornatus; infra annulum superficie in squamas minutas fractiflexas disrupta. Volva libera, alba, cylindrica ad stipitem arcte posita. Structura volvae hyphis angustis, usque 4 pm diam., efibulatis composita; nonnullis segmentis harum hypharum elongatis, inflatis, usque 200 ym longis et 35 wm latis. Cheilocystidia 20-36 X 10-16 um, clavata vel ovata parietibus tenuibus. Basidia 36-50 X 7-11 wm hyalina, clavata, quadrispora parietibus tenuibus. Sporae subglobosae vel ovatae 7.5-8.0 X 6.0- 7.2, vel ovato-ellipticae 9.0-10.0 X 6.5-8.0 um, amyloidea, parietibus tenuibus; sporae ex pulvere sporarum 8.0-9.5 X 6.2-9.0 wm metientes. Holotypus sub Eucalyptus, Hideaway Game Farm, Melkrivier, prope Pretoria, 29 April 1990; V. van Greuning & D.A. Reid. [in PRUM]. Pileus 3.0-7.2 cm convex or shallowly campanulate, becoming shallowly convex or flattened, blotchily grey brown with an innately radially streaky surface. The colour may vary to darker grey-brown or blackish slate-grey toward the centre and become paler toward the margin, although the pileus often develops pallid dirty white blotches at the disc or elsewhere. The streakiness is most conspicuous toward the outer portion of the cap where the radial, dark grey-brown streaks con- trast more sharply against the pale ground colour. __ Pileus lacking all traces of velar tissue. Stipe 3-7 cm high, 0.5-1.5 cm wide at the equal or slightly enlarged 434 O CoV OOD 3 Figs 1-7. Amanita reidii. 1. Spores from print mounted in Melzer's solution. 2. Spores from gill mounted in Melzer's solution. 3. Basidia. 4. Elements from universal veil. All from holotype. 5. Basidia. 6. Cheilocystidia. 7. Spores from print mounted in NH, with congo red. All from Belfast collection. Bar = 10 pm. 2 apex, and up to 2.2 cm at the clavate, nonbulbous base, white to dirty white, the surface with fibrillose scaly zig-zag markings below the annulus. Annulus apical, pendulous, white, membranous. Volva white, saccate, and although free from the stipe, sheathing it rather closely. | Lamellae adnexed to almost free, white to creamy-white. Universal Velar tissue lacking from surface of pileus; structure of volval limb hyphal, comprising narrow hyaline hyphae with a variable proportion of very long inflated segments, but devoid of sphaerocysts; Occasional conducting (gloeoplerous) hyphae present in some collections. The narrow hyphae, to 4 nm wide, lack clamp connexions at the septa, and have thin but rather distinct walls. The inflated segments, often fusoid, are usually of an intercalary position, up to 200 um long, and up to 35 wm wide. Partial veil hyphal. Cheilocystidia 20-36 X 10-16 wm thin-walled, hyaline clavate or ovate. In the Bergvliet material the gill edge is formed of a thick layer of dissociating chains of thin-wailed, hyaline, globose or ovate elements up to 30 um long and to 23 um wide. Basidia 36-50 X 7-11 pm, 4-spored, thin-walled. hyaline, clavate, originating from rounded or 435 wedge-shaped segments, but lacking a basal clamp-connexion. Spores from print: 8.0-9.5 X 6.2-9.0 wm (measured in Melzer's solution), varying subglobose to ovate, more rarely ovate-elliptic, thin-walled, hyaline, amyloid. Spores taken from the gill more variable in size and shape : measuring 7.0-8.0 X 6.0-7.2 ym when subglobose or ovate, but 9.0-10.0 X 6.5-8.0 um when ovate-elliptic. Distribution: under Eucalyptus claeziana, Bergvliet State Forest, Sabie, G.C.A. van der Westhuizen (35), 7 Dec. 1984 [PREM 48618]; under Eucalyptus, Hideaway Game Farm, Melkrivier, V. van Greuning & D.A. Reid, 29 April 1990 [PRUM 3179] [Holotype]; in open situation under Eucalyptus, Pineglades, Bel- fast, 8 Feb. 1991. These gatherings are deposited in PRUM (Mycology Section of the H.G.W.J. Schweickerdt herbarium, PRU, University of Pretoria) or PREM. Edibility nothing is known regarding the edibility of this species, but be- cause of a suggested relationship with A. phalloides it should be treated with very great caution and avoided. DISCUSSION A. reidii is undoubtedly closely related to A. phalloides (Fr.) Link, differ- ing in its prominently streaky grey-brown pileus, without trace of green tints. By contrast the pileus of A. phalloides is yellow-green to olive-green, ornamented with fine, innate, rather densely crowded and slightly darker radial streaks. In A. reidii the streaks are fewer but darker and altogether more pronounced. The white stipe of A. reidii also differs from that of A. phalloides, which is often greenish, in that it is not conspicuously bulbous as in the latter species, and is rather closely sheathed by the volva. Again A. reidii appears to be of slightly smaller stature. Further the known collections of A. reidii have all comprised fruitbodies growing in the vicinity of Eucalyptus trees, and it may be that this fungus will ultimately be shown to require a more specialised habitat than does A. phalloides. In South Africa there has been a trend toward naming the fungus described above as A. phalloides f. umbrina Ferry - at least on field meetings. However the name has not yet appeared in the South African literature. Through the kindness of Dr. Jan Kuthan and Dr. Herink we have received copies of Ferry's original description and comments on this taxon, from which it is clear that he regarded his forma uwmbrina as representing the ultimate stage of ageing of the familiar yellow, greenish or olive fungus. As such it does not merit taxonomic recognition. Discussing the variation of A. phalloides, Ferry (1911) wrote "A ces diverses variétés on pourrait ajouter une forme umbrina ....... Ce n'est pas une variété; les diverses variétés jaune, verdatre ou olive peuvent par- venir a ce stade, qui représenté une état plus avancé : cette teinte bistre survient avant qu'il y ait la moindre trace d'altérative ou ni la moindre odeur." Since we have observed specimens of the fungus under discussion in all stages of development at the Melkrivier locality without finding any evidence of green tints in the young fruitbodies, we feel confident that the South African material represents an autonomous taxon distinct from both Amanita phalloides f. phalloides and A. phalloides f. umbrina. We have, therefore, decided to publish it at specific rank under the name A. reidii. Reference to the publications of Bottomley and Talbot (1954) and van der Westhuizen (1983) suggest that they were aware of this brown to grey brown Amanita, but that they included it in their concept of A. phalloides (Fr.) Link. Thus Bottomley & Talbot (1.c.) in their description of the latter referred to it as having the cap "smooth or streaked with dark innate fibrils and to be variable in colour from "greenish, yellowish olive, greyish brown, umber brown to smoky olive, or even darker ......... ". Indeed their italics suggest that umber brown to smoky olive was the more commonly met with colour form, and further suggests 436 that it was A. reidii rather than the true A. phalloides which was the more familiar species to these authors - at least in the Transvaal. Van der Westhuizen (1983) also noted that A. phalloides, in South Africa, occurred in various shades of greenish yellow olive, smoky brownish-green, umber brown to smoky olive, darker towards the centre." This description again appears to have embraced A. reidii. Having considered the taxonomic relationship between A. reidii and the other species of Amanita known to occur in South Africa it is necessary to consider this relationship in a wider context. It would seem that the South African species is most closely related to A. alauda Corner & Bas (1962), which was described from the Gardens Jungle, Botanic Gardens, Singapore, based on a single collection comprising two half specimens. This species belongs in Amanita, subgenus Lepidella, section Phalloideae, along with both A. phalloides and A. reidii. Fig. 8. Amanita reidii. Two fruitbodies from Melkrivier collection x 1. A. alauda is very similar to A. reidii in having a brownish cap devoid of green colouration, varying from sepia or pale purplish umber to mouse-coloured, darker at the centre. More importantly the cap is "streaked by innate dark fibrils, especially near margin". However the stature of A. alauda differs from that of A. reidii in that whereas the latter tends to be rather short and squat with the length of the stem about equal to the diameter of the cap, A. alauda is taller and more elegant in appearance with the length of the narrow stem approaching twice the diameter of the cap. Further the cap of A. alauda has a distinct umbo seated in a central depres- sion; in contrast the cap of A. reidii is either shallowly campanulate with a flat- tened, sometimes almost truncate disc, or applanate;. umbonate sporophores have not been observed in this species. Another difference is that the cap of A. alauda is more distinctly brown in colour, and the radial streakiness is less complete and less obviously reticulate. Further the cap surface of A. alauda was described as 437 viscid, a feature emphasized in the authors discussion of their new taxon. A vis- cid cap cuticle has not been observed in A. reidii. Again the surface of the stem of A. alaudia was said to be smooth; in A. reidii it disrupts into a zig-zag pattern of fibrillose markings, but this may not be of great taxonomic significance. Spore size and shape are very similar in both taxa as are the basidial features. Cheilocystidia are also similar although these organs are very slightly broader in A. alauda. However comparison of the account of the microstructure of the volval limb of A. alauda with that of A. reidii suggests that there are differences in this respect. Corner & Bas (1962) wrote "Limb of volva mainly consisting of 3-5(-25) pm wide, interwoven sublongitudinal hyphae, with some scarce sphaerocysts up to 100 um wide in the inner part". There is no mention of the occurrence of very long, rather abruptly inflated fusoid hyphal segments such as those found in the volval tissue of A. reidii. Finally there is the difference in habitat to be con- sidered for while A. reidii appears to grow in association with Eucalyptus, which is suggestive of a mycorrhizal relationship, A. alauda was said to grow in jungle with the implication that it grew amongst native vegetation. Finally since A. reidii has, to date, been found only in areas strongly in- fluenced by human activity, it seems that this fungus will prove to be an exotic species in South Africa. If so one might look to Australasia as its possible geographical origin. ACKNOWLEDGEMENTS The authors are grateful to the South African Foundation for Research Development for financial support. We thank the Director General, Department of Water Affairs and Forestry for permission to collect fungi on their property. REFERENCES Bottomley, A.M. & Talbot, P.H.B. (1954). Common Edible and Poisonous Mushrooms in South Africa. Bulletin 324, Botany and Plant Pathology Series No.17, Department of Agriculture, Union of South Africa. Corer, E.J.H. & Bas, C. (1962). The genus Amanita in Singapore and Malaya. Persoonia 2: 241-304. Ferry, R. (1911). Etudes sur les Amanites. A. phalloides, A. verna, A.virosa, Revue mycologique. Toulouse. Suppl. 1. Reid, D.A. & Eicker, A. (1991). South African fungi : the genus Amanita. Mycological Research 95, 80-95. Van der Westhuizen, G.C.A. (1983). Mushrooms and Toadstools. Bulletin 396. Plant Protection Research Institute, Department of Agriculture, South Africa. oe Pe rn ee ae ae, oe Dy - } Me >> el ’ m > ae tS ; as WA A f oh “din fy ( ie i pak) 2/0 . Vapeh ety ht? ec i sit three aa a ay ps . mai v4 ; My my Ae) i J" TV a i ‘Nery “ . x Ny i, ‘i Wn wah Hn a. Pweg pa yi 4 , fice dei we “ {e Pas : (vie . yy ae y ie rs ; Na : hd y asl ; ay ‘ Ane 4) j rm ath na hy pH cet ye i ud . Bey the Manas ! Hi Sa my, MY COTAXON Volume XLVII, pp. 439-460 April-June 1993 PHYTOPHTHORA HIBERNALIS AND P. SYRINGAE H.H. Ho’ and 8.C. Jong’ ‘Department of Biology, State University of New York, New Paltz, New York 12465 *Mycology & Botany Department, American Type Culture Collection, 12301 Parklawn Drive, Rockville, Maryland 20852 ABSTRACT A comparative morphological study of isolates of Phytophthora hibernalis and PP. syringae confirmed that these are two distinct species, readily distinguished by the easily deciduous, elongated ovoid to ellipsoidal sporangia with long pedicels and the amphigynous and paragynous antheridia in varying proportions in the former and the nondecidous, broadly ovoid, obpyriform to ellipsoidal sporangia and the predominantly paragynous antheridia in the latter. Confusions in literature concerning these two species can be attributable to misidentification and/or acceptance of Tucker's treatment of P. hibernalis synonymous with P. syringae which had priority. INTRODUCTION In 1881, Berkeley first described the fungus attacking the leaves of lilac under the binomial Ovularia syringae. The pathogen was later renamed as the type of a new genus: Phloeophthora syringae Kleb. (Klebahn, 1906), then Nozemia syringae (Kleb.) Pethybr. (Pethybridge, 1913) and finally as Phytophthora syringae (Kleb.) Kleb. (Klebahn, 1909), characterized. by sympodially developed sporangiophores, large NOvoOLd, obovoid to 440 ellipsoidal" sporangium with a "flat cap at the énd" sand the ~ productiion “of oogonia: with paragynous antheridia. Despite Sarejanni's attempt (1936) to treat it as a subvariety of P. cactorum, P. syringae has been accepted widely aS a good species. In 1925, Carne reported a new species of Phytophthora, P. hibernalis on the fruit, leaves and smaller branches of Citrus spp. in Australia. It was unique in its "very deciduous, ellipsoidal or lemon shaped conidia with a constant pedicel" and the production of oogonia with amphigynous and occasionally paragynous antheridia. The fungus wasS considered so different from P. syringae that Carne did not even attempt to compare these two species when he discussed the systematic position of P. hibernalis. However, the morphological similarity between P. syringae and P. hibernalis was noted by Petri (1927) and later by Tucker (1931) who showed that these two species were identical in the production of only paragynous antheridia on all culture media, their failure to grow at 25 C, their growth characters, the production of oogonia and oospores of almost the same dimensions, and similarly small sporangia with flattened papilla, sparse on agar medium but more abundant in water. Consequently, he concluded that P. hibernalis should be treated as synonymous with P. syringae which has priority. Leonian (1934) also found that these two species were very similar both morphologically and physiologically but he preferred to keep them as separate species, an opinion shared by Waterhouse (1963) and other researchers. However, Wager (1942), Frezzi (1950)5 Noviello and Snyder (1963) and Novotelnova (1974) subscribed to Tucker's thesis that P. hibernalis and PP. syringae are conspecific. Ther di Chotomy eh bys Op Lntons concerning the taxonomic status of these two species often led to confusion in literature, especially when both have been known to attack the same host, for example, citrus during cool weathers causing identical symptoms (Klotz and Calavan, 1969). In a comprehensive study of the 441 sexual reproduction of Phytophthora species, Savage et al. (1968) pointed out the urgent need for a thorough study of isolates of P. hibernalis and P. syringae. However, most of the studies in the past centered on the pathology and ecology of P. syringae, especially on lilac (De Bruyn 1924a,b; 1928), apple, pear and related Rosaceae fruit trees (Harris, 1979, 85 a,b,c; Harris and Cole, 1982; Upstone, 1978; Upstone & Gunn, 1978; Doster and Bostock, 1988b; Young and Milbrath, 1959), sweet chestnut and beech (Day, 1938) and P. hibernalis on citrus (Favaloro and Sammarco, LO Fst LOt Ze ali 7 3) ¥ The present study was undertaken to compare the morphological characteristics of various isolates of P. hibernalis and P. syringae under uniform conditions in order to define the species more precisely. MATERIALS AND METHODS Isolates and media: Specific information on the isolates of P. hibernalis and P. syringae used is given in Table 1. With the exception of N37 which was provided through the courtesy of Dr. Robert Young, Department of Plant Pathology, West Virginia University, Morgantown, Va., all isolates were obtained from the American Type Culture Collection (ATCC), Rockville, Maryland. Unless otherwise stated, clarified V-8 juice agar medium (Ribeiro, 1978) Supplemented with sitosterol (30mg/1) (CV8) was used for culture. Morphology: Colony characteristics on CV8 were compared after incubating the cultures in darknessaeats) 20, Car foreu7ardays’. The colony diameters were measured at right angles through the inoculum, and the width of primary hyphae measured using a light microscope. Sporangia formation was induced by incubating under light at 20C small mycelial agar discs on cCV8 in freshly collected stream water that had been filtered through 0.45 um pore size membrane discs. Sex organs in single cultures were examined periodically by microsocopy through the 442 bottom of the petri dish. If the isolate failed to produce sex organs in single culture, it was then grown on Tryptophane Medium (Ribeiro, 1978) or cCV8 supplemented with 10% vegetable oil (Doster and Bostock, 1988a) and incubated at 7- SCs RESULTS Colony morphology All isolates were characterized by rather slow growth rate on cCV8 (4-8mm/day for P. hibernalis and 4-6mm/day for P. syringae), appressed to moderately fluffy, often showing distinct growth patterns, commonly rosette (with broad, rounded petalloid sectors) Les. hibernalis and chrysanthemum (with small, narrow, rounded petaloid sectors) in P. syringae. The hyphae were narrow, 3-5 um wide for P. hibernalis and 3-6 wm wide for P. syringae often irregular and wavy with frequent short branches. Small spherical to oval, monilioid hyphal swellings (under 25 um) were commonly produced by some isolates of P. syringae in water or in old cultures but not by isolates of P. hibernalis. No chlamydospores were produced by any isolate under any experimental condition. Sporangia All isolates produced sporangia readily in water within 24-48 hrs, except ATCC 60352 P. hibernalis and ATCC 34002 P. syringae. Isolates ATCC 56352 P. hibernalis and ATCC 14837 P. syringae also produced sporangia on agar medium though not as abundantly as in water. The sporangia of P. hibernalis were rather small (av. 29-53 x 14-22 um), characteristically elongated ovoid to ellipsoidal, often with tapered base and the broadest part near the semi-papillate apex (L/B av. 1.8-2.4), borne terminally on unbranched or long branches of narrow (1-2 um wide), lax sympodially branched sporangiophores and were easily deciduous, each with a long pedicel (23- 73 wm). In contrast, the sporangiophores of P. 443 syringae were either unbranched or distinctly sympodially branched bearing non-deciduous sporangia terminally on short branches often with conspicuous basal plugs and _ subsporangial swellings. The semi-papillate sporangia were in general larger (av. 41-75 x 39-40 um) with mostly rounded base and rather wide and flat apex (apical thickening 1-3 um), ovoid to ellipsoidal in water (L/B av. 1.4-2.1) but broadly ovoid to obpyriform on agar plate (L/B 1.5). No internal proliferation of sporangiophore was observed in any isolate. Nearly all sporangia released zoospores in water through single exit pore, but most sporangia of ATCC 56352 P. hibernalis germinated directly by one or more than one germ tubes through the apex whereas ATCC 14837 P. syringae occasionally formed secondary sporangia in a chain as demonstrated by Lafferty and Pethybridge (1922) Noviello and Snyder (1963) and Edney and Chambers (1981). sex Organs Most isolates produced sex organs in single cultures either readily in 2-3 wks or tardily atter} 2-—5yamontnhs. Isolates ATCC 60352 P. hibernalis and ATCC 34002 P. syringae failed to produce them under any condition. Two isolates of P. syringae, ATCC 46886 and 64876 produced sex organs sparsely only in old cultures but more rapidly and abundantly on V8C supplemented with vegetable oil. The oogonia of both species were colorless to slightly yellowish to tawnish with the single oospore nearly filling the oogonial space, and the oospore wall averaged 1-3 um thick. In general, the oogonia and oospores of P. hibernalis were slightly smaller than those of P. syringae (av. 33-35 um verses 33-41 um diam and 29-32 um versus 30-35 um diam). Whereas the antheridia were distinctly predominantly paragynous in all isolates of P. syringae, they were predominantly amphigynous in isolate ATCC 64708 but variable in isolates ATCC 32995 and 56353 of P. hibernalis. The antheridia were similarly small for both species, averaging 444 10 x 10 wm for P. hibernalis and 11 x 10 um for P. syringae. None of the isolates studied here produced chlamydospores. DISCUSSION Our study of various isolates of Phytophthora hibernalis and P. syringae confirmed Waterhouse's conclusion (1963) that these are two distinct species. There is little doubt that they have much in common, for instance, their homothallism, their low maximal temperature for growth, their narrow, wavy, irregular hyphae, small antheridia, plerotic oospores and their often rosette cultural patterns. However, P. hibernalis can be easily distinguished from P. syringae primarily by the deciduous, elongated ellipsoidal and usually smaller sporangia with a tapered base bearing a long pedicel. Most isolates of P. syringae produced well-defined sympodially branched sporangiophores with broadly ovoid to obpyriform sporangia borne on short stalks and in no instance were the sporangia deciduous. The formation of small moniliform hyphal swellings is a common feature of some P. syringae isolates. These differences were also noted by previous workers (Table 4). The antheridia of P. syringae have been described as paragynous or predominantly paragynous. Our study showed that whereas both types were present, the paragynous antheridia were undoubtedly far more common than amphigynous antheridia. On the other hand, there was much confusion regarding the antheridial configuration in P. hibernalis which has been described as amphigynous, predominantly amphigynous, paragynous or predominantly paragynous (Table 4). Gerrettson-Cornell (1989) stated that the antheridia of P. hibernalis have been reported to be predominantly amphigynous in fresh cultures but predominantly paragynous in old cultures. We have not been able to confirm this in our literature research. In fact, we noticed that in isolates ATCC 32995 and 56353 of P. hibernalis most antheridia appeared initially as paragynous but later on, amphigynous antheridia often prevailed. It is quite possible 445 that the ratio of amphigynous and paragynous antheridia produced in culture may be variable as found in P. porri (Ho, 1983) and P. megasperma (Erwin, 1965), thus explaining the discrepancies in reports regarding the antheridial types of P. hibernalis. The confusions in literature about the morphological characteristics of P. hibernalis and P. syringae can also be attributable, at least in some cases, to misidentification. Thus, Nadel-Schiffmann (1947) reported and described "Pp, hibernalis" from citrus in Palestine and this paper was listed as a major reference in the species description of P. hibernalis (Waterhouse and Waterston, 1964b). Based on the information and the photographs presented by Nadel-Schiffmann in her paper, we agree with kKouyeas) and Chitzanidis (1968) that the fungus should have been identified as P. syringae. Savage (1963) studied an orange isolate of "P. hibernalis" (N37) received from Klotz in California, and found many oogonia with amphigynous antheridia on 3-month lima-bean agar slants. Brennemann (1967) re-investigated N37 along with two other isolates of "P. hibernalis" (N116 received from CMI and N252 received from CBS) and reported that they produced paragynous antheridia on hemp-seed and oatmeal agar plates. He suspected too, that these isolates might be P. syringae, but did not attempt to study them thoroughly to determine their real identity. We have been able to obtain isolate N37 and based on the ~- sporangial characteristics, there is little doubt that it should have been assigned to P. syringae instead of P. hibernalis. Accepting Tucker's treatment of P. hibernalis to be synonymous with P. syringae which has priority, Wager (1942), Frezzi (1950) and Verneau (1953) reported "P. syringae" on citrus even though the pathogen had all or most of the features characteristic of P. hibernalis as described by Carne (1925). Wager described the antheridia of " P. syringae" as paragynous, yet his diagram of two oogonia with attached 446 antheridia showed one to be paragynous and the other, in all likelihood, amphigynous, resembling the amphigynous antheridia of P. cinnamomi and P. parasitica in the same figure. It is puzzling why Tucker (1931) considered P. hibernalis identical as P. syringae. He compared single lilac isolate of P. syringae from Baarn with single isolate of P. hibernalis "from Kew to Baarn" and found that both produced paragynous antheridia in all media, nearly identical oogonia and oospores and similar sporangia with flattened papillae. There was no mention of the caducity of sporangia in the isolate of "P. hibernalis" - - a feature seldom missed by other researchers (Table 4). It is not inconceivable that the "P. hibernalis" isolate that Tucker examined was actually P. syringae. Once these possible sources of confusions have been identified, the differences between these two species reported in literature became apparent. Thus, the sporangia of P. hibernalis have been reported as easily deciduous, ellipsoidal, averaging 37 (10-75) x 18 (8-35) um; L/B 2.1 (1.8-2.5), with long pedicels (2-80 um), closely matching our measurements in present SCUGY 39°" 408 (29-55 LP xe, POR Gl 4220 eeu Meranate4 ee ee (1.8-2-4), pedicel 23-73 um long. In contrast, P. syringae produced non-deciduous, broadly ovoid, obpyriform to elongated, larger sporangia, averaging 4%. (271475) "xs 307@ 1-42) um Bae © (12-233) Vins lLiteraturetand, 547(41=75)ex"32,(24— 40) umeetis/iB yi8 Cle a—oeiye ine Our, Study. Both species prefer low temperature for growth, with the minimum at 0-5 C and maximum 22-27 C. The antheridia of P. syringae are predominantly paragynous. Those authors who reported the antheridia simply as paragynous probably had missed the amphigynous antheridia which could be rare in some isolates. On the other hand, the antheridia of P. hibernalis can best be described as having amphigynous and paragynous antheridia in varying proportions in view of the conflicting reports in literature and based on our observations. There is little difference in the size of sex organs between these two species 447 based on published data. The oogonia, oospores and antheridia of P. hibernalis and P. syringae averaged, respectively, 23 (27-56) vs. 31 (18- 50) wm diam, 28 (20-40) vs. 28 (17-38) um diam and 11.5 (9-21) x 11.5 (10-15) vs. 10 (10-19.6) x 8 (7-10) um. It is interesting to note that in our study, the oogonia and oospores of P. syringae appeared slightly larger than those of P. hibernalis, averaging respectively 37 (33-41) vs. 34.(33-35) um diam and 33 (30-35) vs. 31 (29- 32) um diam. Our measurements of the antheridia of P. hibernalis and P. syringae showed them to bes -simplarnlyvasmalilsalOmex LOM VS 6 al ax y1O0eum. Although both species produce petalloid colonies described commonly as "rosette" many isolates of P. syringae produced "chrysanthemum" type growth patterns, with small narrow rounded sectors. Chlamydospores are absent in P. hibernalis or P. syringae. The chain-like "chlamydospores" reported by Nadel-Schiffman (1947) for "BP. hibernalis" should be interpreted as moniliod hyphal swellings for the lack of delimiting septa (Blackwell, 1949). ACKNOWLEDGEMENTS This work was supported in part by NSF Grant DIRS 9O= 15 ls ie COmoGcd:. The authors thank Elmer Davis and Dr. H.S. Chang for reviewing the manuscript. LITERATURE CITED Al-Hedarthy & Tsao 1979. Sporangium pedicel length in Phytophthora species and the consideration of its uniformity in determining sporangium caducity. Trans. Br. MV.COLMES OChee/ 2c) lia F, Barrachina-Tuset, J.J. 1977. Contribution al conocimiento del genero Phytophthora de Bary en Espana. Ann. Inst. Nacional de Invest. Agrarias Ser. Prot. Veg. 7: 11-106. Bensaude, M. 1929. Note sur le Phytophthora, parasite des Citrus au Portugal. Compt. Rend. soc. Biol. 101: 982-984. Berkeley, M.J. 1881. Lilac fungus. Gard. Chron. 448 Toei 665% Blackwell, E. 1949. Terminology in Phytophthora. Mycol. Pap. 30. Commonw. Mycol. Inst., Kew, Surrey, England. 24pp. Boccas, B. and E. Laville, 1978. Les maladies a Phytophthora, des agrumes. Inst. de Recherches sur les Fruits et Agrumes. 162 Pp. Bolay, A. 1977. Phytophthora syringae - agemt d'une grave pourriture des pommes-~ en conservation. Rev. Vitic. Arbor. Hort. 9: LoL=169.% Bostock, R.M. and M.A. Doster. 1985. Association of Phytophthora syringae with pruning wound cankers of almond trees. Pl. Dis. 69: 568- Sorel Brasier, C.M. and R.G. Strouts. 1978. New or unusual records of plant diseases and pest. Pl. Path. 27: 99-100. Brenneman, J.A. 1967. Patterns of sexuality in ten species of Phytophthora. M.S. Thesis, W.Va. Unin::, Morgantown, W: Va. 9°53) pp” Buddenhagen, I.W. 1955. Various aspects Phytophthora cactorum collar-rot of apple trees in the Netherlands. Tijdschr. Plziekt. Le ES WE 28 ed te Carne, W.M. 1925. AMeDY OWN TOC OLMmCTerus iin Australia (Phytophthora hibernalis n. sp.) J. (Roy. Soctun Wie Australivaivi2 713 —49". Chester, R.S. 1932. A comparative study of three Phytophthora diseases of Lilac and of their pathogens. J. Arnold Arboretum 13: 209-232. Day; W.R.2 1938.) *Root-rot of) sweet) chestnut and beech caused by species of Phytophthora. J. SOC VFOn.MGre (Bri Csr a1 O05. De Bruyn, H.L.G. 1924a. The Phytophthora disease of lilac. | Phytopathology 14" 503=5i7" De’, Bruyn,’ H.L.Gs 29246; De oorzaak van het epidermisch optreden van de Phytophthora ziekte van de seringen. Tijdsch. Plziekt. S30 DLS 12 24 De Bruyn, H.L.G. 1928. Is ontbladering als bestrijidinswijze tegen Phytophthora z ziekte van de seringen gewenscht? Tijydsch':?..Pr* ziekt. 34: 233-238. Doepel, R.F. 1966. Phytophthora species on 449 citrus in Western Australia. Pl. Dis. Reptr. 50: 494-496. Doster, M.A. and R.M. Bostock. 1988a. The effect of temperature and type of medium on oospore production by Phythphthora syringae. Mycologia 80: 77-81. Doster, M.A. and R.M. Bostock. 1988b. Chemical protection of almond pruning wounds from infection by Phytophthora syringae. Pl. Dis. 72: 492-494. Edney, K.L. and D.A. Chambers. 1981. Post- harvest treatments for the control of Phytophthora syringae storage rot of apples. ANN. wappLemBroley O07 si 23 7-241. Erwin, D.C. 1965. Reclassification of the causal agent of root rot of alfalfa from Phytophthora crytogea to P. megasperma. Phytopathology 55: 1139-1143. Favaloro, M. and G. Sammarco. 1973. Ricerche sul marciume del collette e radicale degli agrumi. Specie di Phytophthora presente negli agruneti della Sicilia Orientale. Phytopath. Medit. 12: 105-107. Fawcett, H.S. 1933. New locations for Phytophthora citrophthora and P. hibernalis on citrus. Phytopathology 23: 667-669. Fawcett, H.S. 1936. Citrus diseases and their control. McGraw-Hill, New York. 656 p. Frezzi, M.J. 191507. Las especies de 'Phytophthora' en la Argentina. Rev. Invest. AgGreiGwiB.wAl Festacy 4421354 Gerrettson-Cornell, L. 1989. A compendum and classification of the species of the genus Phytophthora de Bary by the canons of the traditional taxonomy. For. Comm. New South Wales Tech. Pap. 45, Sydney. 103 p. Harris, D.C. 1979. Epidemiology of Phythophthora syringae storage rot. East Malling Res. Sta. Rept.) 1978 5:-193-94°: Harris) auD. Coun l9S5a. Survival of Phytophthora syringae oospores in and on apple erchardsoil:.. Trans wBru.wMyYcoln. Soc, 8 os 153=— nies 5 a Harris, D.C. 1985b. Microdochium fusarioides sp. nov. from oospores of Phytophthora syringae. Trans se pre MYVGOl. SOC. 84 58—36 17 450 Harris, D.C. 1985c. The colonization of fallen apple leaves by Phytophthora syringae in relation to inoculum levels in orchard soil. Ann. Appi. “BLOLE. 1072 179=188. Harris, D.C. and D.M. Cole. 1982. Germination of Phytophthora syringae oospores. Trans. Bre MYCOL. foOC es o..mipie) OV et Himmelbaur, W. 1911. Zur kenntnis' der Phytophthoreen. Jahrb. Hambur. Wiss. Anst. 28: 39-61. Ho, H.H. 1983. Phytophthora porri from stored carrots in Alberta. Mycologia 75: 747-751. Klebahn, H. 1905. Eine neue Pilzkrankheit der syringen. Centbl. Bakt. 15: 335-336. Klebahn, H. 1909. Die neue Zweig-und Knospenkrankheit. Krankheiten des Flieders. Gebruder Borntralger, Berlin. 18-75. KlOUZ, teri. Maloy se Color Handbok of Citrus DiseasesscUnavs) OL Caltts Divs of pAGr=1SCc. E227°ps Klotz, L.J. and E.C. Calavan. 1969. Gum diseases Of CTlUcrus. nie Caliitornia. Unive i Ofmca lite. AGE. EXpve sta. #Clr mo 96. es ender eV. «2. Jan. Kouyeas, H. and A. Chitzanidis. 1968. Notes on Greek species of Phytophthora. Ann. Inst. Phytopath.}Benaki, N.S. 8; 175-192. Krober, H. 1985. Experiences with Phytophthora de Bary and Pythium Pringsheim. Bundesanst fur Land - und Forstw. (Berlin-Dahlem) 255: A sei iy Yl 852 Lafferty, H.A. and G.H. Pethybridge. 1922. On a Phytophthora parasitic on apples which has both amphigynous and paragynous antheridia; and an allied species which show the same phenomenon’... Proc. "Roy. Dublin’ soc. 17: .29— 43. Leonian, LH. 1934. Identification of Phytophthora species. W.Va. Agric. Exp. Sta. Bull. 262. 36 pp. McLennany +E. [7 1935" Notes on the organisms causing’ brown yrot Of = "citrusmetrure, ein Victoria: Austraila (Phytophthora Citropnthorare(sm-.) 7k) om). = Leon strata. hibernalis Carne). Proc. Roy. Soc. Victoria (N.S.) 48: 96-103. Moniz, da Maia, R. 1925. Um nuovo Oomycete 451 parasita de a'rvores do genero Citrus? Rev. Agron (Lisbon) eser.a4% jen —)1y Nadel-Schiffmann, M. 1947. Phytophthora hibernalis. Palestine J. Bot. Rev. Ser. 6: 148-157 Newhook, F.J., G.M. Waterhouse and D.J. Stamps. 1978. Tabular key to the species of Phytophthora de Bary. Mycol. Pap.) 143), Commonw. Mycol. Inst., Kew, Surrey, England. 20ND" NOV1C1.1O wy Commer OnGQueeWsC san GONVOC) mie lO6G. A Phytophthora disease of fennel. Phytopath. zeitsch. 46: 139-169. Novotel 'Nova, N.S. 1974. The Genus Phytophthora. Soviet Acad. Sci. Bot. Inst. Lening. U.S.~S-R. 206 p. eyopukigk=s Say eslchke A fruit rot of apples and pears due to a variety of Phytopthora syringae. Long Ashton Agr. Hort. Res. Sta. Annu. Rept. 1930: 147-150. Peglion, V. and M. Sacchetti. 1927. Intorno alla peronospora del Lilla (Phytophthora syringae, Klebahn). Atti. R. Accad. Naz. Lincei, Rend. Cl. Sci. Fis., Mat. e Nat. 5: 696-698. Pethybridge, G.H. 1913. On the rotting of potatoe tubers by a new species. of Phytophthora having a method of sexual reproduction hitherto undescribed. Sc. Proc. Ree DLLD eS OCLs FL oe OA Petri, L. 1927. Rasseyna del cai fitopatologia pin notevoli osservat: nel, 1926. Bol. R. S Cac baw. VeCO,.m/: lsd oepp. Ribeiro, O.K. 1978. A Source Book of the Genus Phytophthora. J. Cramer, Vaduz, Germany. 417 p. Rosenbaum, J. 1917. Studies of the genus Phyvtophtnora...J. Agnus (Res. (830 233-206. Salih, H. 1974. A Phytophthora species new for Turkey determined in citrus orchards in Adanas Use lLULK. swrny.CODaCh 06:96) ja 1 5 Sarejanni, J.A. 1936. La pourriture du collet des Solanees cultivees et la classification du genre Phytophthora. Ann. Inst. Phytopath. Benake 2:35-52. Savage, 4b. UemmlooS . Sexuality in the genus Phytophthora II. Intra- and interspecific 452 pairings. Ph.D. Thesis, W.Va. Univ., Morgantown, W.Va. 87 p. Savage; E.J.,;,C.W. Clayton, J.H. Hunter,” J.-A. Brenneman, C. Laviola and M.E. Gallegly. 1968. Homothallism, heterothallism and interspecific hybridization in the genus Phytophthora. Phytopathology 58: 1004-1021. Schwinn, Pear ae hove ue Untersuchungen zur systematik der gattung Phytophthora de Bary. ArCHGMIKTODLOl. oS eco oo. Sewell, G.W.F. and J.F. Wilson. 1964. Death of maiden apple trees caused by Phytophthora syringae Kleb. and a comparison of the pathogen with P. cactorum (L. & C.) Schroet. ANN. “ADDL. (DIGle oo... 200 ~2oUR Stamps, D.J., G.M. Waterhouse, F.J. Newhook and G.S. Hall. 1990. Revised tabular key to the species of Phytophthora. Mycol. Pap. 162. COA. B. 2INCern sy MVCOWs VANSG. 7) NeW Vo oULneyy England. 28 °p Tucker, Ce M wees. Taxonomy of the genus Phytophthora de Bary. Mo. Agr. Exp. St. Res. Bulla’: 153.0 208 ip. Upstone, M.E. 1978. Phytophthora syringae fruit rot) (ofmapples Vi PinyPath. 327 524-505 Upstone, M.E. and E. Gunn. 1978. Rainfall and the occurrence of Phytophthora syringae fruit rot of apples in Kent 1973-75. ‘Pl. Path. 27: 30=—3 5 Verneau, R. 1953. Le Phytophthora parasite degli agruni nella Campania. Nota 1. Specie osservate sui frutti. Annali Exp. Agaaria Gems OG Oar. Wager, V.A. 1942. Pythiaceous fungi on citrus. Hilgardia 14: 535-548. Waterhouse, G.M. 1963. Key to the species of Phytophthora de Bary. Mycol... Paps. 92. Commonw. Mycol. Inst., Kew, Surrey, England. Ze Pp. Waterhouse, G.M. and E.M. Blackwell. 1954. Key to the species of Phytophthora recorded in the British Isles. Mycol. Pap. 57. Commonw. Myco..;/Inst.| Kew, Surrey,” England. 9. p,. Waterhouse, G.M. and J.M. Waterston. 1964a. Phytophthora syringae. Descriptions of pathogenic fungi and bacteria 32. Commonw. 453 Mycol. Inst., Kew, Surrey, England. 2 p. Waterhouse, G.M. and J.M. Waterson. 1964b. Phytophthora hibernalis. Descriptions of pathogenic fungi and bacteria 31. Commonw. Mycol.) Inst. Kew, sourrey, England. 1i2 p. VOUNG) eRe Ae aug eocmA. Mi LDA chm. S59 A stem canker disease of fruit tree nursery stock caused by Phytophthora syringae. Phytopathology 49: 114-115. (Abstr. ) 454 : “Stpeuaoqry “d se peATeoeYd “eA ‘umoqUPbION “AAyTsireatun eTUTHATA 382M *KHoToyAed Wuetd Jo quowqyzedeq .s puetbug ‘Aezains ‘mey ‘eqnqtqsul TeoTboToOoAW YRTeSMUOUMTIOD : IWO spueTISyAeN ‘uree” 4goIn{Q[NOToOWMMITYIS IOOA neerangTeez3ueD * SHO zequnu uotTsses0e UOTPOSTTOD ernqtng edAL ueoTtr9uly Aq petytquepl :OoLW epee te ets UR SS re L/v¥9S 23°0T eTUIOJTTeO ebueio ¥ LEN 6L-A 40380EG BTUuIOJTTFO puouTw 91809 DOLV Z10€d YON ABZZOO eTurOFT TRO puouTw TE8b9 OOLW (st6s€ IW) €z°zet sdo zoet Aoupug pueTbug etddy 98896 DOLV 69d won - einty 93 z0eAwQUezZ etTUurIOJTTeO uoulaT ZOObE DOLW LST N ATbetTeS uobe10 [tos 1ved 60L9T JOLW (L9 STTSETAON) ST YOTeY ATe 1 jee, [euued LEBYT DOLW SebutaAe “d LL°@2@sS Sao pue[TeezZ MeN Staeb{ha etbe{ tnby 80Lb9 DOLW (9069€ IWO) Te°o0Lz sao Tebn3104 Stsueuts SsnizTO ZSE09 DOLW O9LPET IWO eTTezqsny °M Stsueuts sn1zzToO €SE9S OOLV Ly9a xreAuQUeZ BeTUIOJT TRO ebueio [eAeN GS6ZE OOLW Stfeuseqrq “d @oaAnOS utbtio d}eaq.eqns/3s8OH Apnys ey ut pesn SeburaAs “q pue sTpeureqry wiouydOYAYG FO soqzeTosI *T eTqeL 455 punoz ATQueutTwoparg (yu) pezede, AT\Aueutwoperzg (L) umtpew zebe uo esoTS ynq 19Q3eM UT xXeT azrOoW ‘q10Yys ATTeOTdAA seyourszgq TetpoduAs y3TtTmM (Zz) seyouerq [etpoduAs Hbuo,T ‘xe{T Area YIM IO payourrzquy (Tt) WIOFJTUN = *eVeTpeY =ey fumtTueYyjQueSsAIYyD =D !aqjesoy =y fe qeTTEIS =sS (p) untpew zre6e uo peonporzd etbuearzods (3) 1a}eM UT peonporzad etbuerzods (q) SZUSUeINSsvVeU OG UO saeseq ‘1017a pizepuejS F UPeW (e) ee a re eo eee A Sabet. beeen te paaiemaietel as! Godin et Seed Eo =p SnNoOnp}Sep-uON | T°0 6°T vb OF 8 SL wa) (2) ++ n LEN ae: SnoNnp Tt Oep-uoNn u CyOntat ESTE "08 2SS Gai (Z) ++ 2) 9LBP9 DOL pz: SnoNnpTtoep-uoNn | 0c € Sc 8 CS ia) (z) lee n LOL‘’9 DOL ak SnNoONnptoep-uON uy T°O b'T € 6€ € TP tas (2) == 2) 9889b ODOLY es snoNnpTOep-uON | CaO mia | 9 LEP wa (Z) == e) TOOPE ODOLY sas SNONpTOsp-uOoN uy OSE rela c 8 2 TT TS (aq) (Zz) or ua 60L9T OOLW = snonpTtoep-uoON aq c°0 GS°T La lt~ 101s 6Sae ted snonpTtoep-uoNn uy €°O 8°T 9 cE OT 6S (aq) (z) ++ ed LE8PT OOLY oebutaAs “gq snonptoed L GeOn ec Ca OTL EES 6 Corcay (T) ae 2) 80L09 OOLW peonpoid umtbueitods on (tT) -- S %S£€09 DJOLW EL-E2 snonptosd L C70 be €¢ Cf G €S (3) S9SLC snonptoed L c°0 8B°T fff O€ OOP a) (T) sobs Re tSE9S OOLY EL-LZ snonptoeqd L €°0 T°2 € bT 9 62 «ay (Tt) ae s SS6ZE DOLW stTeuzeqty “qd (um) yqbuetT (um ) (un ) [Teotpeg Aytonped oeseg q/1I yypeer1g yybuey Bbutyouezrg = sbut[Tems aYeTOs] emia Carew a or eee tae BIT TD UW OOS ota enn) ae ter acca aman, e1o0yd teydAy AuojtoDp /setoeds -otbuerzods “eebutizAs “q pue STyTeuteqry eIOyAYdo AY Jo etbuerods pue seydAy JO SOTQJSTAaVOeAPYD °“% STQeL 456 yueujwopetd Bip} Joy Us SNOUABSIGd = gd {JUGUIwOpPeId B1P1s24 UE snou6 1 yc SUsMIINSesw YG UO paseq ‘JOJJ2 PIePUueIS 4 USI} iT 'su0i1,sodosd BusAsen Uy BYP} Jouue snouABesed pues snouA6i ydwe = d/V eee but bF01 bF0l (WN) YIPIA “eebui Xs “a @buisAS “g Pue SI GUJOQiy 850Yy) WXyg pO SueBsO KIS JO $I19S1I9IIOICY) 201 Z40i L501 (4s) yy Bu2 } eNipt soyjuy d/ eV d/" q d/ adlA j G*Gt 240¢ 9SE 740£ peonposd sue6so Kas ON 7508 9556 2562 pesnposd sue6so Kas OF 2526 ert (un) weip 2100800 cree 2556 , 2398 (un) wep wn 1 u0o6090 rod) 91899 JI1V 20299 II1V 98899 IIIV 200%£ IIIV 60291 IIIV £€89t IIIV sebu aks e019 JIIV 2609 291V £595 JD1¥ 6626 I91V Sy peusaqey *d 29@)0S{/sai2eds “¢ 21981 457 (896T) .dwW (BuoT)aq ¢°Z 8T OF S7> °Z4TYD ¥ Tifosi i/=a dv pe (buot)a 0°Z 61 GE GZ> -- abueirio seofnoy (L96T) d ebuer90 uPeWwoUUeITg (996T) Ol/eed 8Z Ze u Tedsoq (€96T) W ebueiro ebeares (p96T‘E96T) PI/ Z@I/d.W oTIOAeTd Ge (ss-)a 6°T 61 9€ GZ / G> -- u esnoyre7em 2(€S6T) dW b°SZ Eye Lré 6°T c° st b°vEe v7> -- wU/snaqtyp nesureA St T2- u/eTuoUtT 2(0S6T) -OT/S°0OT/,dW LS ze (08-9T)a 9ZL°T GZ G°ePp un €2@ / G sna4ToO TZZe1g c°GE 6°T 4 G°LE o(ZP6T) z7I-6/ 4d 6Z2-LZ ze (Gz-oT)a o°2 8T 9€ Cor ebuez0 rabem (9€6T‘EE6T) dW Gb-72Z 9¢-z7z (9G-Z)a 8Z-OT 9S-OT GZ sna3zToO 22990Me7 ebueio (se6T) (Huot )a Teptosdt{tta uoula'T ueuueTOW (peét) dW GZ> uetTuostT (TEé6T) d L°SZ Dv°S8zd 6°T 6°LT p°ve GZ / G> snaiqto TsyxonL dW TE-TE LT°GE Tit b°PT O€ (6z6T) (os)a T’°c T°6T TP snri4ToO opnesusgd (SzZ6T) eTeW G°7P-77 GTS 8S—GerT == ep zTuoW Led €° PT €°O€ GE 8°Or (9G-z)a €2Z= “GST 6° PE (¢z6T) dW 9°S'-72z 9S-7Z 8Z-OT 9S-LT un (2)++ n/snazqto eure) sT[Teureqry “d (um ) (um ) (um ) (url ) (uri) j6utTyouerq xeW/UTW Y9M/U9T/pedAL *weTd *“weTtd oA¥TONpeD G/T yypeerg yAbueT aizoyd (D)duez 6buttTemsg ,Auojtoo wNnTpTtreyqUuYW e10ds00/untuoboo untbuertods -otbue10ds y Mor» TeydéAy /380OH *jou *einzerezyT]T UT peprzoce1 SebutiAs -q pue StT[eureqty exroyzqdoyAYd JO SOTYSTISZOeIeYD °y STqQeL 458 dW Tt/ tTt/ dv dW dW OT/ @T/ dW Tt=-6/" 8 Ge-ve GZ p°sSd v°vC GZ-LT 07-02 99-77 Ge-SZ TE-8BC OF= EZ CHL 8Z v°97 an 0S°6Z2 an aN 9€-8T aN 0S-0€> (os<)a 95-02 (oz<)a (oz<)a os-8z. (buot)a (g°8Z v°672 -G°Lt)d 9S-SE (cs)a te (ps-7zz)a (buot)a 6b°T aseq pexzoedeq/m ptosogo 61-TT Gp-EE Le Ov €£°S7 98° 6€ Th-OE GL-O¥ Tv-6T GL-¥Ce GL-S¥ 6T-PT TV-OE p°LT OP 87-61 9G-9E LT TP st 9D 8I Le TeprosdTtTla T T‘un Ss I LO EC =o ad 97-SZ/ SG> —— a (LZ6T) *90eS ¥ uot {bed (7z6T) uAnag 2d (ZZ6T) "uqed 3 Ajzreszzel (LT6T) ote unequosod (T1T6T) aneqTSouuiTtH (606T) uyeqeTy aebutaAs ‘d oetTT oeTTI atddw ++ oeTTT oeTTI (0661) “Te Ze sdueqs (6861) TTeurz0) -u0s 43e1199 (6L6T) oesL ¥ sniqto AyArepeH-TW (gL61) *{Te 32 = yOOUMON (8161) U ePURPTUOSMET $3no014S y aretseig (8L6T) o1tTeqod (LL6T) stsuouts qZesnL snaiyto -*yoerizegd (PL6T) yttes (€L6T) ZQ0OT Saya GZ / G> a 97-PC oabueio sn13zTto (penut quod) *py eTgqeL 459 L720 U/e. (w)9°6T/ , LAN GoZT/ =. d peonpoid peonpoid peonpoad peonpord peonpoird peonpoid 6/ 6/ .d¥ dW d S2-LTX9E-8T dw sueb1i0o x9s ON sueb1i0o xos ON suebio xos ON suebio xoes ON suebio xaos ON suebio xos ON 9E-8Z2 9TIOISTd €€ 62 vE TE GE 62 vE O€ PE LZ O€ peonpoid suebio xas ON Se 8°8Z 8°92 6°62 Cale 8°62 9°T 89°TE GZ b°8Zz d an [aman | aN v°T aN b°T aNn v°T aN See aN (SU (eo Saat an 9°T aNn ESS aN Let aN Saat aN 951 aNn Cae aN Siar aN UG Seer v°T €°8C ESSe 8 Gea Cam L Coal GE-6T OF-tT O£€-02 9E Te Sts ce ce 99-PC SSRC 6€ cS 8P 8E OS-OE LS GL 9S 8P cS L@ / G> Le Le Le uN 8t@-97/G> S -€¢ °/ S> s‘un 9E-SC/=0 G7< Ss A Ss VER /SE=0 Seo C=6C/.€-0 Gz / G> +4 ++ +e ++ Se At ++ +e +e ++ ptt +4 (6S6T) uUuTMYOS (896T) °Te 39 obeaes sTunuwood snakd etddy qootady yored (896T) uTiepuen *Z4TUD ¥ uoUusT seoAnoy (7967) uOSTTM ¥ o/etddy T1TeMes (p96T ‘€96T) U esnoyre eM (€96T) aepAus ¥ n/Teuusg OTTOTAON d/ieed (SS6T) etTddy usbeyueppng (pS6T) TLTePMxoeTA ¥ esnoyrzsejemM (€S6T) D/sniy4to nesurzsa (LP6T) 3° 3FTYOS N/snaz4Tod -TePpeNn N/ieed (TE6T) oetddy STATTBO (9€6T) sniqto q22e00Meq (7E6T) OeTTI reyssyo STieb TNA (TE6T) ebutads rayon, (penut quod) *p OTGeL 460 soeyouetg Huot YyyTts TtetpoduAs prottezed = d ‘unuayquesAryo aedAq queuTWOd snonpTooep-UON ED (StL eureqry soezodsopAuweTyo se pe zi0odey Leen Eee ST Teureqry » !snoudAbeired an {(yqbueT Teotped) snonptoed qT {seyouerq ZrOYsS YATM TetpoduAs 7qZ =) Sebutaks “d se pe zr0deu a !snouAbtyduy Tq se poqrodey Ra yl Ras, | Aa « o v0 eal a s {peyourerqun = UN, faqqesor = y twIOFJTUN = Oe (0661) °T? OT>/0T>/,dv 0b-02 dv 62 d Of aw G-8¢ OI>/OT>/ ,dW Ge-SZ Lf -Ot/ d €v-Te d 8E-82 dw 0S-O€ Cece 0S-82 9P-EE an an an an an 9°T> 7-Z°T O€-02 O°C TE 95° T G°62 9°T 9E 9°T> CV-9E O€-6T GL-SP 90-VC T9 GC=LS 0S-772 Ss s‘un LZ-€S> Me tErGyp SE EC /aaeG [HAS dhe 43 CU afi GS GZ €z / G> i mag een 4 4b ++ ++ a je sdureqs (686T) TTeur0D -u0s}3e71eD d/puowT wz u/stunuuod snakd ES eoetuouite ++ ++ +t snunid w/snz3TO a/snz3Td d/etddy o/etddw (penut zuoo ) (ss6T) azeysod 3 yoo sod (ss86T) zeqoiy (8L6T) STTTAePI 3 seo00g (g£6T) "Te We YOOUMEN (8L6T) auo ysdn (8L6T) olteqta (LL6T) Aejtod (pL6T) * TOVZOAON *p eTqed MYCOTAXON Volume XLVI, pp. 461-466 April-June 1993 A NEW CORTINARIUS FROM SOUTHEASTERN TEXAS ORSON K. MILLER JR. Department of Biology Virginia Polytechnic Institute and State University, Blacksburg, VA 24061 SUMMARY INTRODUCTION Southeastern Texas has large areas of mixed pine and hardwood forest with loblolly pine (Pinus taeda L.) intermixed with beech (Fagus grandifolia Ehrh.) and southern oaks including swamp chestnut oak (Quercus prinus L.), sand jack oak (Q. virginia v. geminata Sarg.), willow oak (Q. phellos L.), and swamp red oak (Q. falcata var. pagodaefolia Ell.). Watson (1979) describes several dominant plant communities including areas of beech-magnolia-loblolly pine forest, which is a transitional association between the wetlands and the dry uplands. It occurs in the uplands and hilly terrain interspersed with successional lakes or acid bogs "where underground water seeps to the surface". In this area Watson (1979) refers to the more advanced vegetation in old lake beds as baygalls, which consist of sweet bay (Magnolia virginiana L.) and gall berry holly (Ilex opaca Ait.) along with black gum (Nyssa sylvatica var. biflora ) and red bay (Persea barbonia). These vegetational area are often in close proximity to each other. Within this complex plant community a rather abundant, undescribed, distinctive, bright yellow Cortinarius first came to the attention of David Lewis who made notes on the fresh characteristics, habit and habitat. It is especially common along the boarder of the baygall communities in the transition zone to the higher elevation referred to as the beech-magnolia-loblolly pine community. Since the baygall community has plants that are typically endomycorrhizal, an ectomycorrhizal host must be found elsewhere. David Lewis and I have observed fruitings most often in association with beech, but loblolly pine or the various oaks which are subdominant members of the beech-magnolia-loblolly pine community might also be associated. Methods: Each collection was placed in a separate wrapper to avoid mixing material. The possible mycorrhizal hosts were recorded at the time of collection. Smell and taste were recorded for specimens at the time of collection and again in the laboratory. The color standards used included Ridgway 462 GLODZ5T, (the colors placed in quotes) and Kornerup and Wanscher, (1978) (the colors indicated by (3A7) which in page 3, row A and color block number 7). Microscopic examination was carried out using 3% KOH and Melzer's solution. All collections are deposited at VPI unless other wise indicated. TAXONOMY Cortinarius lewisii sp. nov. O.K. Miller Illustrations: Texas Mushrooms p.177. Pileus: (1-) 2-3-5 cm latus convexus, plano-convexus aut paene convexus umbone lato, squamulis aridis parvis appressis; margo incurvatus luteus aut aurantius. Lamellae adnexae aut adnatae confertae angustae seriebus duobus Gispositae, in juventute albae, in maturitate aurantiacae, in vestustate ferrugineae. Stipes 4-7.5 X 0.3-0.9 (-1.1) cm latus aridus aequalis, fibrillis tenuibus juventute album, luteum in vestutate, annulo luteo prope apicem in maturitate. Odor moschatus aut raphano similis. Sapor mitis. Sporae 6-7.5 X 5-6 um diam., subglobosae, verrucis demissis brunneis. Depositum sporatum ferrugineum. Pileipellis mixocutis est hypharum filamentosarum 3.4-8.4 um diam., fibrillis praesentibus. Trama lamellaris parallela hyalina parietibus crassiusculis. Typum legit. D.P.Lewis-4370, 13 May 1990, Lance Rosier Unit, Big Thicket National Preserve, Hardin Co., Texas. Byleus:, (lO. e2-3 50cm -=broad pconvex;. Dlano-COnvex asco nearly plane with a broad umbo in age, dry, with minutely appressed, squamules, surface bright orange-yellow "Cadmium Yellow", "Light Cadmium", margin "Apricot Yellow", buttons chrome yellow (3A5), buttercup yellow, sunflower yellow (4A8), to some cadmium orange (5A8). Margin incurved at first, plane in age. Lamellae: adnexed to narrowly adnate, close, narrow, with two tiers of lamellulae, nearly white in buttonsto Pisabellat o(2.5Y.6.5/4) ;."Orange=-Citrine: ~sinvage becoming rusty brown from the spores. Stipe: 4-7.5 cm long, 3-29 (-11.), mm; wide, dry, equal; coverediwithjrine fiprits, "Light Cadmium" to cream color with white mycelium over the base Partial veil: dense, cortinous, white in button but soon yellow as pileus matures, forming a bright yellow annulus near the apex but often absent at maturity. lesh: firm, "Deep Colonial Buff", to "Isabella Color". ODOR: musky to radish-musky. TASTE: mild. Pileipellis: 80-180 yum thick, a mixocutis of filamentous hyphae 3.4-8.4 um diam, loosely interwoven at the surface, more compact below, with scattered intracellular incrustations, occasional branching, scattered clamp connections, yellow to orange in Melzer's solution, hyaline in 3% KOH. Pileus trama: of broad hyphae 4-16 um diam, becoming closely packed resembling a textura angularis in 463 Figs. 1-3. 1, 2 and 4 sp'd basidia and a basidiole. 2. basidiospores. 3. filamentous clamped hyphae of the pileipellis. Scale bar = 10 umm. 464 the mid to lower trama, often with common, thick-walls (0.3- 0.6 wm) and scattered clamps, hyaline in 3% KOH, yellow in Melzer's solution. Lamallar trama: parallel, thin-walled, filamentous, hyaline, hyphae 2.5-7.6 um diam slightly thick- walled near the pileus trama, often with scattered clamp connections, hyaline in 3% KOH to light yellowish in Melzer's solution. Subhymenium: a narrow layer of thin- walled, short, filamentous, hyphae 1.7-2.5 um diam, hyaline in 3% KOH, with scattered clamp connections. Basidia: 19-35 X 6-8 um, clavate, laginiform, to clavate-rostrate, thin- walled, 4-spored, with infrequent 1 and 2 spored basidia, hyaline in 3%KOH and Melzer's solution. Partial veil: of filamentous, thin-walled, hyaline, hyphae 2.5-6 (-8.5) um diam infrequently branched at right angles or Y shaped with frequent clamp connections. Universal veil not present. Basidiospores: 6-7.5 X 5-6 um diam, subglobose, globose, thin-walled with low, brown, warts and a small to indistinct, hyaline, apiculus, yellow-brown in 3% KOH and Melzer's solution. Spore deposit brown. Habitat and distribution: terrestrial, common, gregarious, some subcaespitose, in mixed pine and hardwood forest or in the beech, magnolia-loblolly pine mixed forest, often near baygall communities. In Southeastern Texas. Fruiting in spring from May until June and again in October. Material examined. United States. Texas: Orange Co.:Vidor, 23 May 1977, Lewis 974% Vidor, 5 May 1979, Lewis, 1719; Vidor, 16 July 1989, Lewis 4297; Hardin Co., Lumberton, 4 Oct. 519797) Lewis 20357) Vidor) (28) May, 1989s. bewrs 74250; Hardin Co., Lance Rosier Unit, Big Thicket National Preserve, 13 May 1990, Lewis 4370 (Holotype); Orange Co., Sam Houston Nat. Forest, 2 Nov. 1979, O.K. Miller 18684. Austria. Tirol: Vikartal, Cortinarius nothosaniosus, 9 Sept. 1951, M. Moser Nr.51/120 (holotype) (I). Sweden. Femsjo: ost Trollgol, Cortinarius limonius, 8 Sept. 1974, M. Moser, Nr.74/328 (1). Femsjo: Aborrasjo, Cortinarius Fimoneus,;) 4 Augeuloy2, “MayMoser;, Nr wiz /1oo (iy Observations: In the baygall communities Cortinarius lewsii is often gregarious and numerous. The bright, orange-yellow, dry, pileus, yellow flesh, (Metzler et al 1992) and small subglobose to globose spores, and a pileipellis of interwoven hyphae are a distinctive set of characteristics which would place this species in the Subgenus Leprocybe,. The section Limonei has species with vivid yellow to yellow brown pileus coloration and globose, subglobose, to broadly elliptical, warted spores. It appears to be close to C. nothosaniosus Moser which is found in "versauerter Fischtenwald (Picea abies) mit Vaccinium myrtillus"” in Europe, (Moser, 1966). However, C. nothosaniosus has a more acute umbo as illustrated by Moser (1966). In addition, the pileus is smaller (up to 2.5cm broad), the lamellae subdistant and the stipe thin and small (1.5-4cm X 2-4mm thick). Buttons are red brown with a yellowish margin which is fibrillose from the veil. Microscopic features are 465 Similar to those of C. lewisii. The basidioles of C. lewisii are variable in shape as described above and probably result from the maturation of 1, 2 and 4 spored basidia.and distinctive clusters of hyphae are present in the pileipellis with cells 8-30 pum diam. Cortinarius Limonius (Fr. Jex) Fro) Fr. is) similar’ but hasian'|"foxy, red" to "orange red" pileus, is more robust than C. lewisii, and has a clavate stipe. It is a European species also found associated with spruce and pine (Moser & Julich 1986, Moser), 1933)", Species that produce orellanin are found in the Subgenus Leprocybe. Testing for orellanin by Poder and Moser (1989) have revealed that the toxic species are in the Section Orellani. When tested neither C. nothosaniosus nor Cc. limonius, in section Limonei, contain orellanin. It is logical to conclude that C. lewisii also is unlikely to containmene toxin: ACKNOWLEDGEMENTS I thank David Lewis for notes and collections and the Texas Mycological Society for their hospitality. Susan and Van Metzler provided support and help. Dr Harold Burdsall and Dr. Joe Ammirati provided much appreciated comments on the manuscript. Logistical support was provided by the Virginia Museum of Natural History. LITERATURE CITED Fries, E. 1821 Systema Mycologica I: 213. Metzler, S., V. Metzler, & O.K. Miller. 1992. Texas Mushrooms. Univ. of Texas Press, Austin, Texas. 350p. Moser, M. 1966. Einige interessante Pilzfunde aus dem Gebiet von Gotschuchen. Sonderdruck aus Carinthia II, Mitteilungen des Naturwissenschaftlichen Vercines fur Karnten, 76, bzw. DSO e390. Moser, M. 1983. Keys to Agarics and Boleti. Gustav Fischer Verlag, Stuttgart, ‘Germany 535 ). Moser, M. & Julich 1986. Farbatlas der Basidiomycetes. III Agaricales. Cortinarius 3. Gustav Fischer Verlag Kornerup, A. and J.H. Wanscher. 1966. Methuen handbook of SoLlour.ndled. .Metnuenta.CO.,) LUtLa.e) London, Poder, R. und M. Moser. 1989. Eine einfache, empfindliche methode zum makrochemischen nachweis von orellanin. Mycologia Helvetica 3: 283-290. Ridgway, R. 1912. Color standards and color nomenclature. Washington D.C. 44p., I-LIII Pl. 466 Watson, G. 1979. Big thicket Plant Ecology. 2nd Ed. Big Thicket Museum, Saratoga, Texas 10 p. Fig. 4. Cortinarius lewisii xX Le MY COTAXON Volume XLVII, pp. 467-474 April-June 1993 BOOK REVIEWS L. M. Kohn, Book Review Editor Mycota Lithuaniae Volume 5 (1): Rudieciai (Uredinales), by A. Minkevicius & M. Ignataviciute. Hardcover, 223 pp., 240 x 170 mm, 1991. Published by the Lithuanian Academy of Sciences. Requests must be sent to the Institute of Botany (Zaliyju ezery 47, Vilnius, 2021, Lithuania), fax 01 22 359950. This represents the first in a series of of floristic volumes which will eventually cover all major groups of fungi known to occur in Lithuania. The present fifth volume was published out of sequence due to the advanced age of its first author, Professor Minkevicius, who has studied the Basidiomycota at the Lithuanian Academy of Sciences over the last 50 years. Although this first volume contains only line drawings, future plans for the series include color photo illustrations of macrofungi as well as detailed taxonomic descriptions of new taxa. The current volume includes keys and descriptions to about 170 species of rust fungi from Lithuania, with data on their distribution and host biogeography. The introductory chapters also include interesting discussion on rust life history, and particularly the apparent rarity of certain life cycle stages (mostly aecial) in many species owing in part to climatic conditions in this Baltic country. The Mycota Lithuania project was conceived and its first volume published while Lithuania was still a Soviet state. Although economic problems of a free Lithuanian state will certainly delay publication of the remaining volumes already underway, the eventuality of a complete documented mycota for Lithuania will represent a valuable contribution for understanding the biogeography of European fungi. Although this first volume is written in Lithuanian, it contains excellent summaries in both Russian and English, and most of its technical information concerning taxonomic characters and distributions should still be readable and of interest to other professional mycologists working with these fungi. -R. Vilgalys, Department of Botany, Duke University, Durham , North Carolina 27708 U.S.A. Foliicolous Lichens - A Contribution to the Knowledge of the Lichen Flora of Costa Rica, Central America, by Robert Liicking. Beihefte zur Nova Hedwigia, Heft 104. VII: 1-179 + 64 figs. Softcover, 179 pp., 240 x 170, 1992. J. Cramer in der Gebriider Borntraeger Verlagsbuchhandlung, Johannesstr. 3 A, D-7000 Stuttgart 1, Germany, fax (0711) 625001. ISBN 3-443-51026-4. DM 170.--. The principal component of this publication are keys to all (c. 230) obligately foliicolous lichen species known from Costa Rica. Most of these have been found there for the first time by the author during his 468 extensive investigations, including several new species published in a separate paper (Nova Hedwigia 52: 267-304). Added are very useful, good quality, black and white photographs of most species, and many line drawings of spores, asci, paraphyses, etc. Costarican localities and world distributions are provided for each species, and for most species, key, discriminative characteristics are pointed out. The short general part contains a list of collection sites and a glossary. This book transforms Costa Rica into the country with the second largest known foliicolous lichen flora in the world. The size of this flora probably results from the varied topography of the country. It certainly also results from the poor knowledge of the lichen flora in other tropical countries. Facultatively foliicolous lichens are poorly represented in the book. These may be recognized because they tend to also grow on adjacent | twigs. They are by no means rare and are sometimes really conspicuous. Either Lticking happened not to come across them, or he omitted them, not an unusual practice. Further, a number of common neotropical taxa, like Mazosia tumidula, are missing; evidently these have not yet been found, or do not occur, in Costa Rica. For evident reasons the data on world distribution should be considered as provisional. The ecological rather than taxonomic background of the author is reflected in some problematical taxonomic practices. The introduction of "ad interim" names seems undesirable because they may come into wider use through the keys and lead to nomenclatoral confusion. Further, the "isotypes" do not completely reflect current use because they bear different numbers; however, they do not really deviate from the common practice of including in a single collection a number of individuals growing together. In this treatment, individuals which grew on different nearby shrubs have been given different numbers. The main significance of this book is as an identification tool. It forms a very valuable addition to Santesson's monographic treatment of 1952, the main source of information for the group, because it treats numerous species which have been discovered since 1952, and follows the newest taxonomic rearrangements. Moreover, the numerous illustrations are a welcome complement to Santesson's sparsely illustrated book. The keys contain most of the important characters, making descriptions largely superfluous, and the illustrations capture subtle characters which are difficult to aptly convey in words. H. Sipman, Botanischer Garden & Botanisches Museum, Konigin- Luise-Stragpe 6-8, D-1000 Berlin 33, Germany. 469 The Polypores of China, by Zhao Ji-Ding & Zhang Xiao-Qing. Bibliotheca Mycologica Band 145. Softcover, 524 pp., 225 x 140 mm, 1992. J. Cramer in der Gebriider Borntraeger Verlagsbuchhandlung, Johannesstr. 3 A, D-7000 Stuttgart 1, Germany, fax (0711) 625001. ISBN 3-443-59046-2. DM 190.--. This flora, written in English, treats 351 species. One new combination is proposed, Phellinus quercinus (Zhao) Zhao, and no new species are described. A short introduction is followed by a list of the Chinese polypores. There is a key to the 76 genera and keys to species appear under the treatment of each genus. The generic concepts in most cases follow Gilbertson and Ryvarden's North American Polypores, 1986-1987. The polypores in the major works on Chinese fungi by Tai (Sylloge Fungorum Sinicorum, 1979) and Teng (Fungi from China, 1963) have been incorporated. The species’ descriptions are original having been prepared from _ Specimens studied by the authors. They are detailed and follow a consistent format. The principal microscopic features of 318 species are illustrated by line drawings. Each species treatment includes data on habitat, specimens examined, and distribution. The sources for the notes on habitat were probably comments accompanying the specimens, but some comments seem to be from the literature. The distribution simply lists some countries where the species has been reported. A surprisingly large number of the species, e.g., 53 of the first 100, also occur in the northern United States and Canada. Those using this text to identify collections should be aware of a few potential problems. First, in the key to the genera each generic name appears only once. Thus, a genus having some species with and others without clamp connections, appears only once under one key choice, e.g., with clamp connections, and it will be difficult to correctly key-out those species lacking clamp connections. Second, Meruliporia violacea with hyaline, thin-walled spores is obviously misplaced. The prevailing concept of Meruliporia is of a genus with distinctly yellow- brown spores. Third, in some drawings the spore shapes differ somewhat from North American specimens, i.e., the spores are overly attenuated at the apiculus end. Fourth, the size of the spores in the drawings sometimes differs from the sizes given in the text, e.g., for Dichomitus squalens the text says spores 7-8 Um long but the drawing has spores 6-9.5 um long. Finally, the spore ornamentation in Bondarzewia montana is correctly described as "short, irregularly arranged ridges," but the illustration shows five spores with conical to hemispherical warts. This is an impressive volume. It is well-organized, detailed and readable, except for some instances where the choice of words is confusing. It will be an important reference in the identification of specimens from Asia. Students of the polypores will find this a useful volume, as will biogeographers, forest pathologists and plant quarantine scientists. J. Ginns, Centre for Land and Biological Resources Research, Ottawa, Canada. 470 Pilze auf Pilzfruchtkorpern. Untersuchungen zur Okologie, Systematik und Chemie, by Wolfgang Helfer. Libri Botanici 1:1-157 + 8 color figs. Softcover, 240 x 170 mm, 1991. [HW-Verlag, Bert- Brecht-StraBe. 18, D-8057 Eching, Germany, fax 089-3192257. ISBN 3-9802732-2-9. DM 88.--, US $55.--. This initial volume of a new series concerns the fungi that occur on other fungi, sometimes referred to as the fungicolous fungi or fungicoles. The book consists of two basic sections. One section presents a taxonomic account of the 140 species of Bavarian fungicolous fungi encountered in the study including seven new species. Although primarily Ascomycetes and Fungi Imperfecti, there are 9 species in the Zygomycetes and 13 Basidiomycetes. Fungi occurring on myxomycetes are listed separately. Many of these species occur widely in temperate regions. Descriptions are included for only the newly described species with taxonomic literature for each species abundantly cited. A few taxonomic inconsistencies exist, for example, Nectria vilior Starb. (as N. violor) is listed as a synonym of N. berkeleyana on p. 52 while on p. 47 it is not and should not be in my estimation. But for the most part the taxonomy is accurate and up-to- date. This section combined with the list of substrates, a key to genera, and a key to the known or suspected anamorphs of Hypomyces make this book an extremely useful synopsis of fungicolous fungi. The key to anamorphs of Hypomyces is especially useful given that many Hypomyces are encountered only rarely as the teleomorph but may have one or two distinctive anamorphs. Results of a chemotaxonomic analysis of anamorphs of Hypomyces bring into question current concepts of these anamorph genera. The second section presents experimental data on factors that influence infection and host specificity of Hypomyces chrysospermus, a fungicolous species thought to be restricted to members of the Boletales. In one series of experiments, extracts of various host fungi are shown to affect the germination rate of aleurioconidia and phialoconidia differentially. Other experiments involve the effects of extracts of Paxillus involutus on aleurioconidia under various conditions. Finally, an attempt was made to isolate and identify the factor that stimulates germination of aleurioconidia. Although unsuccessful, this research is unique and fascinating with implications for related fields, for example, circumscription of the Boletales. A. Y. Rossman, USDA-ARS Systematic Botany and Mycology Laboratory, Beltsville, Maryland 20705-2350 U.S. A. 471 Ektomykorrhizien an Fagus sylvatica: Charakterisierung und Identifizierung, okologische Kennzeichnung und unsterile Kultivierung, by Felix Brand. Libri Botanici 2: 1-229. Softcover, 240 x 170 mm, 1991. IHW-Verlag, Bert-Brecht Strafe 18, D-8057 Eching, Germany, fax 089-3192257. ISBN 3-9802732-4-5. DM 105.--, U.S. $ 65.--. Since the mid 1980's, Reinhard Agerer and his students have published several studies on the anatomy and morphology of various ectomycorrhizae in their well known series of publications entitled "Studies on Ectomycorrhizae" as well as in the Colour Atlas of Ectomycorrhizae. Now, a compilation of studies by Felix Brand is available on the ectomycorrhize of European beech. Keys are provided for the identification of 23 distinct beech ectomycorrhizae along with their fungal symbionts. A detailed description including line drawings and black and white photographs of the morphology and anatomy is provided for each ectomycorrhiza in question. In addition, the manual contains sections on how to prepare and examine field-collected ectomycorrhizae and how to interpret the descriptions provided in the text. Detailed line drawings aid in this respect. Comparisons are made with reports of ectomycorrhizae on beech by other authors. A whole chapter provides detailed information on the edaphic factors and general ecology of the ectomycorrhizal fungi associated with beech, while another deals with the synthesis of beech ectomycorrhizae under laboratory conditions. The specialized scope of the book may limit its usefulness. However, the detailed information presented cannot be overlooked by any researcher working on the anatomy and morphology of ectomycorrhizae found on other tree species. L. J. Hutchison, Department of Forest Science, University of Alberta, Edmonton , Alta., Canada. Book Review Editor's note: MYCOTAXON has received three additional volumes in this series. Libri Botanici 3: 1-215; Allgemeine Taxonomie und Chorologie der Pflanzen, by Werner Rothmaler, a 1992 reprint of a work originally published in 1955 (ISBN 3-9802732-5-3, DM 36.--). Libri Botanici 4: 1-115 + map; Die Vegetationsentwichlung in NaBbaggerungen der bayerischen Donauebene, by Andreas Otto, 1992 (ISBN 3-9802732-7-X, DM 68.--). Libri Botanici 5:1-288 + 16 color photographs, Pilzflora von Bayreuth und Umgebung, by Wolfgang Beyer, 1992 (ISBN 3-9803083-1-6, DM 88.--). Note that the Pilzflora includes many ascomycetes (with some nice color photos) as well as basidiomycetes; Red List status is indicated for some entries. 472 An Illustrated Manual on Identification of some Seed-borne Aspergilli, Fusaria, Penicillia and their Mycotoxins, by K. Singh, J. C. Frisvad, U. Thrane, and S. B. Mathur. Softcover, 133 pp., 296 x 210 mm, 1991. Danish Government Institute of Seed Pathology for Developing Countries, P.O. Box 34, Ryvangs Allé 78, DK-2900 Hellerup, Copenhagen, Denmark. ISBN 87-7026-3175. Price outside Denmark is DKK 220.-- (excl. VAT) + postage. This manual was written to provide a means of identifying seedborme species of Aspergillus, Fusarium, and Penicillium, with an emphasis on mycotoxins and other secondary metabolites as identification criteria. The book begins with information on techniques for isolating these fungi into pure culture, for growing them under standard conditions for morphological observations, and for detecting mycotoxins in culture. A compilation of informationon physical characteristics of mycotoxins is provided, e.g., UV spectra, TLC data and HPLC retention indices. Descriptions illustrated with mostly color plates and line drawings are provided for 15 aspergilli, 9 fusaria, and 18 penicillia. A synoptic key is provided for species treated in each genus. The final chapter details methods for detecting and identifying mycotoxins in seeds. The book ends with a glossary of over 40 terms and a bibliography arranged by subject, with comments on some of the works cited. The authors state that they have followed the modern taxonomic and nomenclatural approach outlined in Samson and Pitt, Eds. (Plenum Press. 1990). I find two main limitations to this manual. First, the color terminology is general and lacks reference to a standard. For example, “creamish yellow" describes the color of the reverse of a number of colonies that are shown in the plates with colors that differ significantly from each other. Additionally, for some species the color seen in the plate does not correlate with the color indicated in the description. Second, measurement data of microscopic structures are not provided, presumably because the information is available elsewhere and because this book emphasizes the use of mycotoxins for identification. This emphasis assumes a lack of variation in mycotoxin production within a species. It also requires that additional references be used to identify an isolate with certainty. For these reasons, the book is less useful as a lone identification manual, especially for users not already familiar with these organisms. Despite these limitations, this book successfully addresses the needs of a user group that must identify species in this difficult group. The color photomicrographs are generally of high quality. The most valuable aspect of the book is the fact that it is a concise source of identification information for these three genera, especially regarding mycotoxin detection techniques. Users will find this book a worthwhile addition to their collection of identification tools. M. E. Palm, USDA/APHIS, Systematic Botany and Mycology Laboratory, Beltsville, Maryland 20705-2350 U.S.A. 473 Pleurotus unter Ste}. Okophysiologische Untersuchungen zu Wasserhaushalt und Sporulation, by Angelika Achhammer. Bibliotheca Mycologica Band 141. Softcover, 206 pp., 225 x 140 mm, 1992. J. Cramer in der Gebriider Borntraeger Verlagsbuchhandlung, Johannesstr. 3 A, D-7000 Stuttgart, Germany, fax (0711) 625001. ISBN 3-443-59042-X. DM 130.--. This doctoral thesis reports on the role of water relations in vegetative growth, fruiting, and sporulation of several isolates of Pleurotus ostreatus and one of P. pulmonarius. Emphasis is on the response to water stress induced either directly or indirectly. Long term and daily sporulation patterns were determined in the field and under various combinations of controlled temperature, humidity and duration of daylight and darkness. Water loss, uptake, and transport were studied with detached fruitbodies taking into account differences among major fruitbody regions. Although sporulation continued despite considerable dehydration, even very high relative humidity could not revive dry fruitbodies. Liquid water was needed for resumption of sporulation, and to be effective had to be supplied through the cap because water movement through the stipe was generally too slow. Total water and osmotic potentials of fruitbodies were determined, and the effects of drying and freezing were studied by scanning electron microscopy of different fruitbody regions, especially the hymenium. Vitality was tested with fluorescein diacetate. Tolerance of osmotic stress was studied for mycelial growth, fruiting, sporulation and spore germination in cultures grown on media with added osmotica (NaCl and for some tests glycerol). The content of trehalose and several polyols was determined in fruitbodies subjected to osmotic stress and freezing because these compounds may protect against water stress. The book will interest researchers studying the physiology of water relations and ecology of Hymenomycetes. It will also interest Pleurotus growers and forest pathologists. There is a three-page summary in English and an extensive bibliography. Few typographical errors were noted. Unfortunately, readability suffers from the fact that the results are subdivided into many sections each with introductory remarks, literature review and a lengthy discussion. In consequence it is difficult to focus on the results and to get a clear overview of the material. The problem is increased by the large number of abbreviations and the excessively complex numbering of the figures. H. E. Gruen, Department of Biology, University of Saskatchewan, Saskatoon, Sask., Canada. Texas Mushrooms: A Field Guide, by Susan Metzler and Van Metzler; Scientific Advisor, Orson K. Miller, Jr. Hard or Softcover, 304 pp. with 249 color photographs, 5.75 x 8.25 inches, 1992. University of Texas Press, P.O Box 7819, Austin, Texas 78713-7819. ISBN (cloth hardcover) 0-292-75125-7, U.S $39.--; ISBN (paper softcover) 0-292-75126-5, U.S. $17.95. 474 The authors describe their book, Texas Mushrooms, as a pictorial guide for beginners to common Texas mushrooms and fungi. It contains 222 major entries, each with at least one color photograph. Nearly 40% of the descriptive part of this book covers just the boletes, Amanita, Lactarius, and Russula. Several of the taxa included here cannot be found illustrated in other field guides. Many of the photographs are quite striking and the book is well bound and printed on good paper. Also, this is the first mushroom field guide of Texas. In fact, despite its short-comings, some particular to this book, some common to the field guide genre, Texas Mushrooms is a welcome addition to the growing literature of regional guides, and can give residents and non-residents alike a sense of this particular geographically defined mycoflora that is unavailable from any of the national field guides. | The book is divided into five parts: and introduction to mushroom hunting in Texas; how to use the book to identify mushrooms (illustrations, charts, etc.); a section on mushroom toxins by Robert Harvey, M.D.; a section on cooking and eating wild mushrooms (with 10 recipes); and the bulk of the book, some 271 pages, with text and accompanying photographs on the same page. There is a short list of books for further study and an index. There is also an appendix of spore data, despite the authors’ stated preference for choosing "species that can be clearly distinguished by field characteristics," and their statement that "when (microscopic) examination is the only way to distinguish between closely related species, (they) provided the name of the most commonly found form" (p. 3). Because field guides are part of taxonomic literature but are not scientific literature (usually no voucher collections are cited, etc.), the most pressing question becomes: what is an acceptable level of imprecision, and for whom? Texas Mushrooms lacks authors' citations, specific references to technical descriptions with each major entry, and voucher collections. Nowhere does it state that all the photographs were even taken in Texas, and, despite this guide's reliance on the photography for species determinations, some of the photos are not diagnostic, or they are murky, or badly off-color, or misdetermined. The species descriptions are less than adequate, and they are wrong in detail for some entries and neglect essential field characters of others. In fact, many species descriptions seem intended to cover whole complexes or even genera rather than species. There are also examples of the same species represented by different entries (different names and photographs). The section (and scattered notes) on mushroom poisoning is badly misleading and in error in several instances. G. Lincoff, The New York Botanical Garden, Bronx, New York, U.S.A. MY COTAXON Volume XLVI, pp. 475- 479 April-June 1993 NOTICE: A NEW COMPUTER PROGRAM FOR THE IDENTIFICATION OF LICHEN SUBSTANCES ESTHER MIETZSCH Institut fiir Physiologie, Abt. Biokybernetik, Ruhr-Universitat Bochum, D-4630 Bochum, Germany H.THORSTEN LUMBSCH Botanisches Institut, Universitat Essen, Postfach 103 764, D-4300 Essen 1, Germany JOHN A. ELIX Department of Chemistry, The Faculties, Australian National University, GPO Box 4, Canberra, ACT 2601, Australia Abstract: A program for the identification of lichen substances based on TLC RF values, HPLC RI-values, colours of developed spots under visible and UV light, as well as the results of lichen spot tests has been developed for Microsoft WINDOWS. The program is based on a similar program for Apple Macintosh computers. It accepts RF values from six standard solvents, a RI-value from a gradient HPLC with reverse phase column and a range of possible colours for an unlimited number of TLC spots and will generate answers within a user defined error range. A list of synonyms of lichen substances is included as well as an RF and RI-value calculator. Introduction Chemical investigations of lichens form an integral part of all serious taxonomic studies on lichen-forming fungi today. About 550 lichen substances are known, most of them specific to lichens. The significance of chemical characters in the taxonomy and systematics of lichenized fungi has often been discussed (W. CULBERSON 1970, 1986, C. CULBERSON 1986, CULBERSON & CULBERSON 1970, HAWKSWORTH 1976, BRODO 1978, 1986, EGAN 1986). However, even the more experienced lichenologist 476 encouters TLC spots or HPLC peaks that are unfamiliar and difficult to identify. Lists of RF-values in standardized TLC solvent systems (CULBERSON & KRISTINSSON 1970, CULBERSON 1972, CULBERSON, & AMMANN 1979, ELIx et al. 1987b) and RI-values of gradient elution HPLC (FEIGE et al. 1993) have been published. These lists, however, cannot be kept current since new substances are continuing to be discovered and a large bulk of scattered literature exists on the RF-values of recently described substances. Associated information such as the colour of the spots, mass spectra data etc. is not available in lists, but is scattered within the literature. In an effort to make the information more readily accessible, and to keep such a library of information current as more and more lichen metabolites are identified and characterised, a data bank suitable for storing such information was developed by ELIX et al. (1987a, 1988) for APPLE MACINTOSH computers. Discussion In the past it has been very difficult to produce a user-friendly menu environment in the Dos system. However, with the improvement in personal computers and the introduction of the WINDOWS system this situation has changed. Moreover powerful computer equipment is now available at reasonable prices. The design of the menu environment and windows of the new program resembles that of the MACTABOLITES program, but three further options were added (MIETZSCH et al. 1992). These are a "Calculate" menu to calculate RF- and RI-values, "Synonyms", which gives a list of synonyms and the appropriate names in current use and a "Help" menu. The substance classes and the HPLC RI-values are additional features of WINTABOLITES. The program makes full use of the user friendliness of the WINDOWS system as illustrated in Figure 1. The program accepts RF-data from six standard solvents, an RI-value for a gradient HPLC with reverse phase column and a range of possible colours for an unlimited number of TLC spots. A list of possible answers is generated within a user defined error range of RF/RI-values and colours. The data set operates primarily on RF-values obtained by TLC on silica gel in any number of six standard solvent systems as well as the gradient HPLC system. Thus listed along with the name of each lichen substance is the following information: 1. The TLC RF-values of the compound in six standard solvent systems: A, toluene/ dioxane/ acetic acid (180 : 45 : 5) B, hexane/ diethyl ether/ formic acid (130 : 80 : 20) B’, hexane/ methy!] tert-butyl ether/ formic acid (140 : 72 : 18) C, toluene/ acetic acid (170 : 30) E, cyclohexane/ ethyl acetate (75 : 25) G, toluene/ ethyl acetate/ formic acid (139 : 83 : 8) 2. The HPLC RI-value from the gradient system described by FEIGE et al. (1993) for phenolic compounds. 3. Whether the TLC spot is coloured in visible (natural) light and/or detectable in 477 short wavelength ultraviolet light. 4. The colours of the TLC spot in both visible (natural) and long wave length ultra- violet light after spraying with H2SO4 and charring. 5. The colour of the TLC spot after spraying with Archers solution. 6. The results of medullary spot tests with K, C, KC, PD (when a compound gives a K + or C + reaction then KC has been omitted unless a different colour is produced by KC). 7. Three major peaks in the mass spectrum plus molecular ion (if observed). 8. The substance class. =z Wintabolites File Edit Data Search Calculate Synonyms Help | 6-O-Methyithiophanic BIOSYNTHETICALLY RELATED COMPOUNDS 65 x 71 60 2.5, 7-Trichloro-3-O-methylnorlichexanthone 72 x 55 66 3-0-Methylasemone lisohypertatolic acid 5,7-Dichloro-3-o-methyinorlichexanthone lsoarthothelin Thiophanic acid Thuringione 12,4-Dichlorolichexant 78 x «676 O75 iHyperlatolic acid 53 x 74 52 2,5, ?-Trichlorolichexanthone V: + UV: + 87 x 74 85 58 x 54 Fig. 1: WINTABOLITES - Compounds Biosynthetically Related to 3-O-Methylthio- phanic acid; Screen Display. 9. Up to nine biosynthetically related compounds. 10. Notes, a one hundred character long message containing additional information regarding the characteristics of the particular substance. However, it is not necessary to have all such information about an unknown spot - the search program will operate on as little information as a single RF or RI-value. Qualifying information simply reduces the number of possible answers generated. 478 The program was written in TURBO PASCAL for WINDOWS (Borland) using object- oriented programming techniques. The size of the source code is 66 KB. As all data are kept in the main memory by using dynamic data structures during the running of the program, a very high speed of retrieval is achieved. After the program has started, the data are loaded from the data file and all records are sorted into a binary tree which ensures quick access to the data (WIRTH 1983). At the same time seven arrays are build up, there being a separate array for each solvent system and one for the HPLC. The arrays are indexed by the RF-value or the RI-value of the compounds and the elements of the arrays each contain a dynamic list of pointers to all the records which have an RF or RI of that value for that solvent. When a search is started all the lists in the array of the first solvent system from RF - error index to RF + error index of the first spot are checked and all records, where the additional information given in agreement, are regarded as possible answers. The number of data sets as well as the number of spots to be examined is unlimited. WINTABOLITES is a WINDOWS application. Its use requires WINDOWS 3.0 or 3.1. the computer must have a 80286 or higher (i.e. 80386 or 80486) CPU and at least 2 MB RAM. A VGA graphic card and monitor is recommended. Availibility of the program: The program and an illustrated manual are available from the German authors for a price of US $ 149. Private persons not attached to an institution may ask for a reduced price of US $ 79. Updates will be distributed to the WINTABOLITES users yearly for DM 10 to 15,- if they wish to receive them. We wish to thank Dr. Siegfried Huneck (Halle) who kindly reviewed this manuscript. References BRODO, I.M. (1978): Changing concepts regarding chemical diversity in lichens. Lichenologist 10: 1-11. BRODO, I.M. (1986): Interpreting chemical variation in lichens for systematic purposes. Bryologist 89: 132-138. CULBERSON, C.F. (1972): Improved conditions and new data for the identification of lichen products by a standardized thin-layer chromatographic method. J. Chromatogr. 72: 113-125, CULBERSON, C.F. (1986): Biogenetic relationships of the lichen substances in the framework of systematics. Bryologist 89: 91-98. CULBERSON, C.F. & K. AMMANN (1979): Standardmethode zur Diinnschichtchromato- graphie von Flechtensubstanzen. Herzogia 5: 1-24. CULBERSON, C.F. & H.D. KRISTINSSON (1970): A standardized method for the identifi- cation of lichen products. J. Chromatogr. 46: 85-93. CULBERSON, W.L. (1970): Chemosystematics and ecology of lichen-forming fungi. Ann. Rev. Ecol. Syst. 1: 153-170. CULBERSON, W.L. (1986): Chemistry and sibling speciation in the lichen-forming fungi: ecological and biological considerations. Bryologist 89: 123-131. CULBERSON, W.L. & C.F. CULBERSON (1970): A phylogenetic view of the chemical 479 evolution in the lichens. Bryologist 73: 1-31. EGAN, R.S. (1986): Correlations and non-correlations of chemical variation patterns with lichen morphology and geography. Bryologist 89: 99-110. ELIx, J. A., JOHNSTON, J., and PARKER, J. L. (1987a): Mactabolites. Users Manual. 2nd ed. Canberra. ELIx, J.A., J. JOHNSTON & J.L. PARKER (1987b): A catalogue of standardized thin layer chromatographic data and biosynthetic relationships for lichen substances. Australian National University, Canberra. ELIx, J.A., J. JOHNSTON & J.L. PARKER (1988): A computer program for the rapid identification of lichen substances. Mycotaxon 31: 89-99. FEIGE, G.B., LUMBSCH, H.T., HUNECK, S. & ELIX, J.A. (1992): A method for the identification of aromatic lichen substances by gradient high performance liquid chromatography. J. Chromatogr., in prep. HAWKSWORTH, D.L. (1976): Lichen Chemotaxonomy. S. 139-184. In: D.H. BROWN, D.L. HAWKSWORTH & R.H. BAILEY (Hrsg.): "Lichenology: Progress and Problems", Aca- demic Press, London. MIETZSCH, E., H.T. LUMBSCH & J.A. ELIX (1992): Wintabolites (Mactabolites for Windows) Users Manual. Universitat Essen. WIRTH, N. (1983): Algorithmen und Datenstrukturen. 3rd. edn. Stuttgart, Teubner Verlag. ie } “Tt: , See iy are \¢ ws ‘ 4 ‘ \Eat aah ght F b ee tare ml are Mere rt ee Pe ey PN tg gL et a dey Bt tes Ae % r if — @") Rate ee if e a4 ty 481 Author Index to Volume Forty-seven Abdullah, N. see Ho, Y. W., D.J.S. Barr, N. Abdullah, and S. Jalaludin see Ho, Barr, Abdullah, Jalaludin and Kudo Alvarez, Isabel F. Javier Parladé, James M. Trappe and Michael A. Castellano Hypogeous mycorrhizal fungi of Spain. 201-217 Bakalova, Ganka G. see Sameva and Bakalova Banerjee , Partha and Walter J. Sunberg. Three new species and a new variety of Pluteus from the United States. 389-394 Barr, D. J.S. see Ho, Y. W., D.J.S. Barr, N. Abdullah, and S. Jalaludin see Ho, Barr, Abdullah, Jalaludin and Kudo Blackwell , Will H. and Martha J. Powell. Taxonomic history of the oomycete genus Thraustotheca. 183-192 Boqueras , Montserrat and Paul Diederich. New or interesting lichenicolous fungi. 3. Karsteniomyces llimonae sp. nov. and Sclerococcum SeruSiauxii sp. nov. (Deuteromycotina). 425-431 Castellano, Michael A. see Alvarez, Parladé, Trappe and Castellano Corlett, Michael Taxonomic studies in the genus Mycosphaerella. 2. Notes on some additional species occurring on Brassicaceae. 131-146 Diederich, Paul see Boqueras and Diederich Eicker, Albert , J. V. Van Greuning and Derek A. Reid. Amanita reidii - a new species from South Africa. 433-437 Elix, John A. New species in the lichen family Parmeliaceae (Ascomycotina) from Australia. 101-129 and Debra A. Venables. 4-O-Methyllividic acid, a new lichen depsidone. 275-281 Frizzi, G. see Pacioni, Frizzi, Miranda and Visca Galan, R. Moreno and Galan Gomez-Bolea, A. see Muntafiola-Cvetkovic and Gémez-Bolea Grosse-Brauckmann., Helga see Hjorstam and Grosse-Brauckmann Guzman, Gast6n see Ryvarden and Guzman Haines, John see Olsen, Haines and Sivertsen Halling, Roy E. Two new Crinipellinae (Tricholomataceae: Marasmiae) from South America. 379-385 Hess, W. W. see Strobel, Stierle, Stierle and Hess Hjortstam, Kurt and Leif Ryvarden. Two new South American species of Corticiaceae (Basidiomycetes) with amyloid spores. 81-85 and Helga Grosse-Brauckmann. Two new species of Cristinia (Basidiomycotina, ~ Aphyllophorales) and a survey of the genus. 405-410 Ho, Y. W., D. J. S. Barr, N. Abdullah, and S. Jalaludin. Anaeromyces, and earlier name for Ruminomyces. 283-284 and H. Kudo. A new species of Piromyces from the rumen of deer in Malaysia. 285-293 _____and§&. C. Jong. Phytophthora hibernalis and P. syringae. 439-460 Ialongo, Marco T. Biostatistical characterization of the genus Oidium. 193-199 Jalaludin, S. see Ho, Barr, Abdullah, and Jalaludin see Ho, Barr, Abdullah, Jalaludin and Kudo Jayachandra , J. A. see Muthumary, Jaychandra and Preetha Jong , Shung-Chang and Candace McManus. Computer coding of strain features of the genus Rhizopus.. 161-176 see Ho and Jong Ju, Y..-M. Felipe San Martin Gonzalez and Jack D. Rogers. Three xylariaceous fungi with scolecosporous conidia. 219-228 Kohn, L. M. Book Reviews. 467-473 482 Kudo, H. see Ho, Barr, Abdullah, Jalaludin and Kudo Kuthubutheen, A. J. see McKenzie and Kuthubutheen McKenzie, E. H. C. and A. J. Kuthubutheen. Dematiaceous Hyphomycetes on Freycinetia (Pandanaceae). 4. Cryptophiale. 87-92 McManus, Candace see Jong and McManus Miller, Orson K. A new Cortinarius from Southeastern Texas. 461-466 Miranda, M. see Pacioni, Frizzi, Miranda and Visca Moreno, G. and R. Galan Hydnangium pila Pat., an older name for Martellia mediterranea Moreno, Galan & Montecchi. 157- 159 Morgan-Jones, G. see Phelps and Morgan-Jones Muntanola-Cvetkovic, M. and A. Gémez-Bolea. Chaetospermum chaetosporum (Coelomycetes). First record from the Iberian Peninsula. 59-65 Muthumary, J., J. A. Jayachandra and M. Bhagavathy Preetha. Development of conidiomata in the Phyllosticta state of Guignardia mangiferae Roy and observations on the fine structure of the conidium. 147-155 Nograsek, A. see Scheuer and Nograsek Olsen, Sigurd, John Haines and Sigmund Sivertsen. Three new hyaloscyphaceous fungi from Norway and Greenland. 177-182 Pacioni, G. , G. Frizzi, M. Miranda and C. Visca. Genetics of a Tuber aestivum population (Ascomycotina, Tuberales). 93-100 Parladé, Javier see Alvarez, Parladé, Trappe and Castellano Phelps, Ryan A. and G. Morgan-Jones. Systematic and biological studies in the Balansieae and related anamorphs. III. Ascospore and macroconidial germination as a taxonomic criterion. 41-57 Powell, Martha J. see Blackwell and Powell Preetha, M. Bhagavathy see Muthumary, Jaychandra and Preetha Ramaley, Annette W. New fungi from Yucca: Planistromella yuccifoliorum, gen. et sp. nov., and its anamorph Kellermania yuccifoliorum, sp. nov. and Planistromella uniseptata, sp. nov., the teleomorph of Kellermania yuccigena. 259-274 Randlane, Tiina and Andres Saag. World list of cetrarioid lichens. 395-403 Reid. Derek A. see Eicker, Van Greuning and Reid Rogers, Jack D. see Ju, San Martin Gonzdélez and Rogers see San Martin Gonzalez and Rogers Ryvarden, Leif and Gast6n Guzman. New and interesting Polypores from Mexico. 1-23 see Hjortstam and Ryvarden Saag, Andres see Randlane and Saag Sameva , Ekaterina F. and Ganka G. Bakalova. Septoria thymi sp. nov. from Bulgaria. 387-388 San Martin Gonzalez , Felipe and Jack D. Rogers. Biscogniauxia and Camillea in Mexic« 229-258 see Ju, San Martin Gonzdlez and Rogers Scheuer, Christian . Lectotypification of Ophiobolus trichellus (Dothideales, Ascomycetes). 67-69 and A. Nograsek. Trematosphaeria pachycarpa and Herbampulla crassirostris gen. et spec. nov. (Ascomycetes) 415-424 Sivertsen, Sigmund see Olsen, Haines and Sivertsen Stierle, Andrea see Strobel, Stierle, Stierle and Hess Stierle, Don see Strobel, Stierle, Stierle and Hess Strobel, Gary , Andrea Stierle, Don Stierle and W. W. Hess. Taxomyces andreanae, a proposed new taxon for a bulbilliferous hyphymycete associated with Pacific Yew (Taxus brevifolia). 71-80 Sunberg, Walter J. see Banerjee and Sunberg Toyazaki , Noritsuma see Udagawa, Toyazaki and Tsubouchi Trappe, James M. see Alvarez, Parladé, Trappe and Castellano 483 Treu, R. Studies on Boletus section luridi. 367-377 Tsubouchi, Haruo see Udagawa, Toyazaki and Tsubouchi Udagawa, Shun-ichi , Noritsuma Toyazaki and Haruo Tsubouchi. Neosartorya primulina, anew species of food-borne ascomycetes. 359-366 Upadhyay, H. P. Comments on recent work on Ophiostoma and its synnematous anamorphs. 411-413 Van Greuning , J. V. see Eicker, Van Greuning and Reid Venables, Debra A. see Elix and Venables Visca, C. see Pacioni, Frizzi, Miranda and Visca Wong, Pak Yau. Type specimens of lichens and lichenicolous fungi in the Canadian Museum of Nature (CANL). 295-357 Wu, Chi-Guang. Glomales of Taiwan: III. A Compartative study of spore ontogeny in Sclerocystis (Glomaceae, Glomales). 25-39 Te eh are See rus ir aw aneurin chetalot pares “ ie aie we Aa ace A ia iti gph Nee (sea ie ay “” wan ka rye A Ths ay) & iN re ae a P “Daf: Midi oes ttt leh Mt a. . sits ingle kine ae hee * saphiny ae ear ware Pees. * ‘ ple nian ra rage hn Ae. Ah at ee 4 | spas eis iy : ny : an er eases thee Rul bi | » A Fras Mahe 2 si! we) hy, hd Senapang EO Mak ayer) Bayi! he oa ay aioe shel as ut Sega: ey nie’ wre Hers | ie uaa ts . Cait ane Ae alde Hl beh? a iy inter ata Meat ha i « ecw aM Ay av? iets Ae Ne | ue Pie hil in a ON) pie ra ee ange He cra men “ Bos hast te | ‘ ie mo ris 1 emf awe £, ny Sa eR: a te ie i ae My at oan uw ind Csi: a ie (hia d ota t ys Bacd Me My At hay a ihe yo ai) Aa bight Y "ii } i tite ‘ 5 “he Sta Et ei Reale 7: BROS tot 5s apres mes he 4 t : ' WE Bh 4 At 7) Hi; a re easy i ub Sipe ' . ae ‘ hey ; ‘ J x ” r ‘! ite Thy a Ve ® yt r ete Le mae ee og > i boty 1 Rn Ae é Hespught es Bag Rk ea pee! i Re fy Wig Gah is < Cah ada aca Uy eahntertal rualaes Aah had a me nad, enki: ied ie eat § I Mectivd Sit yhagaded he gcd ys tie ad Age Sn) Rat Pay ae Reece on ony pal » 1/¥p Pai r A YP) Hi " ites re alt wa mrtie mina Ne iad ; ay a" PASSE 1 E ai, vid eae tl a Ww Ts ag a senae a eee heed ‘eae an, Ae ae Le Rn a . af eat watt ate WAP, lea tii’ el i sre . eT i ay 1 NR A A Oa ean : eT eee ore pie Ry oe BEM aN NN eA KIN! cms HOY (eg 9) a Piacher ei Taitay RN he, REN Senge vers ven ss ee vA, heer hte bat Me a f) es ieee ‘Pe ankaliey | wy Uses ra i aN 4 i $i a pats Corte nna bet A (a bis je yoo ae ry ) | A) Cae ‘- nae maid ohne a in oui bonne ew) iy rh ay, 2 | ne 14) c ; A ‘ , 5 ha Po ¢. iy a ; fer dete Nha inh on MA Noes Ween fie ey i te a i Hi shasta A ee ve eiiples ties ’ wa ees al is SUA See oie saa in te ny ‘sy alte uty he ey a snip Laan ja (ote hue ma ’ i i Y, My Dd | : fi Lak cM tanathay nba et, Af Ce owiiy) gr vit Fae ashes anaes EOE : - eu - Pe ye AN 4 Mine aa, 7? NN aL j uh eh Pea yet eer ie tf th ( a te EN i o ; Aiaite v a ane vis oe ake “ | ey: a | or ie . ay ae ee eda | ne mish ve. yee et aan eel atk aime ain AT aig ges ee ry an a ane Mg uae ak 7 ae * a ar Pe oe Moly AAs ame tb lib he * one x pag JANE» . ne penne iy? ‘ NSS pacid eran’ Baty A nm ee ee INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME 47 This index includes the names of genera, infrageneric taxa, species, and infraspecific taxa. New names are in boldface, as are page numbers on which new taxa are proposed. Absidia 175 Acanthodochium 219 Acarospora canadensis 296 punae 296 Acaulospora scrobiculata 39 Achlya 183, 186, 189, 191 bispora 191 dubia 188-189 Acolium bolanderi 296 Acrostaphylus 233 Agaricus brunnescens 97 Ahtiana 395, 403 sphaerosporella 401 wallichiana 402 Albatrellus 23 subrubescens 2, 20 Alectoria ambigua 296 arctica 297, 312 cornicularioides 297 corymbosa 297 fuscescens 297 glabra 297 imshaugii 297 mexicana 297 pseudofuscens 350 sarmentosa ssp. sarmentosa 298 stigmata 297 Aleurodiscus 410 aurantius 85 exasperatus 81, 83-85 Allocetraria 395, 403 ambigua 396 cucullata 397, 399 isidiigera 398 nivalis 400 potaninii 400 stracheyi 398, 401 Alternaria 59 Amagdalaria consentiens var. japonica 298 continua 298 haidensis 298 Amanita 385, 433, 435-437 subg. Lepidella 436 alauda 436-437 phalloides 433, 435-437 f. phalloides 435 f. umbrina 435 reidii 433-437 verna 437 virosa 437 Amauroderma 8, 21 camerarium 3, 20 Amphinema 407 Amyloathelia amylacea 83 aspera 81-82 Amylosporus campbellii 3, 17 Anaeromyces 283 elegans 283-284 mucronatus 283-284 Anaptychia fragilissima 298 japonica var. reagens 298 Anisomeridium nyssaiegenum 299 Anomoporia albolutescens 3, 17 myceliosa 3 Anthostoma 220, 222 Anthostomella sinensis 223 Anthracothecium japonicum 298 thelomorphum 339 Antrodia gossypina 4, 17 lenis 7 malicola 4, 20 vaillantii 4, 20 Antrodiella 485 486 [Antrodiella] angulatopora 4, 17 versicutis 5, 17 Anzia afromontana 298 Aphanomyces 183, 192 Arthonia rufidula 306 Arthopyrenia submuriformis 298 willeyana 299 Asahinea 395, 402 chrysantha 397, 399, 401 scholanderi 399, 401 Ascochyta 148-150 cypericola 76 Aspergillus 365 sect. Fumigati 365 Aspicilia 318 quartzitica 299 Asteromella brassicae 132 Atkinsonella 41, 56-57 hypoxylon 43, 54, 56 var. texensis 56 texensis 43, 56-57 Aurificaria liteo-umbrina 5, 20 Bacidia colchica 299 finkii 300 lisowskii 299 nivalis 299 scutellifera 299 spirospora var. patagonica 299 vasakii 299 violascens 300 Baeomyces weberi 300 Bahianora poeltii 300 Bainieria 412 Balansia 41-43, 56-57 aristidae 41, 43-44, 46, 48-50, 56-57 atramentosa 56 claviceps 42, 57 epichloe 41, 43-44, 50, 52-54, 57 henningsiana 57 linearis 42 obtecta 42 strangulans 41, 57 Balansiopsis 56 Balsamia 214 vulgaris 203 Basidiobotrys 233, 241 Belemnospora 92 Bilimbia finkii 300 Biscogniauxia 226-227, 229-231, 233- 234, 238 atropunctata 231-232, 240 cinereo-lilacina 236 comedens 231 divergens var. macrospora 231, 233 exutans 238 fuscella 229, 231, 233 grenadensis 231, 236 mediterranea 232, 236 nummularia 231-232, 238-240 var. exutans 232, 238 var. merrillii 232, 238-239 var. nummularia 239 var. "pseudopachyloma" 232, 239- 240 sp. 231-232, 234, 236, 240 Blumeria graminis 197 Boletus 367-368, 375-376 sect. Luridi 367, 376-377, 385 calopus 376-377 piedmontensis 376 pseudo-olivaceus 367, 369-371, 374, 376 queletii 376 roseobadius 367, 371-374, 376 rufocinnamomeus 367, 372-373, 376 subluridellus 367, 369, 373-377 subvelutipes 367 Botanamphora 415, 419 pachycarpa 415-416, 419 Botryodiplodia theobromae 150 Botryosphaeria dothidea 271 Brevilegnia 184, 186, 188, 191-192 bispora 186, 191 unisperma 188 var. litoralis 188 Bryoria carlottae 300 cornicularioides 297 friabilis 300 furcellata 398 fuscescens 297 glabra 297 [Bryoria] pikei 300 pseudocapillaris 301 salazinica 301 spiralifera 301 trichodes subsp. americana 297 subsp. trichodes 343 Buellia excellens 301, 340 galapagona 301 imshaugii 301 multispora 301 rivas-martinezii 301 tephrodes 301 Buelliella eximia 302 physciicola 302 Buergenerula 271 Bulbothrix microscopica 101-102, 104 queenslandica 101, 126 subtabacina 101-102, 127 Burgoa 76 Byssocorticium 407 coprophilum 407 lutescens 407 Byssoloma amazonicum 302 anomalum 302 Caecomyces equi 293 Cainia graminis 422 Calicium adspersum ssp. australe 302 constrictum 302 cryptocroceum 302 curtisii var. splendidula 302 fuscipes 303 obscurum 303 parvum 303 Caloplaca bisagnonis 303 brattiae 303 britannica 303 citrina var. arcis 303 coralloides 334 fraxinea 303 hensseniana 303 487 litoricola 304 marina ssp. americana 304 sbarbaronis 304 sorediata var. tenuis 304 spotornonis 304 trabicola 304 Calyptralegnia 186 achlyoides 187 Camillea 227-229, 233, 241, 247, 254- phe: broomeiana 250 cyclisca 243 flosculosa 250 fossulata 236, 242-243, 250 gigaspora 242-243, 248 guzmanii 229, 243-244, 246, 248 hainesii 242, 246 hyalospora 246 labellum 242, 244, 246-247 magnifica 229, 234, 243, 246-247, 248 mexicana 229, 234, 242, 248, 250- 251 mucronata 242, 250-251 obularia 242, 251 punctulata 242, 250-251 stellata 242, 252 tinctor 243, 252 verruculospora 241, 253 Candelariella canadensis 304 faginea 304 hudsonica 304 lambii 304 plumbea 305 Canomaculina melanochaeta 101, 127 Canoparmelia 103, 105 herveyensis 101-102, 103-104 macrospora 103 owariensis 101, 127 pruinata 103 pustulescens 101, 127 raunkiaeri 102-103 subarida 101, 103-104 whinrayi 101, 104-105 Catapyrenium caeruleopulvinum 305 tuckermanii 312 Catillaria corymbosum 297 italica 305 488 [Catillaria] melanopotamica 305 delavayi 397 Ceratocystiopsis 411, 413 delisei 397-398 Ceratocystis 411-413 denticulata 397 davidsonii 412 dermatoidea 397 piceae 412 diffusa 397 sagmatospora 411 elenkinii 397 ulmi 412 endocrocea 398 Cetraria 395, 403 endoxantha 398 aculeata 396 endoxanthoides 398 agnata 396 epiphorella 398 alaskana 305, 396 ericetorum 397-399, 401 albopunctata 396 eriophylla 398 aleurites 396 everniella 398 alvarensis 396 fahlunensis 398 ambigua 396 fallax 398 americana 396 fastigiata 398 andrejevii 396, 401 fendleri 398 annae 396 f. coralligera 305 antarctica 396 formosana 398 arborialis 396 furcellata 398 arctica 396 gilva 398 arenaria 396 glauca 398 asahinae 396 globulans 398 atlantica 396 gracilenta 398 aurescens 396 halei 398 australiensis 305, 396 hepatizon 398, 400 bavarica 396 herrei 398 billardieri 396 hiascens 398 bohemica 396 hypotrachyna 398 braunsiana 396 iberica 305, 398 californica 396-397 idahoensis 398 canadensis 397 inermis 398 caperata 397 inflata 398 capitata 397 isidioidea 399 chicitae 397 islandica 399-400 chlorophylla 397 var. arborialis 305 chrysantha 397 japonica 399 ciliaris 397 juniperina 399 citrina 397 kamczatica 399 clarkii 397 komarovii 399 f. clarkii 397 kurodakensis 399 collata 397 kurokawae 399 commicta lacera 399 f. sorediella 401 lacunosa 399 commixta 397-398 laevigata 399 complicata 397 lanata 399 coralligera 305, 397 laureri 399 corallophora 397 laxa 399 corniculata 397 leucostigma 399, 401 corrugis 397 libertina 399 crispa 397 madreporiformis 399 cucullata 397 magnussonii 399 culbersonii 397-398 megaleia 399 daibuensis 397 melaloma 399-400 [Cetraria] merrillii 399 microphylla 399 microphyllica 399 muricata 399 nepalensis 399 nephromoides 399 nigricans 397, 399 nigricascens 396-397, 399-401 nipharga 399 nipponensis 399 nivalis 400 normoerica 400 norvegica 400 nova-zelandiae 400 nuda 400 oakesiana 400 ochrocarpa 400 odontella 400 orbata 400 ornata 400 pachysperma 400 pallescens 400 pallida 400 pallidula 400 perstraminea 400 pinastri 400 placorodia 400 platyna 400 platyphylla 400 platyphylloides 400 polyschiza 400 potaninii 400 pseudocomplicata 400 pubescens 400 ramulosa 400 rassadinae 400 regenerans 400 rhizophora 400 rhytidocarpa 401 richardsonii 401 rugosa 401 sanguinea 401 saviczii 401 scholanderi 401 scutata 401 sepincola 401 septentrionalis 401 sibirica 401 sikkimensis 401 simmonsii 401 sorediella 401 stenophylla 401 stracheyi 401 straminea 401 489 stuppea 401 stygia 401 subalpina 306, 396, 401 subfendleri 401 subscutata 401 sulphurea 401 tenuifolia 401 tenuissima 401 terrestris 401 teysmanni 401 thomsonii 402 thyreophora 402 tilesii 402 togashii 402 tristis 402 tubulosa 402 tuckermanii 402 ulophylla 402 ulophylloides 402 urceolata 402 viridis 402 wallichiana 402 weberi 402 xizangensis 402 yunnanensis 402 Cetrariopsis 395, 403 wallichiana 402 Cetrelia 395, 403 alaskana 305, 396 braunsiana 396 cetrarioides 397 chicitae 397 collata 397 davidiana 397 delavayana 397 isidiata 398 japonica 399 monachorum 399 nuda 400 olivetorum 396, 400-401 orientalis 400 pseudocollata 400 pseudolivetorum 400 sanguinea 399, 401 sinensis 401 Cetreliopsis 395 rhytidocarpa 399, 401 Chaenotheca brunneola 303 Chaenothecopsis nana 306 nigropedata 306 sagenidii 306 tasmanica 306 490 Chaetocalathus 379, 384-385 sect. Holocystis 384 liliputianus 384 magnus 379, 382-384 Chaetoconis 272 "Chaetonaevia archangelica" 180 Chaetospermella 62 Chaetospermum 59, 62-64 camelliae 63-64 carneum 63 chaetospermum 64 chaetosporum 59-60, 62-64 gelatinosum 62-63 tubercularioides 63 Charcotia rufidula 306 Cheiromycina petri 306 Chithramia 64 Choiromyces magnusii 204 meandriformis 204 venosus 204 Ciliochorella mangiferae 150 Ciliospora 62 Cladidium bolanderi 318 Cladina arbuscula subsp. beringiana 307 boliviana 307 conspicua 307 terrae-novae 309 f. cinerascens 309 stygia 309 Cladonia alpestris f. aberrans 307 anitae 307 arbuscula subsp. beringiana 307 boliviana 307 botrytes 307 convoluta var. vagans 307 crispata var. infundibulifera f. albopunctata 307 var. cetrariiformis 307 favillicola 307 gracilis subsp. vulnerata 307 hedbergii 308 kauaiensis 308 labradorica 308 libifera 308 magyarica 308 norvegica 308 patagonica 308 physodalica 308 poeciloclada 309 pollicum 308 rangiferina f. caerulescens 309 f. leucosticta 309 squamosa 308 subchordalis 309 terrae-novae 309 f. cinerascens 309 thomsonii 309 wainii 309 Cladorrhinum 74, 76 foecundissimum 74 Cladosporium 59, 146 herbarum 142 Clinostomum luteolum 309 vitellinum 309 Coccocarpia stellata 309 Coccotrema maritimum 310 Coelocaulon aculeatum 396-397 muricatum 399, 401 Coelopogon epiphorellum 398 Coleophoma cylindrospora 150 Collema leptaleum 310 mircoptychium 310 Collybia 385 subg. Rhodocollybia 385 Coltricia 13 Combea californica 342 Confertobasidium 410 Coniella fragariae 150 Coniocybe amabilis 310 Coniothecium 431 Conotremopsis weberiana 310 Coriolopsis [Coriolopsis] caperata 7 caperatiformis 7 floccosa 6, 20 rigida 6, 20 Coriolus ochrotinctellus 5 Cornicularia 403 normoerica 400, 402 Coronicium 410 Corticium amylaceum 83 coprophilum 407 filium 409 vallum 83 Cortinarius 461, 465 subg. Leprocybe 464-465 sect. Limonei 464-465 sect. Orellani 465 lewisii 462, 464-466 limonius 464-465 nothosaniosus 464-465 Creosphaeria 219, 222 pinea 222 riograndensis 222-223 sassafras 219, 222-223, 224, 226-227 verruculosa 222 Crinipellis 379, 382, 385 sect. Iopodinae 382 sect. Ionopodinae 382 hygrocybioides 382 insignis 382 purpurea 382 ticoi 379, 380-382 Cristinia 405-407, 409-410 artheniensis 405-406 brevicellularis 405-406 coprophila 405-407 filia 409 gallica 406-407 helvetica 405-406 mucida 407 rhenana 405-407, 408 sasae 405 sonorae 409 Crocicreas 178 Cryptophiale 87, 91-92 aristata 91 caudata 87-88, 91 enormis 91 kakombensis 91 novae-caledoniae 87, 89-91 secunda 91 Cryptophialoidea 91-92 Cytospora 491 turgida 220 Dacryobasidium 405, 407 Dactylina arctica 396 beringica 310 madreporiformis 399 ramulosa 400 Daedalea ambigua 8 elegans 6 microsticta 6, 19-20 Daldinia 254 Datronia caperata 7, 20 Delastria 214 rosea 204 Dentipellis fragilis 407 Dermatocarpon corticola 310 "corticolum" 310 gorzegnoense 310 vagans 310 Diatrype hypophloea 226 stigma 224 Dictyuchus 186 clavatus 184-185, 187 Didymella 271 Dilophospora alopecuri 64 Dimerella chiodectonoides 310 degeneri 311 frederici 311 isidiigera 311 pocsii 311 subdentata 311 subfallaciosa 311 usambarensis 311 Dingleya 217 Diplomitoporus lenis 7, 17 Diploschistes 129 actinostomus 312 conceptionis 312 hypoleucus 312 montevidensis 312 sbarbaronis 312 straminescens 312 Dirina californica 312 492 Dolichocarpus chilensis 312 Drechslera 59 Elaphomyces 201 anthracinus 204 granulatus 204 muricatus 201, 204 personii 204 trappei 204 Endocarpon mailae 312 tuckermanii 312 Endogone microcarpa 205 Entomopatella 62 Entophylyctis variabilis 289, 292 Eopyrenula intermedia 338 Ephelis 41-44, 48, 50, 52, 56 mexicana 42 Epichloe 41 typhina 43, 54, 56-57 Erioderma knightii 398 Erysiphe convolvuli 194, 197 Esslingeriana 395 idahoensis 398 Eutypa armeniaceae 228 Evernia perfragilis 297, 312 stracheyi 401 Everniastrum fragile 312 Fellhanera endopurpurea 312 Flavodon flavus 7, 17 Flavoparmelia 129 euplecta 105-106 kantvilasii 101, 105-106, 109 Fomes ohiensis 11 Fomitopsis cubensis 11 melliae 8, 17, 19 Fulgensia canariensis 313 Gaertneriomyces 292 Ganoderma 23 coffeatum 8, 19-20 Gautieria mexicana 204 morchelliformis 204, 214 Genea 214 klotzschii 204 verrucosa 205, 214 Geniculosporium 219 Geolegnia 186 Geopora arenicola 205 cervina 205 cooperi 205 foliacea 205 nicaensis 205 sumneriana 205 tenuis 205, 215 Glomus 25-26, 34, 201 ambisporum 26, 34, 39 constrictus 39 dimorphicum 34 heterosporum 25, 34, 39 macrocarpum 201, 205 microcarpum 205 Glyphis achariana 313 Gonohymenia cribellifera ssp. macrocarpa 313 lusitanica 313 undulata 313 Grammothele fuligo 8, 20 Graphilbum 411-412 Graphina aibonitensis 313 muscicola 313 subvirginalis 314 Graphis subvirginalis 313 Graphium 411-412 penicilloides 412 Guignardia mangiferae 147, 149 Gyalectidium colchicum 314 Gyalideopsis alnicola 314 kalbii 314 trapperi 314 [Gyalideopsis] vezdae 314 Gyrophoropsis dwaliensis 314 Haematomma inexpectatum 314 pustulatum 314 Hafellia fosteri 315 Heppia uruguayensis 315 Herbampulla 419, 422 crassirostris 415, 419, 421-422, 424 Heterodermia desertorum 315 stichensis 315 Heteropatella 272 Heterothecium pachycheilum 315 Homothecium sorediosum 315 Hubbsia californica 315 lumbricoides 315 Huilia circumnigrata 315 Humphreya 8 Hyalopesotum 411 Hyalopeziza 180 subg. Unguicularia 180 subg. Unguiculella 181 citrinus 206 muticus 201, 206 niveus 201, 207 olivaceus 207 populetorum 207 remyi 207 sp. No. 1 206 sp. No. 2 206 tener 201, 207 vulgaris 207 Hyperphyscia mobergii 316 Hyphoderma 410 Hyphodontia subs. Ellipsosporae 409 Hypocenomyce foveata 275, 277, 279 leucococca 316 Hypochniciellum 407 Hypogymnia austerodes 330 billardieri 281, 396 heterophylla 316 hokkaidensis 316 kiboensis 316 mollis 316 oceanica 316 rugosa 330 Hypotrachyna 275, 280 booralensis 101, 106, 109 densirhizinata 106-107 heterochroa 107 archangelica 177-178, 180 immaculata 275, 277-279 groenlandica 177-178, 181 livida 275, 277 raripila 180 osseoalba 275, 279 Hydnangium 158 proserpinensis 101, 107-109 aurantiacum 205 pustulifera 108 carneum 206 subpustulifera 101, 108-109 carotaecolor 205 pila 157-158, 206 Hydnocystis clausa 214 Hydnotrya tulasnei 206 Hydnum helveticum 405, 407 mucidum 407 Hydrothyria venosa 316 Hymenogaster 201 albus 201, 206 bulliardi 206 calosporus 206 cerebellum 206 virginica 331 Hypoxylon 219, 222, 226, 228, 230, 234, 238-241, 254-255 sect. Entoleuca 228 atropurpureum 234 bifrons 223 callostroma 223 cinereo-lilacinum 234, 236 cinereum 225-226 comedens 233 diathrauston 222 divergens 233 var. macrospora 233 flosculosum 250 fossulatum 250 fuscellum 233 494 {[Hypoxylon] glycyrrhiza 246, 254 grisea 226 hainesii 246 hemisphaericum 244, 248 hypophlaeum 224, 226 maculum 239 microplacum 224, 226-227, 232, 238 nummularium 231, 239 var. exutans 238 var. merrillii 238 var. pseudopachyloma 239 var. theissenii 241 punctulatum 251, 254 sassafras 222-223 tinctor 233 valsarioides 223 Hysterangium 201, 215 clathroides 201, 207 gardneri 201, 207 membranaceum 210 rickenii 207 separabile 207 stoloniferum var. rubescens 207 thwaitesii 207 Immersaria carbonoidea 322 Imshaugia evernica 101, 127 placorodia 397, 400 Incrustoporia nivea 9, 20 Inocybe geophylla 392 Inonotus hispidus 9, 20 munzii 9, 20 Involucrothele ligurica 316 Ionaspis ventosa 317 Jenmania osorioi 317 Jongiella 254 Juleila fallaciosa 335 Junghuhnia nitida 10, 20 Kalbiana brasiliensis 317 Karsteniomyces 425-426, 428 llimonae 425-426, 427, 429, 431 peltigerae 426 tuberculosus 426 Kellermania 65, 259, 265-266, 268, 272-274 anomala 265-266, 270, 273 yuccifoliorum 259-260, 262, 265- 266, 268-273 yuccigena 259, 265, 267-268, 270, 272-273 Kiessleriella 271 Kochiomyces 292 Labyrinthomyces 217 donkii 208, 215 "Laetinaevia archangelica" 181 Laetiporus sulphureus 14-15 Lecanactis chloroconia 317 premnea var. chloroconia 317 subdryophila 317 Lecania brialmontii 349 constricta 317 macrocarpa 317 sbarbaronis 317 Lecanora aeruginascens 317 austrooceanica 318 bolanderi 318 caesiorubella subsp. merrillii 318 caesiosora 318, 321 californica 318 christoi 318 cinerofusca var. appalachensis 318 concinna 318 "concinnum" 318 congesta 318 freyi 319 fuliginosa 319 geophila 321 gisleri 319 homalea 319 imshaugii 319 insolata 319 {[Lecanora] kutakii 319 laatokkaensis 343 luteovernalis 319 mellea 320 microbola 320 "olea" 320 oleae 320 opiniconensis 320 orae-frigidae 320, 322 pachysoma 320 perplexa 320 pseudopinguis 320 pycnocarpa 321 salicicola 321 soralifera 318, 321 superfluens 321 texana 321 thallophila 321 utahensis 321 weberi 321 Lecidea (Psora) 322 baffiniana 321 carbonoidea 322 grassiana 322 gypsicola 322 hyotheja 322 leproloma 322 patagonica 322 santensis 322 shushanii 322 sorediata 320, 322 ullrichii 323 Lempholemma dispansum 323 Leprocaulon arbuscula 343 pseudoarbuscula 347-348 subabicans 349 Leproloma diffusum var. chrysodetoides 323 Leptogium byssinum 323 caesiellum 323 Leptographium 412 Leptoguignardia 271 Leptosphaeria 178, 271, 416 hazslinskyana 415-416 pachycarpa 415-417, 424 Letharia togashii 323 wandelensis 323 Leucogaster floccosus 208, 215 Libertella 222 Lichen candelarius 323 cavernosus 397 salazinus 323 Lichenoconium parasiticum 323 Lichenostigma maureri 324 Lichina macrospora 324 polycarpa 324 rosulans 324 Lindquistia 219 Lobaria fulva 324 fuscotomentosa 324 gyrophorica 324 oregana 350 tuberculata 324 Loculotuber 201 gennadi 201, 208, 214 Lopadostoma 219 495 turgidum 219-220, 222, 226-227 var. minus 220 Loweporus 11 Macrophomina phaseolina 148-150 Marasmius hygrocybioides 382 Maronella laricina 324 Martellia mediterranea 157, 208 mistiformis 157 pila 157 Masonhalea 395 richardsonii 401 Mastigonema 62 Mazosia pilosa 325 pseudobambusae 325 Megalospora foersteri 325 pachycheila 315 Melanaria macounii 325 var. meizotoca 325 Melanelia 403 albertana 328 stygia 401 496 [Melanelia] tominii 330 trabeculata 331 Melanogaster ambiguus 208 broomeianus 208 variegatus 208 Melaspilea fuscolimitata 325 Melogramma 422 Micarea muhrii 325 vulpinaris 325 Microdochium fusarioides 449 Microporellus obovatus 10, 20 Microsphaeria euonymi-japonici 195 trifolii 195 Microthelia atramentea 325 Milospium graphideorum 430 Minimedusa 76 Monochaetiella hyparrheniae 272 Monochaetiellopsis 272, 274 Mucor 176 Muhria urceolata 325 Mycocalicium fuscipes 303 Mycoglaena quercicola 325 Mycosphaerella 131-133, 135, 142, 144-146 allicina 142 alyssi 132, 144-145 brassicicola 131-132, 144-146 carniolica 133, 144-145 confinis 133-134, 144-145 cruciferarum 131, 134, 137, 144-145 denigrans 131, 135-136, 144-145 densa 131, 136, 144-145 hambergii 136, 144-145 isatidis 137, 144-145 napicola 144-145 pachyasca 131, 137-139, 144-145 pashkiensis 131, 139-140, 144-145 phlomidicola 135 pyrenaica 131, 136, 141, 144-145 tassiana 131, 133-134, 136, 138-140, 142-145 vesicariae-arcticae 142-145 Myriogenospora 56 atramentosa 41-44, 46, 54, 56-57 Nectria 43, 47 aquifolii 43 coryli 43 Neocallimastix 292 frontalis 289, 292-293 variabilis 293 Neofuscelia 112, 115 delisea 112 imitatricoides 101, 110-111, 113 imitatrix 110-111 incantata 110 kondininensis 101, 110, 113 loxodella 111-112 luteonotata 112, 114 melanobarbatica 115 parasitica 101, 111-113 parviloba 114-115 pulla 111 pyrenaica 112 scabrella 114 scabrosina 101, 112-114 subbarbatica 101, 114, 117 subincerta 112, 128 verisidiosa 111-112 Neosartorya 359-360, 364-365 fischeri 364-365 hiratsukae 359, 364, 366 primulina 359-360, 361-362, 364 pseudofischeri 365 Nephroma americana 396 chubutense 326 kuenhnemannii 326 lepidophyllum f. "hypomelaena" 326 f. hypomelaenum 326 occultum 326 Nephromopsis 395, 403 asahinae 396 californica 397 delavayi 397 ectocarpisma 397, 399 endocrocea 397-398 endoxanthoides 398 globulans 398 isidioidea 399 komarovii 399-400 kurokawae 399 laureri 397, 399 laxa 397, 399 [Nephromopsis] morrisonicola 399 nipponensis 399 ornata 398, 400 pallescens 397, 400-401 pseudocomplicata 400 rugosa 401 stracheyi 401 yunnanensis 402 Neuropogon rohmederi 326 f. ushuaiensis 326 trachycarpus f. elatior 326 Nodulisporium 219, 229-230, 233-234, 252 Nozemia syringae 439 Nummularia 230, 241, 254 broomeiana 254 fuscella 233-234 "Nummularia" grisea 226 hypophloea 226 sp. 219, 224, 226-227 viridis 219, 224, 226 Numulariola 233, 238, 254 atropunctata 232 broomeiana 251 exutans 238 mediterranea 236 pseudopachyloma 240 Ochrolechia gowardii 326 juvenalis 326 montana 327 pseudopallescens 327 subisidiata 327 trochophora var. pruinirosella 327 tuckermanii 327 yasudae 327 Octaviania asterosperma 208 pila 206 Oidium 193-194, 196-197, 199 Oligoporus fragilis 10, 20 Omphalodina bullata 327 Opegrapha albidoatra 327 astraea 327 497 pertusariicola 327 subabnormis 328 Ophiobolus trichellus 67-68 Ophiochaeta trichella 68 Ophiostoma 411, 413 davidsonii 412 sparsum 412 Oropogon colibor 328 diffractaicus 328 mexicanus 328 Orpinomyces bovis 292 Ovularia syringae 439 Pachnodium 411 Paecilomyces 365 Pannaria leucostictoides 328 Papulaspora 76 Paraparmelia 129 alabamensis 328 lithophiloides 115 yamblaensis 101, 115, 117 Paraphaeosphaeria 271 Parmelaria 395 subthomsonii 401 thomsonii 402 Parmelia subg. Xanthoparmelia 128 alabamensis 328 albertana 328 ambigua var. halei 328 cetrarioides 397 chlorochroa 328 crinita f. varians 328 crowii 101, 116-117 cryptoxanthoides 329 dubia var. scrobiculata 329 elabens 329 erumpens 116 formosana 281 frondifera 329 frostii 398 gloriosa 329 haitiensis 127 hygrophila 329 498 [Parmelia] jejunga 118 livida 281 melanochaeta 127 monachorum 399 multipartita 126 olivetorum 400 omphalodes subsp. glacialis 329 owariensis 127 pacifica 329 physodes var. enteromorpha f. rugosa 329 f. subisidioides 330 pseudolivetorum 400 pseudotenuirima 116 pseudovirens 127 pustulescens 127 queenslandica 126 ralla 330 reptans 330 rubsecens 401 saximontana 330 semansiana 330 skultii 329 sphaerosporella 401 spinibarbis 330 squarrosa 330 subdistorta 330 subtabacina 127 sulcifera 331 trabeculata 331 virginica 331 xanthosorediata 127 Parmeliella concinna 331 granulata 331 Parmelina 118, 129, 425 conlabrosa 118-119 endoleuca 119 euplectina 101, 116-118 johnstoniae 101, 118-119, 121 labrosa 105 pastillifera 425, 428-429 pseudorelicina 116, 118 quercina 425-426, 429 tiliacea 425, 428, 430 Parmelinopsis evernica 127 minarum 119 protocetrarica 101, 119, 121 subambigua 328 Parmentaria astroidea 338 Parmotrema 402 demethylmicrophyllinicum 331 indicum 331 merrillii 120 perforatum 397 pseudovirens 101, 127 queenslandense 331 submerrillii 101, 120-121 subtinctorium 328 zollingeri 120 Peltigera 426 kristinssonii 331 malacea f. hasimotoi 332 pacifica 332 Peltula cylindrica 332 Penicillium 365 Perenniporia ohiensis 11, 20 tephropora 11, 17 Pertusaria glaucomela 333 gymnospora 332 macounii 325 neoscotica 332 panyrga 333 paramerae 332 santamonicae 332 suboculata 332 subpupillaris 332 subvelata 333 sulcata 333 victoriana 333 Pesotum 411-412 ulmi 411 Pestalotiopsis palmarum 150 Pestalozziella 65 Phaeocalicium asciiforme 333 curtisii 303 Phaeocollybia 385 Phaeolopsis veraecrucis 14 Phaeopeccania australiensis 333 hispanica 333 Phaeophyscia cernohorskyi 334 spinellosa 333 Phaeosphaeria 424 Phellinus 12 chryseus 11, 17 [Phellinus] contiguus 12, 17 ferrugineo-velutinus 12, 17 gilvus 19 grenadensis 12, 17 rhytiphloeus 12, 17 rimosus 13, 20 Phialocephala 412 Phialographium 411-412 sagmatosporae 411 Phloeophthora syringae 439 Phlyctis pulveracea 333 Phlyctochytrium arcticum 292 Phomatospora 422 Phylloporia chrysita 13, 20 frutica 13 spathulata 2, 13-14 veraecrucis 2, 14, 20 Phylloporus 385 boletinoides 385 Phyllopsora corallina 322 Phyllosticta 147, 149-150, 154 Physcia duplicorticata 333 hirsuta var. echinella 334 Phytophthora 163, 175, 440-441, 447- 452 cactorum 440, 448, 452 cinnamomi 446 citrophthora 449-450 crytogea 449 hibernalis 439-451, 453-457, 460 megasperma 445, 449 parasitica 446 porri 445, 450 syringae 439-457, 460 Picoa juniperi 208 Pilidiella quercicola 150 Pilophorus cereolus var. hallii 334 clavatus 334 Piptarthron 259, 265-266, 272-273 pluriloculare 274 uniloculare 259, 265, 274 Piptoporus soloniensis 14, 17 499 Piromonas communis 285-286, 293 minima 286 Piromyces 285-286, 289, 292 communis 286, 292 dumbonica 286, 293 mae 292-293 minutus 285-286, 287-288, 290, 292 rhizinflata 286, 293 Placodium coralloides 334 Placopsis alphoplacoides 334 var. clavidera 334 contortuplicata f. fuegiensis 334 dusenii 334 ; gelidoides 334 lateritioides 334 perrugosa f. activa 335 salazina 335 trachyderma 334 var. Clavifera 334 Placynthium petersii 337 Plagiocarpa septemseptata 335 Planistroma 259, 271, 273 obtusilunatum 265, 273-274 yuccigena 274 Planistromella 259-260, 271, 273 uniseptata 259, 264, 267-268, 271 yuccifoliorum 259-260, 261-262, 264-267, 271 Platismatia 395, 403 erosa 398, 400 formosana 398 glauca 398 herrei 398, 402 interrupta 398 lacunosa 397, 399 norvegica 399-400 regenerans 400 stenophylla 401 tuckermanii 396, 402 Platysma thomsonii 402 subperlatum 401 Piuteus 389-390, 394 sect. Celluloderma 389 sect. Hispidoderma 389 sect. Pluteus 389-390, 392 cervinus 390 500 [Pluteus] cinerascens 389-390 Psilolechia heterocystis 389-390, 392 leprosa 336 laricinus 389-390, 392 Psora magnus 390 cerebriformis 336 salicinus 393-394 montana 337 var. achloes 394 pacifica 337 var. americanus 389-390, 393-394 texana 337 var. salicinus 394 Psoroma Polyblastia internectens 337 cucurbitula 335 Psorotichia sbarbaronis 335 rimosa 336 Polyblastiopsis Pterygiopsis quercicola 335 australiensis 337 Polycauliona canariensis 337 coralligera 335 convoluta 337 Polyporus 13 Pterygium caperatus 7 petersii 337 elegans 15 Punctelia pargamenus 16 bolliana 329 pseudosulphureus 15 perreticulata 329 semipileatus 9 rudecta 397 soloniensis 15 semansiana 330 varius 15, 20 Pustularia veraecrucis 14 insignis 215 Polystictus 13 Pyrenastrum Poria 22 gemmeum 337 myceliosa 3 Pyrenopsis vaillantii 4 sojakii 338 vincta 15 Pyrenula Porina cinerella corruscans 335 var. quadriloculata 338 mangiferae 335 leucoplaca Porocyphus kalbarrensis 335 Porogramme fuligo 8 Porpidia carlottiana 335 thomsonii 336 Protoparmelia laricata 336 Pseudephebe pubescens 399-400 Pseudocyphellaria anomala 336 coriacea 341 exanthematica 336 hirsuta f. leucosticta 336 rainierensis 336 Pseudoidium 194 Pseudoparmelia aradensis 396 crystallicola 336 var. pluriloculata 338 macounii 338 maculata 338 neglecta 338 subsp. occidentalis 338 nitidella var. maculata 338 occidentalis 338 pseudobufonia 338 santensis 355 shirabeicola 338 thelomorpha 338 Pyroporellus subblinteus 5 Pythium 163, 175, 450 Pyxine daedalea 339 Radulodon 409 americanus 409 Radulum [Radulum] gallicum 407 mucidum 407 Ramalina americana 339 atlantica 339 bicolor 339 cactacearum 339 euxini 339 inflata 398 sideriza 339 superfraxinea 339 tenuis 339 tigrina 340 Ramonia intermedia 340 kandlerii 340 Reddellomyces 217 Relicina 128 Rhizocarpon alaxensis 340 atlanticum 340 barilochense 340 compositum 301, 340 constrictum 340 cumulatum 340 hensseniae 340 inimicum 340 renneri 341 Rhizophlyctis rosea 289, 292 Rhizoplaca bullata 327 Rhizopogon 201, 212 aestivus 209 colossus 215 luteolus 208 provincialis 209 roseolus 209, 213 rubescens 209 subareolatus 201, 209, 213 ventricisporus 201, 209 vinicolor 215 vulgaris 209 Rhizopus 161-163, 175-176 arrhizus 175 azygosporus 163 homothallicus 162 microsporus 162, 175-176 oryzae 163, 175-176 sexualis 162 stolonifer 162, 176 Rhodocybe 384 Ricasolia beckettii 341 Rigidoporus vinctus var. cinerea 15, 17 var. vinctus 15 Rimeliella haitiensis 101, 127 Rimularia fuscosora 341 Rinodina afghanica 341 ascociscana 341 calculiformis 341 conradii 342 dakotensis 341 humilis 341 milliaria f. obscura 341 oregana 342 pallida 342 sabulosa 342 Robillarda depazeoides 150 Rocella galapagoensis 342 Rocellina luteola 342 olivacea 342 Rosellinia diathrausta 222 linderae 223 prinicola 223 Rozites 385 Ruminomyces 283 elegans 283-284 Sagenidiopsis merrotsii 342 Saprolegnia 183, 192 Sarrameana tasmanica 342 Sawadaea 197 Schismatomma californicum 312 Schistophoron variabile 342 Schizopelte californica 342 Sclerococcum 425, 430-431 epiphytorum 430 serusiauxii 425, 428-430 sphaerale 430 Sclerocystis 25-26, 28, 30, 32, 34, 36, 38-39 501 502 [Sclerocystis] clavispora 25-28, 30, 32, Stachybotrys 92 36, 39 Stagonospora 271-272 coremioides 25-28, 30, 32, 34, 36, 38 gigantea 259, 271 liquidambaris 25-28, 30, 34, 36 Stephanocyclos rubiformis 25-27, 30, 32, 34, 36, 38 henssenianus 343 sinuosa 25-28, 30, 32, 34, 36, 38 Stereocaulon taiwanensis 25-28, 30, 32, 34, 36, 38 alpinum Scoleconectria var. erectum 343 cucurbitula 43 arbuscula Scolecosporiella 272 var. aberrans 343 Scutula 426 argodes 343 aggregata 426 argus 343 miliaris 426 var. stenospermum 344 Septoria 150, 387 thymi 387-388 azulense 344 botryosum Sepultaria f. depressum 344 arenicola 205 f. pygmaeum 344 arenosa 205 capense 344 foliacea 205 colensoi Setaria var. raegens 344 trichodes 343 condensatum Skeletocutis var. sorediatum 344 nivea 9 coralloides 344 Skyttea var. flabellatum 344 tephromelarum 343 var. occidentale 345 Sphaerella cornutum alyssi 132 var. corallizans 345 brassicicola 132 corticatulum carniolica 133 confinis 133-134, 136 var. hambergii 136 cruciferarum 134 denigrans 135 densa 136 napicola 137 pachyasca 137, 139 pyrenaica 141 tassiana 138, 142 vesicariae-arcticae 142 Sphaeria var. capense 344 var. complanatum 345 var. procerum 345-346 crambidiocephalum 345 dactylophyllum 344 var. flabellatum 344 var. occidentale 345 dendroides 345 denudatum var. umricola 345 var. "umbricolum" 345 depressum 344, 346 brassicicola 132 dusenii 345 callostroma 223-224 esterhuysenae 345 cruciferarum 134 evolutoides prinicola 223 var. paschaleoides 345 sassafras 222-224 exutum 346 turgida 220 fastigiatum Sphaerophorus var. dissolutum 346 notatus 343 ramulifer 343 Spizellomyces 289, 292 f. confluens 346 glabrum 344, 348 f. flabellans 347 Sporothrix 411-412 glareosum Squamaria var. brachyphylloides 346 configurata halei 346 f. diffracta 343 intermedium [Stereocaulon intermedium] f. compactum 346 japonicum var. subfastigiatum 346 var. tokioense 346 leprocauloides 346 melanopotamicum 345-346 meyeri f. tucumanum 346 montagneanum 347 myriocarpum var. altaicum 347 nanodes f. schadeanum 347 novoarbuscula 347 octomerelloides 347 octomerellum 347 papuanum 347 paschale 348 subsp. evolutoides f. laxatum 347 f. paschale 347 var. alpinum f. flabellans 347 var. serpens 348 v. evolutoides f. sorediatum 348 patagonicum 348 f. subirregulare 348 philippinense 348 pileatum 348 var. nipponicum 348 f. macrum 348 proximum var. gracilius 348 pseudoarbuscula 348 pseudomassartianum 349 pygmaeum 349 ramulosum 350 var. gracilius 348 var. pulvinare f. crebratum 349 salazinum 323 saxatile f. paschaleoides 346 f. sorediatum 348 spathuliferum f. congestum 346 f. pygmaeum 344 speciosum 349 var. surreptans 349 stenospermum 344 subcoralloides f. sorediascens 349 503 tennesseense 349 var. nigrofastigiatum 349 tomentosum var. Capitatum 350 var. compactum 349 verruciferum 349 var. surreptans 349 verruculigerum var. formosanum 350 var. verruculigerum 350 vesuvianum var. nodulosum f. umbricola 345 f. verrucosum 350 vimineum 350 weberi 350 Sticta lacera 399 oregana 350 Stioclettia 422 luzulina 422 Strangospora senecionis 350 Strigula submuriformis 299 Sulcaria badia 350 isidiifera 350 Talaromyces flavus 364-365 Taphrophila 69 Taxomyces 74 andreanae 71-72, 73-75, 77-80 Tephromela elixii 351 muscicola 351 nashii 351 septentrionalis 351 tropica 351 velloziae 351 Terfezia arenaria 209 berberiodora 210, 215 cadevalli 210 claveryi 210 hispanica 210 leonis 209 leptoderma 210 olbiensis 210 oligosperma 212, 215 pallidum 210 Thelidium 504 [Thelidium] umbilicatum 315 Thelomma mammosum 296 Thelotrema brasiliana 351 Thraustotheca 183-190, 192 achlyoides 186-188 caucasica 187, 191-192 clavata 183-190, 192 irregularis 188 primoachlya 183, 186-189 unisperma 186, 188 var. litoralis 188 var. unisperma 188 Thyronectria 43 balsamea 43 berolinensis 43 lamyi 43 Tiarosporella paludosa 150 Toninia arctica 351 bullata 352 ruginosa subsp. pacifica 352 var. andicola 352 tristis subsp. canadensis 352 subsp. pseudotabacina 352 Toxosporiopsis 65 Trametes malicola 4 Trapeliopsis hainanensis 352 pseudogranulosa 352 Trappea 215 darkeri 210 Trechispora farinacea 407 Trematosphaeria 415-417 pachycarpa 415-418 Trichaptum abietinus 16 biforme 16, 19-20 Tricholomopsis 385 Triparticalcar 292 arcticum 289 Tuber 93-97, 201 aestivum 93-97, 99-100, 210 asa 211 borchii 210, 215 brumale 211 excavatum 211 ferrugineum 212 foetidum 211 gennadii 201, 211 levissimum 201, 211 lutescens 210 maculatum 201, 211 malenconii 201, 211 maresa 211 melanosporum 94, 96-97, 211, 213 mesentericum 95, 97, 211-212 multimaculatum 201, 212, 214 nitidum 212 oligospermum 212 panniferum 212 puberulum 212 rufum 212 sinuosum 204 uncinatum 93-94, 96 Tubercularia chaetospora 62-63 Tubeufia 67, 69 trichella 67-68 Tuckermannopsis 395-396 americana 396, 398 aurescens 396 aokesiana 396 chlorophylla 397, 401-402 ciliaris 397 coralligera 397 fendleri 398 gilva 398 hepatizon 398 merrillii 396, 398-402 microphyllica 399 oakesiana 400 orbata 396-397, 400 pallidula 400 platyphylla 400 platyphylloides 400 sepincola 401 ulophylloides 402 Tylopilus 374, 377 Tyromyces fragilis 10 fumidiceps 16-17 galactinus 16-17 humeana 16-17 semipileatus 9 Umbilicaria badia 314 lambii 352 phaea 352 Urceolella 178 505 [Urceolella] aasii 177-178 tubulosus 396, 402 crispula 178 viridis 402 tuberculiformis 178 Urohendersonia pongamia 150 Wawea Usnea fruticulosa 355 acromelana 352 durietzii 353 eulychniae 353 Xanthoparmelia 129 kuehnemanrii 353 austroconstrictans 101, 120-122 neoguineensis 353 canobolasensis 101, 122-123, 125 neuropogonoides 353 chlorochroa 328 nidularis 353 constrictans 120, 122 perpusilla 326 cylindriloba 355 pseudoceratina 353 glareosa 122-123 tanzanica 353 hyposalazinica 101, 123, 125 trachycarpa 326 kiboensis 316 Ustulina 233 lipochlorochroa 355 masonii 101, 124-125 mollis 356 Verrucaria mougeotina 124 bagliettoi 353 multipartita 124, 126 bisagnoensis 353 reptans 330 boccana 354 rubrireagens 122 contardinis 354 subdistorta 331 diplotommoides 354 subnuda 123-124 durietzii 354 sulcifera 331 f. rhabdota 354 trirosea 101, 124-126 fuscella victoriana 124 f. omblensis 354 xanthofarinosa 101, 127 gorzegnoensis 354 xanthosorediata 101, 129 haeyrenii 354 Xanthopeltis "ha{umlat]yrenii" 354 rupicola 356 imperfecta 354 Xanthopsorella incompta 354 texana 337 lacustris 354 Xanthoria langhensis 355 candelaria 323, 335 latebrosoides 355 filsonii 356 omblensis 354 fulva 324 ornata 355 polycarpa pseudomacrostoma 355 var. maritima 356 putnae 355 sorediata 304 rapallensis 355 Xylaria 228, 254 santensis 355 Xylocladium 219, 229, 233, 241, 243, savonensis 355 247 Verticladiella 411-412 Vezdaea rheocarpa 355 Zahlbrucknerella Vuilleminia 85 patagonica 356 Vulpicida 395, 403 Zelleromyces canadensis 397 stephensii 157 juniperinus 399 pinastri 397, 400 tilesii 401-402 Tm Cn ny yd ee i) ae ori ae Ua a a Be eee ‘ rca? ye =r : 7 Rh iad is i Y i 507 ERRATA, VOLUME FORTY-SIX Pages 437-440 for.) Lema read Timah ERRATA, VOLUME FORTY-SEVEN Page 102 for _ P. subtabacina read B. subtabacina PUBLICATION DATES MYCOTAXON Volume 46: 1-508 (January-March) was published on February 4, 1993 Correction: MYCOTAXON Volume 45: 1-548 (October-December) was published on October 30, 1992 508 Reviewers, Volume Forty-Seven The Editors express their appreciation to the following individuals who have, prior to acceptance for publication, reviewed one or more of the papers appearing in this volume: T. Ahti J. Ammirati A. W. Archer allan ec) ‘Chang Ef Grane . Fogel . Guarro . Hawksworth R. A. Humber I. Karnefelt J. W. Kimbrough R R,E D. M. Langsam G.-z Ma D. Malloch G. Morgan-Jones G. M. Mueller A. Nawawi D. E. Padgett O. Petrini E. Punithalingam A. Rambelli L. Ryvarden E. Sérusiaux A. Strid F. A. Uecker G. Van der Westhuizen S. G. Vanev D. Verdon M. A. Vincent C. Walker R. Watling C. B. Wolfe EDITORS oF MYCOTAXON JEAN BOISE CARGILL, Editor-in-Chief Harvard University Herbaria 22 Divinity Avenue, Cambridge, MA 02138, USA ASSOCIATE EDITORS ROBERT DIRIG G. L. HENNEBERT LINDA M. KOHN Index Editor French Language Editor Book Review Editor Bailey Hortorium, Mann Library UCL, Place Croix du Sud 3 Botany Dept., Univ. of Toronto Cornell Univ., Ithaca, NY 14853 B-1348 Louvain-la-Neuve Mississauga, Ont. LSL 1C6 USA Belgium Canada MYCOTAXON is a quarterly journal devoted to all phases of mycological and lichenological taxonomy and nomenclature. It seeks to publish all papers within 5 months of submission, using photo-offset lithography. All articles are reviewed by specialists prior to acceptance. Publication is open to all persons. Papers may be in French or in English. Summaries in those or in any additional languages desired by the authors are given for longer articles. KEYWORDS are provided for each, article to facilitate library and computerized access. Printing is on high quality, acid-free, recycled book paper. EDITORIAL SERVICES & INFORMATION FOR PROSPECTIVE AUTHORS Authors prepare their own camera-ready copy after having received critical comments from pre-submission reviewers. Detailed Revised Instructions to Authors appeared in MYCOTAXON 26: 497-510 (1986). A copy of these instructions will be sent upon request to the Editor-in-Chief. Neither BJOPLATE transfer letters nor SPECIAL MANUSCRIPT PAPER are any longer available, and will not be restocked unless there is a strong demand from our authors. SUBSCRIPTION INFORMATION Each volume, beginning with volume 3, contains at least 512 pages, and consists of an irregular number of quarterly issues (rarely an additional issue, a Festschrift, may also be included in a volume). Each issue of MYCOTAXON varies in number of pages. Subscriptions are normally on a per volume basis, but subscribers may choose an annual basis to avoid frequent billing. Currently this would involve prepaying three volumes. Personal subscriptions are available at a substantially reduced rate for individuals who agree not to deposit their copies in another library than their personal one within 3 years of receipt. Address orders to the Mycotaxon Order Department, not to the Editors. Prices for the current volume are: USA Canada/Mexico Other Foreign (Air) REGULAR (multiuser) $60.00 $62.00 US $65.00 US PERSONAL (individual) $28.00 $30.00 US $35.00 US (All back volumes are still available.Volumes 1 through the latest complete volume are available at $25.00 per volume when shipped by surface mail, $40 per volume by air mail.) Place subscriptions through the Order Dept., MYCOTAXON, LTD., P.O. Box 264, Ithaca, NY 14851-0264, U.S.A. or through your agent. MYCOTAXON may also be obtained on a journal-exchange basis. This may be arranged with journals, institutions, or individuals who have difficulty in obtaining foreign currency. TWENTY-VOLUME CUMULATIVE INDICES, 1974-1984, & 1984-1991 MYCOTAXON CUMULATIVE INDEX FOR VOLUMES I-XX (1974-1984) by Richard P. Korf & Susan C. Gruff (ISBN 0-930845-00-5) is availabie at $17.50 postpaid, and MYCOTAXON CUMULATIVE INDEX FOR VOLUMES XXI-XL (1984-1991) by Richard P. Korf & Susan C. Gruff (ISBN 0-930845-01-3) is available at $30.00 postpaid, from MYCOTAXON, LTD., P.O. Box 264, Ithaca, NY 14851-0264, U.S.A. AVAILABILITY IN MICROFORM, TEAR SHEET, & PHOTOCOPY MYCOTAXON is also available in microfiche and in microfilm from University Microfilms, 300 North Zeeb Road, Ann Arbor, MI 48106, U.S.A., or 30-32 Mortimer Street, London WIN 7RA, England, from whom prices may be obtained. Tear sheets or photocopies of individual articles may be obtained through The Genuine Article™, 1.8.1., 3501 Market Street, Philadelphia, PA 19104, U.S.A., from whom prices may be obtained. CONTACTING MYCOTAXON'S EDITOR-IN-CHIEF BY E-MAIL OR BY FAX To reach the Editor-in-Chief regarding manuscripts, you may use this electronic mail INTERNET address: Cargill@HUH.Harvard.Edu. or you may FAX to Jean Cargill at (617) 495-9484. CONTACTING MYCOTAXON'S ORDER DEPARTMENT BY FAX To reach the Order Department for information or placing orders, you may FAX to Richard Korf at (607) 273-4357 ¥, 25 ex i an : cea) +h va \ a ve cet As eth ; Wee PAR cae 4 f dah te wie ‘ : ee ye | iy sonny ReLaOMNNC a y J a nds At | Ni ny gf)! # f ie ere ba i ie , ; my) oe x i Lyad i Nee a hy Ce , 0) al Age + Yeh i j Nie: LA eh aP me ae eT , Pn ae me ce) W 5 A Chis ud t ht hee ahi Y. ue ‘i % oh a hia ‘J PS ae -- a