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Phallus pygmaeus, a new minute species from a Bra

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Notes on Pestalotiopsis from southern China -siaxiang 1ang, Tong X

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Taxonomic and nomenclatural clarification algae neck rotting Botryti David S. Yohalem, Karsten Nielsen, & | Volvolepiota and Macrolepiota ~ = Macrolepiota vellosa, a new species fro ‘Else C. V ‘| A study of Peniophora species with simple-sepate hype "

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[Contents continued from front cover]

Type studies on Phellinus baumii and Phellinus linteus Young Woon Lim, Jin Sung Lee, & Hack Sung Jung Microfungi from Venezuela. A new species of Brachydesmiella, anew combination, and new records R. F. Castafieda Ruiz, T. Iturriaga, D. W. Minter, M. Saikawa, G. Vidal, & S. Velazquez-Noa A new species of Minimelanolocus and some hyphomycete records from rain forests in Brazil R. F. Castafieda Ruiz, J. Guarro, S. Velazquez-Noa, & J. Gené Notes on the genus Pithomyces (Hyphomycetes) from China Xiu-Guo Zhang & Tian-Yu Zhang Taxonomic studies of Stemphylium from China Xiu-Guo Zhang, Yue-Ming Wu & Tian-Yu Zhang A new species of Memnoniella De-Wei Li, Chin S. Yang, R. Haugland & S. Vesper Type studies in Agaricaceae - Chlorophyllum rachodes and allies Else C. Vellinga Two new species of the genus Pholiota from south India K. Natarajan & C. Ravindran Protousnea fibrillatae sp. nov. (Parmeliaceae, lichenized Ascomycota) from Argentina, southern South America _ S. Calvelo, E. Stocker-Wértgétter, S. Liberatore, & J. A. Elix New taxa in the lichen genus Pertusaria (lichenized Ascomycotina) from Thailand Sureeporn Jariangprasert, Alan W. Archer, John A. Elix & Vilaiwan Anusarnsunthorn A new species and new records of Gymnopilus from India K. Agretious Thomas, Laura Guzman-Davalos, & P. Manimohan

Microbotryum silybum sp. nov. (Microbotryales) Kalman Vanky & Dana Berner Yuea, a new genus in Xylariales Ove E. Eriksson New species of the genus Asterina from China III Bin Song

New species of Parmotrema and Relicina (Ascomycota, Parmeliaceae) from Thailand Kawinnart Noicharoen, Wetchasart Polyiam, Kansri Boonpragob, John A. Elix & Patricia A. Wolseley New and interesting inoperculate discomycetes from Korea Ain Raitviir & Hyeon-Dog Shin A new species of Memnoniella from India T. S. Keshava Prasad, L. G. Asha & D. J. Bhat Saprobic fungi on dead wild banana W. Photita, P. Lumyong, E. H. C. McKenzie, K. D. Hyde & S. Lumyong Notes on Hyphomycetes. XC. Fusicladosporium, a new genus for Cladosporium-like anamorphs of Venturia, and the pecan scab-inducing fungus E. Christopher Partridge & Gareth Morgan-Jones Notes on Hyphomycetes. XCI. Pseudoacrodictys, a novel genus for seven taxa formerly placed in Acrodictys William A. Baker & Gareth Morgan-Jones Studies on Lactarius: a new combination and two new species from Mexico Leticia Montoya & Victor M. Bandala Additions to our knowledge of the genus Geastrum (Phallales: Geastraceae in Brazil [uri Goulart Baseia, Maria Auxiliadora Queiroz Cavalcanti & Adauto Ivo Milanez Hyalophytophthora elongata, a new marine species from Taiwan H. H. Ho, H. S. Chang & S. H. Huang A checklist of mangrove-associated fungi, their geographical distribution and known host plants John Paul Schmit & Carol A. Shearer Book reviews and notices David L. Hawksworth

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AN INTERNATIONAL JOURNAL OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS

VOLUME LXXXvV, 2003

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EDITOR-IN-CHIEF

Pavel Lizon Department of Cryptogams, Institute of Botany Dubravska 14 SK-845 23 Bratislava, Slovakia

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TABLE OF CONTENTS, VOLUME EIGHTY-FIVE

Taxonomical studies in Ustilaginales. XXII Kalman Vanky Three interesting thermophilous taxa of Gymnopus (Basidiomycetes, Tricholomataceae): G. pubipes sp. nov., G. pubipes var. pallidopileatus var. nov. and G. dryophilus var. lanipes comb. nov. Antonio Ortega, Vladimir Antonin, & Fernando Esteve-Raventés Phallus pygmaeus, a new minute species from a Brazilian tropical rain forest luri Goulard Baseia, Tatiana Baptista Gibertoni, & Leonor Costa Maia Paraceratocladium polysetosum, a new record from Brazil Luis Fernando Pascholati Guzmao & Fabio Fernandes Barnes Biosystematics of the myxomycetes Didymium squamulosum, Physarum compressum, and Physarum melleum: additional isolates Jim Clark & Steven L. Stephenson Notes on Pestalotiopsis from southern China Jiaxiang Zhang, Tong Xu, & Qixin Ge Polypore fungi newly recorded in Taiwan Sheng-Hua Wu & Leif Ryvarden New species of Marasmius (Basidiomycetes, Tricholomataceae) from tropical Africa I. Sect. Epiphylli, Fusicystides, Globulares, Hygrometrici and Neosessiles Vladimir Antonin A new species of Pithomyces Xiu-Guo Zhang & Yue-Ming Wu Tuckermanella, a new cetrarioid genus in western North America Theodore L. Esslinger Two new species of the genus Entoloma from South India K. Natarajan & C. Ravidran A new rust fungus (Uredinales) on Penaeaceae: Uredo sarcocollae on Saltera sarcocolla Mechthilde Mennicken, Reinhard Berndt, & Franz Oberwinkler A list of discomycetes of China. Supplement II Wen-ying Zhuang Tricholoma zangii, a new name for T. quercicola M. Zang (Basidiomycetes: Tricholomataceae) Z.-M. Cao, Y.-J. Yao, & D. N. Pegler New and rare rust fungi (Uredinales) from Anatolia (Turkey) Zeliha Bahcecioglu & Halvor B. Gjaerum Taxonomic and nomenclatural clarification of the onion neck rotting Botrytis species David S. Yohalem, Karsten Nielsen, & Mogens Nicolaisen Volvolepiota and Macrolepiota Macrolepiota vellosa, a new species from China Else C. Vellinga & Zhu L. Yang A study of Peniophora species with simple-septate hyphae occuring in Taiwan Sheng-Hua Wu Type studies on Phellinus baumii and Phellinus linteus Young Woon Lim, Jin Sung Lee, & Hack Sung Jung Microfungi from Venezuela. A new species of Br achydesmiella, a new combination, and new records R. F. Castafieda Ruiz, T. Iturriaga, D. W. Minter, M. Saikawa, G. Vidal, & S. Velazquez- -Noa A new species of Minimelanolocus and some hyphomycete records from rain forests in Brazil R. F. Castafieda Ruiz, J. Guarro, S. Velazquez-Noa, & J. Gené Notes on the genus Pithomyces (Hyphomycetes) from China Xiu-Guo Zhang & Tian-Yu Zhang Taxonomic studies of Stemphylium from China Xiu-Guo Zhang, Yue-Ming Wu & Tian-Yu Zhang

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A new species of Memnoniella De-Wei Li, Chin S. Yang, R. Haugland & S. Vesper Type studies in Agaricaceae - Chlorophyllum rachodes and allies Else C. Vellinga - Two new species of the genus Pholiota from south India K. Natarajan & C. Ravindran Protousnea fibrillatae sp. nov. (Parmeliaceae, lichenized Ascomycota) from Argentina, southern South America S. Calvelo, E. Stocker-W6rtgétter, S. Liberatore, & J. A. Elix New taxa in the lichen genus Pertusaria (lichenized Ascomycotina) from Thailand Sureeporn Jariangprasert, Alan W. Archer, John A. Elix & Vilaiwan Anusarnsunthorn A new species and new records of Gymnopilus from India K. Agretious Thomas, Laura Guzman-Davalos, & P. Manimohan Microbotryum silybum sp. nov. (Microbotryales) Kalman Vanky & Dana Berner Yuea, a new genus in Xylariales Ove E. Eriksson New species of the genus Asterina from China II] Bin Song New species of Parmotrema and Relicina (Ascomycota, Parmeliaceae) from Thailand Kawinnart Noicharoen, Wetchasart Polyiam, Kansri Boonpragob, John A. Elix & Patricia A. Wolseley New and interesting inoperculate discomycetes from Korea Ain Raitviir & Hyeon-Dog Shin A new species of Memnoniella from India T. S. Keshava Prasad, L. G. Asha & D. J. Bhat Saprobic fungi on dead wild banana W. Photita, P. Lumyong, E. H. C. McKenzie, K. D. Hyde & S. Lumyong Notes on Hyphomycetes. XC. Fusicladosporium, a new genus for Cladosporium- like anamorphs of Venturia, and the pecan scab-inducing fungus E. Christopher Partridge & Gareth Morgan-Jones Notes on Hyphomycetes. XCI. Pseudoacrodictys, a novel genus for seven taxa formerly placed in Acrodictys William A. Baker & Gareth Morgan-Jones Studies on Lactarius: a new combination and two new species from Mexico Leticia Montoya & Victor M. Bandala Additions to our knowledge of the genus Geastrum (Phallales: Geastraceae in Brazil Iuri Goulart Baseia, Maria Auxiliadora Queiroz Cavalcanti ) & Adauto Ivo Milanez Hyalophytophthora elongata, a new marine species from Taiwan H. H. Ho, H. S. Chang & S. H. Huang A chercklist of mangrove-associated fungi, their geographical distribution and

known host plants John Paul Schmit & Carol A. Shearer Book reviews and notices David L. Hawksworth Nomenclatural novelties proposed in Mycotaxon volume 85 Errata

Instructions to Authors (updated January 2003) and Author’s Checklist Author index, volume eighty-five

Index to fungous and lichen taxa, volume eighty-five

Reviewers, Volume Eighty-Five

Publication Date for Volume Eighty-Four

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MYCOTAXON

Volume LXXXV, pp. 1-65 January-March 2003

TAXONOMICAL STUDIES ON USTILAGINALES. XXIIT.

KALMAN VANKY

Herbarium Ustilaginales Vanky (HUV) Gabriel-Biel-Str. 5, D-72076 Tiibingen, Germany e-mail: VANKY.K@cityinfonetz.de

Key words: smut fungi, Ustilaginomycetes, new species, new combinations, synonyms, excluded species, lectotypifications.

ABSTRACT

Fifteen NEW SPECIES of smut fungi are proposed: Entyloma frondosa, Macalpinomyces elionuri-tripsacoidis, M. ugandensis, Restiosporium dapsilanthi, Sporisorium absconditum,_ S. berndtii, S. compactum, S. cymbicum, S. divisum, S. eriochrysis, S. rhytachnes-rottboellioidis, S. sphacelatum, S. tothii, S. tristachyae-nodiglumis and S, zambianum. NEW NAME proposed: Sporisorium spegazzinii. NEW COMBINATIONS are: Macalpinomyces elymandrae, M. pretoriensis, Moreaua apicis, Sporisorium anadelphiae, S. argentinum, S. barberi, S. bengalense, S. cymbopogonis, S. densiflorum, S. lanigeri, S. mildbraedii, S. mutabile, S. nardi, S. panici- carthaginense, S. parodii, S. pennisetinum, S. spermoideum, S. urelytri, Tranzscheliella halophila, T. halophiloides, T. jacksonii, T. macrochloae, T. minima, T. sparti and T. ventanensis. EXCLUDED SPECIES are: Entyloma ameghinoi, E. nectrioide and Ustilago lycoperdospora. LECTOTYPES are designated for Entyloma chilense, Tilletia eremophila and Ustilago paraguayensis. Revision of different groups of smut fungi resulted in the establishment of twenty TAXONOMIC SYNONYMS. KEYS to eleven groups of smut fungi are compiled.

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THE GENUS TRANZSCHELIELLA (USTILAGINACEAE)

Lavrov (1936:29) proposed the genus 7ranzscheliella on the basis of two, small, supposed "cells" attached at opposite sides of the spores of 7. otophora. What Lavrov considered to be cells are actually the oppositely situated opercula (circularly broken parts of the thick exospore, standing off from the spore). This character alone is perhaps insufficient for recognising a separate genus. However, T. otophora, together with the closely related Ustilago hypodytes complex, differs markedly in several aspects from Ustilago hordei, the type of the genus Ustilago. Therefore, I am proposing to widen the circumscription of Tranzscheliella, including in it several related species.

Tranzscheliella Lavrov, emend. Vanky Sori superficial, naked, surrounding the distal internodes or the aborted inflorescence branches of Gramineae, which are covered by powdery spore masses. Infection systemic. Spores solitary, pigmented (brown), operculate or not, small (usually less than 8 pm long). Spore germination results in phragmobasidia. Host-parasite interaction by intracellular hyphae lacking interaction apparatus. Mature septa are poreless. Currently, 7ranzscheliella has three species: T. comburens (F. Ludw.) Vanky & McKenzie, type on Danthonia sp., T. hypodytes (Schlitdl.) Vanky & McKenzie, type on Elymus arenarius L., and the type of the genus, 7. williamsii (Griffiths) Dingley & Versluys, type on Stipa richardsonii Link (for synonyms, description and illustration see Vanky, 2002b:162-163). There are further Ustilago species which are better to be placed into this genus.

Tranzscheliella halophila (Speg.) Vanky, comb. nov. Basionym: Ustilago halophila Spegazzini, Anales Mus. Nac. Buenos Aires, Ser. 3, 1:58, 1902. Type on Distichlis scoparia (Kunth) Arechav., Argentina, Santa Fé, near Rufino, XII.1900, C. Spegazzini, LPS 3198. Isotype HUV 13520!

Ustilago stipae-barbatae Maire, 1917:139. Cintractia stipae-barbatae (Maire) Maire, in Maire & Werner, 1937:44. Lectotype (design. by Vanky, 1985:256) on Stipa barbata Desf., Morocco, Taourirt, 4.VI.1916, L. Ducellier, MPU, Herb. Maire 4226! Syntype on Stipa gigantea Link, Algeria, Miliana, near the top of Zaccar Rharbi, 12. V1I.1917, R. Maire, MPU, Herb. Maire 4810! (syn. by Vanky, 1988:372). For description and ill. see Vanky, 1994:359 & 404.

On Gramineae: Distichlis scoparia (Kunth) Arechav., D. spicata (L.) Greene, (D. maritima Raf.), D. stricta (Torr.) Rydb., Stipa barbata Desf., S. gigantea Link (Macrochloa arenaria (Brot.) Kunth), S. pennata L.; Africa, Asia, N. & S. America.

Tranzscheliella halophiloides (G.W. Fischer) Vanky, comb. nov. Basionym: Ustilago halophiloides G.W. Fischer, Res. Stud. State Coll. Wash. 20:9, 1952. Type on Distichlis stricta (Torr.) Rydb., USA, Oregon, Albert Lake, 28.VII.1950, G.W. Fischer & R. Sprague. Isotypes in Fischer, Gramin. smuts N. Amer. no. 215, HUV 9945!

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Sori surrounding the internodes, affecting all but the lowermost, first covered by a delicate, silvery membrane which flakes away disclosing the dark brown, powdery mass of spores. Spores globose, subglobose to ovoid, often depressed on one side, 4—6 um long, yellowish-brown; wall even, c. 0.5 wm thick, without polar thickenings, minutely verrucose, spore profile smooth to finely wavy.

On Gramineae: Distichlis stricta (Torr.) Rydb.; N. America (USA).

The isotype in HUV (no. 9945), besides 7. halophiloides, contains also plants with smooth spores, typical for 7. minima.

Tranzscheliella jacksonii (Zundel & Dunlap) Vanky, comb. nov. Basionym: Ustilago jacksonii Zundel & Dunlap, in Zundel, North American Flora 7:982, 1939b. Type on Stipa lettermanii Vasey, USA, Colorado, Tolland, 13. VII.1921, E. Bethel, BPI 162078, 162079.

Sori surrounding the upper internodes and the aborted inflorescence axis, covered only by the enveloping leaf sheaths, dark brown, powdery. Spores variable in shape and size, globose, subglobose, ovoid, oblong, 8-12 x 8—13.5(—16) um, olivaceous-brown; wall evenly thick (0.8—1 um), finely, densely verrucose, warts often fusing into irregular groups, spore profile nearly smooth to finely wavy.

On Gramineae: Stipa lettermanii Vasey; N. America (USA).

Tranzscheliella macrochloae (Pat.) Vanky, comb. nov.

Basionym: Ustilago macrochloae Patouillard, Bull. Soc. Mycol. France 22:199, 1906. Sphacelotheca macrochloae (Pat.) Maire, 1916:279. Type on Stipa tenacissima L., Tunisia, El Haffey, 25.11.1891, N. Patouillard 112, FH. Isotypes PC, HUV 14996!

For description see Vanky, 1994:365 & 415 (as Ustilago macrochloae).

On Gramineae: Stipa tenacissima L. (Macrochloa tenacissima (L.) Kunth); N. Africa (Algeria, Morocco, Tunisia).

Tranzscheliella minima (Arthur) Vanky, comb. nov. Basionym: Ustilago minima Arthur, lowa Agr. Coll. Dept. Bot. Bull. 1884:172, 1884. Type on Oryzopsis cuspidata Benth. ex Vasey (= O. hymenoides (Roem. & Schult.) Ricker), USA, Iowa, Ames.

Sphacelotheca valesiaca Schellenberg, 1911:61. Type on Stipa pennata L., Switzerland, Sitten, 1901, A. Volkart.

Ustilago stipae Ciferri, 1931:52. Lectotype (design. by Hirschhorn, 1947:75) on Stipa spartea Trin., USA, Iowa, Ames, 9.VII.1892, F.C. Stewart. Isolectotypes in Seymour & Earle, Econ. fgi., Ser. C., no. 70, HUV 9662!

Sorosporium reverdattoanum Lavrov, 1934:86. Type on Lasiagrostis splendens (Trin.) Kunth (= Achnatherum splendens (Trin.) Nevski), Kazakhstan, Semipalatinsk Prov., Buran, 7. VII.1928, N.N. Lavrov, LE. Isotype HUV 1210!

For description and illustrations see Vanky, 1994:366 & 418 (as Ustilago minima). :

On Gramineae: Achnatherum splendens (Trin.) Nevski (Stipa splendens Trin.,

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Lasiagrostis splendens (Trin.) Kunth), Oryzopsis hymenoides (Roem. & Schult.) Ricker (O. cuspidata Benth. ex Vasey), Sitanion hystrix (Nutt.) J.G. Smith, Stipa blackii C.E. Hubb., S. dasyphylla (Lindem.) Trautv., S. eminens Cav., S. joannis Celak., S. mollis R. Br., S. occidentalis Thurb., S. pennata L., S. pulcherrima C. Koch, S. spartea Trin., S. stuposa Hughes, S. verticillata Nees ex Spreng.; cosmopolitan. Tranzscheliella sparti (Massenot) Vanky, comb. nov.

Basionym: Ustilago sparti Massenot, in Guyot, Malengon & Massenot, Rev. Pathol. Vég. Entomol. Agric. France 34:215, 1955. Lectotype (design. by Vanky, 1994:376) on Lygeum spartum L., Tunisia, near Hadjeb-el-Aioun, 1.X.1953, L. Guyot, PC. Isolectotype HUV 15938!

For description and illustrations see Vanky, 1994:376 & 434 (as Ustilago sparti).

On Gramineae: Lygeum spartum L.; N. Africa (Algeria, Tunisia).

Tranzscheliella ventanensis (Hirschh.) Vanky, stat. et comb. nov. Basionym: Ustilago jacksonii Zundel & Dunlap var. ventanensis Hirschhorn, Notas Mus. La Plata, Bot. 8:171, 1943. Type on Stipa sp., Argentina, Prov. Buenos Aires, Sierra de la Ventana, La Pileta, 5.X1.1941, A.L. Cabrera 7343, Herb. E. Hirschhorn. Isotypes LPS 21560, HUV 15445!

Sori surrounding the upper internodes and the aborted inflorescence, naked, dark brown, powdery. Spores globose to ellipsoidal, 5-7 x 5.5—8(—-9) um, yellowish-brown; wall even, 1-1.5 um thick, finely, densely verrucose, spore profile wavy, in SEM warts often confluent into irregular groups.

On Gramineae: Stipa sp.; S. America (Argentina).

Key to the Tranzscheliella species

1. Spores 8—13.5(—16) wm long................ RESTS GeO Ne ae T. jacksonii Spores: smaller! gies yes ORE IN LRP RINE pant My ane Pere 2 2 SPOres WithicoOnspicuoUs Operculartrni sede ees T. williamsii Spores without conspicuous opercula but "polar caps" may be present............ 3 SU SPOTES VETTUCOSEN 2) Le. ee oh re Pre reer AR nee 4 Spores smooth, punctate or punctate-verruculOse.............ecceesecereeseeeeteeeteeenees ) 4) Spores'4—6 tum longi yee a ne eee T. halophiloides mSPOres D15--O(-9) Lem LONG eR CRs ae ee een ean ree tee T. ventanensis 5; Peridtum around the sor presenti) Wii rere ee eee ete 6 iPReriditm absent an ay. eae rane etc te ey er RL SO nes 8 6. Spores 3.5—6 um long, polar caps absent .............0... cc ccececcceeeeeeseees T. minima ~ Spores 5.5-8 um long, polar caps may be present.....................00008 Pe 7 Te Sporessmooth Pare. area are cies ate eee T. halophila =" SPOTes DUNCAtEKVEITUCUIOSE. 1) crater fierttes te rciicdiy ete eomeee T. macrochloae S$: Spores'2:5-4:5(-5) um jong ae ee T. comburens ~YOPOTES AATLET: 52 Ht. LC cea ee ieren Manta Se cn se ee pee ee 9

9. Spores 3.5—7 um long, polar caps may be present. Sori on culms . 7. hypodytes Spores 5.5—8 um long, polar caps absent. Sori in the inflorescence...... T. sparti

THE ENTYLOMA SPECIES OF BIDENS (COMPOSITAE)

Several Entyloma species have been published on Bidens. The oldest name is fk. guaraniticum Spegazzini (1884b), followed by /. bidentis Hennings. (1895) and &£. bidentis Spegazzini (1899). Saccardo and Sydow discovered the homonymy and renamed the fungus as /. spegazzinii Sacc. & P. Sydow (in Saccardo, 1902). Sawada (1922, n.v.), described an Entyloma on Bidens pilosa L. from Japan as E. bidentis Sawada (later homonym, not Hennings, 1895, not Spegazzini, 1899). Later, Sawada (1943:43) himself attributed the fungus to E. bidentis Henn., but Ling (1953b:344) considered it to be #. guaraniticum Speg. Ciferri (1928:38), published (invalidly, ICBN 34.1) the name of E. incertum Cif., "n. sp. ad interim", without’ any description (nom. nud.), on Bidens chrysanthemoides Michx. (= 8. laevis (L.) B.S.P.). North American authors often treated the Hntyloma species of Bidens (e.g. B. frondosa L., B. laevis (L.) B.S.P., B. tenuisecta Gray, B. vulgata Greene) under the collective name of /'. compositarum Farlow (comp. G.W. Fischer, 1953:85). The following four Entyloma, including a new species, could be recognised on Bidens:

1. Entyloma bidentis Hennings, in Engler, 1895:49. Type on Bidens pilosa L., Tanganyika Territory [= Tanzania], Mt. Kilimanjaro, Marangu, alt. 1500 m, V.1894, G. Volkens 2283, BR!

Sori on the leaves forming first whitish, later yellow to brown, flat, rounded to angular spots, 1—5 mm in diameter or larger by confluence. Spores embedded in the leaf tissue, globose, subglobose, ellipsoidal to irregular, 9-14 x 10-16 um, subhyaline to pale yellowish-brown, content granular; wall even to -slightly uneven, !—1.5(—2.5) um thick, smooth.

On Bidens bipinnata L., B. chilensis DC, (B. leucantha Willd.), B. pilosa L.; cosmopolitan in the tropics and subtropics.

2. Entyloma guaraniticum Spegazzini, 1884b:127. Type on “Araliaceae ?" (= Bidens pilosa L.), Paraguay, Guarapi, XII.1882, B. Balansa 3731, LPS 3361! Isotypes in Roumeg., Fgi. gall. exs. no. 4031, HUV 1030!

Sori on the leaves, first as whitish, rounded spots, later yellowish or light brown, bullate, cup-shaped, hypertrophied areas, 1-5 mm in diam. With age, the host tissues in the central part of the sori become dark brown, necrotic and the sori may appear perforated. Spores embedded in the host tissue, subglobose, ellipsoidal to irregular, with one or two flattened sides, (8—)9—-13 x 9-16 um, subhyaline to pale yellow, content granular; wall uneven, 1-3(-4) um wide, smooth.

On Bidens bipinnata L., B. pilosa L. and its var. radiata Schultz Bip.; N., C. & S. America, S. Africa, Australasia (Papua New Guinea).

E. guaraniticum was often mistaken for the more common £. bidentis Henn. However, besides spore wall thickness, the sorus morphology serves as a good

.

6

differentiating character: sori flat in E. bidentis, bullate, cup-shaped in EF. guaraniticum. ;

| 3. Entyloma spegazzinii Saccardo & P. Sydow, in Saccardo, 1902:376. Entyloma bidentis Spegazzini, 1899:211 (later homonym, non Hennings, 1895).

Type on Bidens bipinnata L., Argentina, Cérdoba, XII. 1887, C. Spegazzini.

Sori forming rounded, flat to slightly thickened leaf spots, 4 mm in diam. or larger by confluence, first pale, with indefinite margins, later brownish. Spores embedded in the leaf tissue, densely agglutinated, subglobose, ellipsoidal or irregular by mutual pressure, 9-16 x 12-20 wm, subhyaline, content homogenous; wall uneven, 2—5.5 um thick, smooth.

On Bidens bipinnata L., B. pilosa L., B. subalternans DC.: S. America, C. & S. Africa.

4, Entyloma frondosa Vanky, sp. nov. Typus in matrice Bidens frondosa L., Zambia, Southern Province, 162 km NE urbe Livingstone, 16°56'40" S, 26°48'12" E, alt. cca. 1270 m.s.m., 14. 1V.2001, leg. C., T. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19693, isotypi in Vanky, Ust. exs. no. 1142.

Sori sicut maculae rotundae, depressae foliorum, diametro 1-4 mm, primo pallide virides, serius in medio brunneae, in latere abaxiali foliorum proper conidia albidae. Sporae in telis foliorum innatae, satis dense situatae et variae, globosae, ovoideae usque irregulares, cum latere uno doplanato, (8—)9,5—14(—16) x 11—18(—20) pm, subhyalinae usque pallide flavidobrunneae; pariete inaequali, 1-3(-4) um crasso, levi. Anamorpha praesens.

Sori (Fig. 1) as rounded, flat, leaf spots, 1-4 mm in diameter, first pale green, later brown in the centre, whitish on the abaxial side of the leaves due to conidia. Spores (Fig. 5) embedded in the leaf tissue, rather crowded and variable, globose, ovoid to irregular, with a flattened side, (8-)9.5—14(—16) x 11—18(-20) um, subhyaline to pale yellowish-brown; wall uneven, 1—-3(-4) um thick, smooth. Anamorph present.

On Bidens frondosa L., C. Africa (Zambia). Probably, the North American "Entyloma compositarum Farlow" on B. frondosa represents also this species.

The main differentiating characters of the Entyloma species of Bidens are shown in a key. Key to the Entyloma species of Bidens

Loon bullates CUp-SDaNed Sas ne oie ee neo te ie oy E. guaraniticum (SOLE a Ale eh 1s be neh an AERA oh teetee Ar cebu ota NY eoekn es g tae 2 2. Spores 12-20 um long, wall 2-5.5 pm thick...........cccccccccseceeceeeee E. spegazzinii --xopores smaller wall thinneta as tater tern tla ANC WS Uke oes oi 3 3. Spores 11—18(—20) um long, wall 1-3(-4) um thick... E. frondosa

Spores 10-16 um long, wall 1-1.5(—2.5) um thick ...00000.0. E. bidentis

Fig. 1. Sori of Entyloma frondosa Vanky on a leaf of Bidens frondosa L. (type). Bar = 1 cm.

8

THE SMUT FUNGI OF PENNISETUM (GRAMINEAE)

This is the third and last part of the smut fungi of the subtribe Cenchrinae (see Vanky, 1998:330-337, and 2002a:391-398).

Pennisetum Rich., with c. 80 species throughout the tropics, is not sharply delimited from Cenchrus. On Pennisetum at least 26 smut fungi have been reported. Several of them are taxonomic synonyms (see below). In some cases the host plant is not a Pennisetum or the fungus is not a smut. For example, Tuburcinia penniseti J.N. Mishra (type on Pennisetum hordeoides (Lam.) Steud., India) is a rust fungus, Phakopsora apoda (Pat. & Har.) Mains (Vanky, 1999:159). The 13 recognised smut fungi of Pennisetum, including three new species, are:

1. Moesziomyces bullatus (J. Schroter) Vanky, 1977:133. Sorosporium bullatum J. Schréter, 1869:6. Type on Panicum crus-galli L., Germany. |

Tolyposporium senegalense Spegazzini, 1915:118. Type on Penicillaria typhoidea (L. Rich.) Fig. & de Not. (= Pennisetum glaucum (L.) R. Br.), Senegal, Dakar, 8.XII.1913, C. Spegazzini, LPS 4553!

For description, further synonyms and illustrations see Vanky, 1994: 163-166.

On species of Echinochloa, Leersia, Panicum, Paspalum, as well as on Pennisetum glaucum (L.) R. Br. (P. americanum (L.) Schum., P. typhoideum L. Rich.), and P. purpureum Schum.; cosmopolitan.

2. Sporisorium cenchri (Lagerh.) Vanky, 1985:114. Ustilago cenchri Lagerh., in Patouillard & Lagerheim, 1895:62. Type on Cenchrus sp., Ecuador, Riobamba, VIII.1891, G. Lagerheim, S!

Sorosporium chardonianum Zundel, 1942:125. Type on Pennisetum bambusiforme (Fourn.) Hemsley ex B.D. Jackson, Venezuela, near Merida, alt. 1600 m., 4. XII.1936, C.E. Chardon, Myc. Explor. Venezuela no. 1844 III, BPI 195142! (syn. here).

Sorosporium texanum Zundel, 1944:409. Type on Pennisetum nervosum (Nees) Trin., USA, Texas, Brownsville, Fort Brown, 23.XII.1942, Hansel 52794. Holotype BPI 180809!, isotypes BPI 180810, 195125. (syn. here).

For description, further synonyms and illustrations see Vanky, 1994:200 & 211, and 2002a:392.

On Cenchrus brownii Roem. & Schult., C. ciliaris L. (Pennisetum ciliare (L.) Link; Pennisetum cenchroides Rich.), C. echinatus L., C. gracillimus Nash, C. incertus M.A. Curtis (C. carolinianus Walt.), C. myosuroides H.B.K., C. pauciflorus Benth., C. pilosus H.B.K. (C. tribuloides auctt. eur. non L.), ?Panicum species, Pennisetum bambusiforme (Fourn.) Hemsley ex B.D. Jackson, and P. nervosum (Nees) Trin.; cosmopolitan.

Study of the types of Sorosporium chardonianum Zundel, and S. texcanum Zundel revealed that they are identical with Sporisorium cenchri.

Fig. 2. Sori of Sporisorium dubiosum (Zundel) Vanky producing stunting and witches' brooms on Cenchrus ciliaris L. (Pennisetum cenchroides Rich.; Namibia, HUV 19680). Habit. To the left a healthy inflorescence. Enlarged, three spikelets with sori of different developmental stages.

Bars = 1 cm for habit, 2 mm for the detail drawings.

3. Sporisorium divisum Vanky, sp. nov. Typus in matrice Pennisetum divisum (Forssk. ex Gmel.) Henr., W. Pakistan, Mankera prope Bhakkar, 21.X1.1976, leg. S. Ahmad. Holotypus in Herbario -Ustil. Vanky, HUV 9009!

Sori in spiculis, cylindrici, 1,5—-3 x 4-12 mm, primo peridio pallide brunneo cooperti, quo iregulariter rupto massam semiagglutinatam usque pulveream glomerulorum sporarum et columellas 4-6, sat crassas, apicaliter tenuiescentes, saepe breviter ramificatas ostendentes. Spiculae omnes eiusdem spicae infectae. Glomeruli sporarum globoidei, ellipsoidales, elongati usque irregulares, 30-80 x 40-90 um, atro-rufobrunnei, e sporis multis, faciliter separabilibus compositi. Sporae subglobosae, ellipsoidales usque subpolyedrice irregulares, 8-10,5 x 9-12 um, flavidobrunneae; pariete aequaliter crasso (cca. 1 um), conspicue levi usque leniter, dense punctato.

Sori (Fig. 10) in the spikelets, cylindrical, 1.5—3 x 4-12 mm, first covered by a pale brown peridium which ruptures irregularly disclosing the semiagglutinated to powdery mass of spore balls and 4—6, rather stout, apically narrowing, often shortly ramifying columellae. All spikelets in a spike are infected. Spore balls (Figs. 6, 7) globoid, ellipsoidal, elongated to irregular, 30-80 x 40—90 pm, dark reddish-brown, composed of numerous, easily separating spores. Spores (Figs. 6, 7) subglobose, ellipsoidal to subpolyhedrally irregular, 8-10.5 x 9-12 um, yellowish-brown; wall evenly thick (c. 1 pm), apparently smooth to finely, densely punctate.

On Pennisetum divisum (Forssk. ex Gmel.) Henr. (P. dichotomum (Forssk.) Delile); S. Asia (W. Pakistan). Known from the type collection only.

4, Sporisorium dubiosum (Zundel) Vanky, 2002a:396. Cintractia dubiosa Zundel, 1931:299. Type on Pennisetum sp. (= Cenchrus ciliaris L., teste K. Vanky), Kenya, Nairobi, 9.[X.1920, H.L. Shantz. Holotype in BPI 171508!, isotypes in BPI 171509 and 171510. For revised description see Vanky, 2002a:396. See also Fig. 2. On Cenchrus ciliaris L. (Pennisetum cenchroides Rich.); E. & S. Africa (Kenya, Namibia).

5. Sporisorium ehrenbergii (Kiihn) Vanky, 1990:270. Sorosporium ehrenbergii Kiihn, 1877:87. Tolyposporium ehrenbergii (Kithn) Patouillard, 1903:254. Type on Sorghum cernuum (Ard.) Host (= S. bicolor (L.) Moench), Egypt, Cairo.

For description and illustrations see Vanky, 1994:201 & 213.

On Sorghum bicolor (L.) Moench. Recently, a morphologically identical fungus was collected on Pennisetum glaucum (L.) R. Br. (P. typhoideum L. Rich.) in Eritrea (HUV 19512, det. K. Vanky), which is under investigation; Africa, Asia.

1]

6. Sporisorium penniseti (Rabenh.) Ershad, 1994:18. Ustilago penniseti Rabenhorst, 1871:18. Sphacelotheca penniseti (Rabenh.) Reichert, 1921:679. Neotype (design. by Vanky, 2000:211) on Pennisetum orientale L. C. Richard, "Taurus Catasnicus, 1865", C. Haussknecht, JE!

Ustilago pappiana Baccarini, 1906:272. Sorosporium pappianum (Bacc.) L. Ling, 1953a:192. Lectotype (design. by Ling, 1953a:192) on Pennisetum ruppellii Steud. (= P. setaceum (Forssk.) Chiov.), Eritrea-Amasen, Addi-Baré, along the river Mareb, 29.X.1902 (as "22.XI.1902"), A. Pappi, FT (RO 6168)! Syntype on Pennisetum orientale Rich. [var. altissimum Chiov.] (= P. setaceum), Eritrea, Ingal-Ceccaharat near Oculé-Cusai, alt. c. 1600 m, 22.11.1893 (as "22.1V.1893"), A. Pappi, FT (RO 4797)! (syn. here).

Sorosporium catharticum Maire, 1931:359. Sporisorium catharticum (Maire) Vanky, 1989:155. Type on Cenchrus catharticus Delile (= C. biflorus Roxb.), Algeria, South Sahara, near Tilemsi, coll. T. Monod 467, MPU! (syn. by Vanky, 2000:211).

Sphacelotheca panjabensis H. Sydow, in Sydow & Ahmad, 1939:442. Ustilago panjabensis (H. Sydow) L. Ling, 1950:76. Type on "Cenchrus biflorus Roxb.” (= C. setigerus Vahl, teste K. Vanky), India, Punjab, Sargodha, sine die, leg. S. Ahmad 33, HUV 17298! (syn. by Vanky, 2002a:397).

Sorosporium penniseti Mundkur, 1939:116. Type on Pennisetum ciliare (L.) Link (= Cenchrus ciliaris L.), India, Delhi, 3.VIII.1938, M.A. Khan. Holotype HCIO 7749!, isotype IMI. (syn. by Vanky, 2000:211).

Sphacelotheca_ stewartii Mundkur, 1944a:290. Type on Pennisetum flaccidum Grieseb., Kashmir, Baltistan, on the way from Kasurmik to Doghani, alt. 9000 ft, 16. VIII.1940, R.R. Stewart 20793. Holotype HCIO 10017, isotype HUV 17309! (syn. here).

Ustilago penniseti Rabenh. var. verruculosa Massenot, in Guyot, Malengon & Massenot, 1969:217. Type on Pennisetum ciliare (L.) Link, Morocco, near Atchana, Valley of Oued Guir, 12.VI.1932, G. Malencon 140. Holotype PC, isotype HUV 13658! (syn. by Vanky, 2000:211).

Sori in the spikelets, cylindrical, 1-1.5 x 2-7 mm, hidden by the glumes, covered with a greyish-brown peridium which flakes away exposing the semi- agglutinated to granular-powdery mass of spore balls and a central columella with a few, short, apical branches. All spikelets in a spike are infected. Spore balls globose, ovoid to ellipsoidal, 40-110 wm long, dark reddish-brown, composed of many spores which separate rather easily. Spores dimorphic; those on the periphery of the balls globose to ellipsoidal, 9-13 x (9—)10—13.5(-14.5) um, medium reddish-brown, finely and densely echinulate, those in the centre subpolyhedral, smaller (7-10 ym), lighter coloured and finely punctate or apparently smooth. Sterile cells of the peridium variable in shape and size, often elongated, 4-16 ym long, arranged in parallel rows, hyaline, smooth.

On Cenchrus biflorus Roxb. (C. catharticus Delile), C. ciliaris L. (Pennisetum ciliare (L.) Link), C. prieurii (Kunth) Maire (Pennisetum prieurti Kunth; P. cenchroides Rich.), C. setigerus Vahl (C. biflorus auct. non Roxb.), Pennisetum flaccidum Griseb., P. macrourum Trin., P. natalense Stapf,

12

P. orientale Rich. (P. fasciculatum Trin.), P. setaceum (Forssk.) Chiov. (P. orientale Rich. var. altissimum Chiov.; P. ruppellii Steud.); S. Europe (Mediterranean area), Africa (incl. Madagascar), S. Asia.

Study of the type of Ustilago pappiana Bacc., and that of Sphacelotheca stewartii Mundkur revealed that they are identical with, and synonyms of Sporisorium penniseti (Rabenh.) Ershad.

In the original description of Ustilago penniseti (Rabenhorst, 1871:18), the spores are globose, smooth, 9-11 um in diam. Baccarini, 1906:272, describing U. pappiana, wrote that it differs from U. penniseti in having larger spores (12-15 um) and no columella. The spore measurements given for U. penniseti are too small (comp. the description above) and those of U. pappiana too large (comp also Ling, 1953a:192). A columella, typical for Sporisorium penniseti, is also present in the sori of U. pappiana. Small variations (+ 1 pm) in the spore measurements may occur in samples from different host plant species or from different geographical areas.

7. Sporisorium penniseti-japonici (Hennings) Vanky, in Vanky & Guo, 1986(1987):234. Ustilago penniseti-japonici Hennings, 1904:140. Sphacelotheca penniseti- japonici (Henn.) S. Ito, 1935:91. Type on Pennisetum japonicum Trin. ex Spreng. (= P. alopecuroides (L.) Spreng.; P. purpurascens (Thunb.) Makino), Japan, Tokyo, 25.[X.1900, Kusano 443. Isotype in HUV 15794!

Ustilago penniseti Rabenh. var. tonkinensis Patouillard, 1890:57. Type on Pennisetum sp. (= P. alopecuroides (L.) Spreng., teste K. Vanky), Indochina [= North Vietnam], Tonkin Region, Mts. of Elephants, 1.1886, B. Balansa, FH! (syn. here).

Ustilago penniseti Saccardo, 1921:596. Type on Pennisetum sp., China, Fukien, Foochow, VI.1918, Skvortzow 3962 (later homonym, not Rabenh., 1871:18; syn. by Ling, 1953b:344).

Sori in the central part of the spikelets, infecting all spikelets in the inflorescence, ellipsoidal, 1.5—-2 x 5-7 mm, more or less hidden by the outermost floral envelopes and the bristles, first covered by a pale brown peridium which ruptures from its apex disclosing the first agglutinated later granular powdery mass of spore balls and spores surrounding several flagelliform columellae about the length of the sorus. Spore bails globoid, ellipsoidal, oblong, irregular, 40-80 x 50-100 um, reddish-brown, composed of tens of spores which separate easily by pressure. Spores subglobose, ovoid, ellipsoidal to subpolyhedrally irregular, 10.5—13.5(-14.5) x 12—15(-16) um, yellowish-brown; wall slightly uneven, 0.5—1 ym thick, densely verrucose-echinulate, spore profile undulate to finely serrulate. Sterile cells absent.

On Pennisetum alopecuroides (L.) Spreng. (P. purpurascens (Thunb.) Makino; P. japonicum Trin. ex Spreng.); E. Asia (Japan, N. Vietnam).

Study of the type of Ustilago penniseti Rabenh. var. tonkinensis Pat. revealed that the host plant is Pennisetum alopecuroides, and its smut identical with Sporisorium penniseti-japonici.

13

8. Sporisorium pennisetinum (S. Ahmad) Vanky, comb. nov. Basionym: Sphacelotheca pennisetina S. Ahmad, Mycol. Pap. 64:7, 1956. Type on Pennisetum flaccidum Grieseb., Pakistan, Swat State, Madian, 14. VII.1952, S. Ahmad 4596, IMI 57436!

Sori destroying the whole inflorescence, first covered by a peridium which ruptures to expose the dark brown, powdery mass of spores intermixed with groups of sterile cells, and numerous filiform columellae. Spores when mature single, subglobose, ovoid, ellipsoidal to slightly subpolyhedrally irregular, 9-13 x 10-14.5(—-16) um, yellowish-brown; wall even, c. 0.5 pm thick, densely, finely verrucose-echinulate; spore profile finely serrulate. Sterile cells in irregular groups, collapsed in old specimen, single cells 7-14 um long, subhyaline to pale yellowish-brown.

On Pennisetum flaccidum Grieseb.; S. Asia (Pakistan). Known only from the type collection.

9. Sporisorium sphacelatum Vanky, sp. nov.

Typus in matrice Pennisetum sphacelatum (Nees) Dur. & Schinz., South Africa, East Cape Province, Mts. Drakensberg, inter oppid. Barkley East et Lady Grey, 30°43'08" S, 27°20'50" E, alt. cca. 960 m, 22.X11.1996, leg. C. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19733, isotypi in Vanky, Ust. exs. no. 1145. Paratypus in matrice P. sphacelatum, Lesoto, Leribe Distr., Nkaobee Pass, cca. 15 km N Katse Dam Wall, 29°18'57" S, 28°32'10" E, alt. cca. 2400 m, 25.XI1.1996, leg. C. & K. Vanky, HUV 19734.

Sori inflorescentiam totam destruentes, longe cylindrici, 1-3 x 40-150.mm, a vagina folii supremi protrudentes, primo peridio cinerascentibrunneo cooperti, quo longitudinaliter rupto massam atrobrunneam, pulveream glomerulorum sporarum et sporarum cum columellis nonnullis, longis, filiformibus immixtarum ostendentes. Glomeruli sporarum globosi, ellipsoidales, elongati usque irregulares, 30-80(—90) x 40—150(—190) um, atro-flavidobrunnei usque subopaci, e sporis pluries decem usque pluries centum facile secedentibus compositi. Sporae subglobosae, ellipsoidales, raro parum irregulares, 5—6,5 x 5,5—7,5 pm, flavidobrunneae, ad latus unum pallidiores; pariete inaequaliter crasso, 0,5-0,8 wm, ad dimidium pallidum tenuiore, subtiliter, dense punctato- verruculoso; imago obliqua sporarum levis. Cellulae steriles non observatae.

Sori (Fig. 3) destroying the whole inflorescence, long cylindrical, 1-3 mm x 4-15 cm, protruding from the uppermost leaf sheath, at first covered by a greyish-brown peridium which ruptures longitudinally disclosing the dark brown, powdery mass of spore balls and spores intermixed with several, long, filiform columellae. Spore bails (Figs. 8, 9) globose, ellipsoidal, elongate to irregular, 30-80(—90) x 40-150(—190) pm, dark yellowish-brown to subopaque, composed of tens to hundreds of easily separating spores. Spores (Figs. 8, 9) subglobose, ellipsoidal, rarely slightly irregular, 56.5 x 5.5—7.5 um, yellowish-brown, paler on one side; wall unevenly thick, 0.5-0.8 ym, thinner on the pale half, finely, densely punctate-verruculose; spore profile smooth. Sterile cells not seen.

14

On Pennisetum sphacelatum (Nees) Dur. & Schinz.; S. Africa (Lesoto, South Africa). Known only from the type collections.

10. Sporisorium tothii Vanky, sp. nov. Typus in matrice Pennisetum glaucum (L.) R. Br., Sierra Leone, Central Province, Njala, XII.1929, leg. C.F. Deighton. Holotypus in Herbario Ustil. Vanky, HUV 7114, isotypus in BPI 192487.

Sori in ovariis nonnullis inflorescentiae eiusdem, obovoidei vel globoidei, pedicellati, 1,5-2,5 x 3-4,5 mm, cum apice brevi, acuto, peridio crasso, atrobrunneo cooperti, quo irregulariter rupto massam atrobrunneam, primo agelutinatam, deinde granulosopulveream glomerulorum sporarum columellam brevem, simplicem, complanatam circumdantem ostendentes. G/omeruli sporarum subglobosi, ellipsoidales, elongati, plerumque irregulares, 40-120 x 50—140(—190) um, atro-flavidobrunnei usque subopaci, e sporis pluries decem usque pluries centum, pressione separabilibus compositi. Sporae globosae, ovoideae, ellipsoidales usque parum trregulares, 7-9,5 x 8-11(-12) pm, flavidobrunneae; pariete aequali, cca. 0,5 pm crasso, levi. Cellulae steriles nullae.

Sori (Fig. 4) in some ovaries of an inflorescence, obovoid or globoid, 1.5—2.5 x 3—4.5 mm, with a pedicel and a short acute tip, covered by a thick, dark brown peridium which ruptures irregularly disclosing the first agglutinated, later granular-powdery, dark brown mass of spore balls surrounding a short, simple, flattened columella. Spore bails (Figs. 15, 16) subglobose, ellipsoidal, elongate, usually irregular, 40-120 x 50—140(-190) wm, dark yellowish-brown to subopaque, composed of tens to hundreds of spores which separate by pressure. Spores (Figs. 15, 16) globose, ovoid, ellipsoidal to slightly irregular, 7-9.5 x 8—11(—12) pm, yellowish-brown; wall even, c. 0.5 um thick, smooth. Sterile cells absent.

On Pennisetum glaucum (L.) R. Br. (P. typhoideum L. Rich), cultivated; W. Africa (Sierra Leone). Known only from the type collection.

Etymology: This fungus is named in the honour of my friend of more than half a century, Dr. Sandor Toth (83; Géd6ll6, Hungary), an exceptional, helpful, modest human being, who despite losing his arm in the Second World War, became the best contemporaneous specialist of Hungarian microfungi.

11. Tilletia barclayana (Bref.) Sacc. & P. Sydow, in Saccardo, 1899:422, s. lat.

Neovossia barclayana Brefeld, 1895:170. Type on Pennisetum triflorum Nees, India, Himachal Pradesh, Simla, coll. A. Barclay. Tilletia pennisetina H. Sydow, 1929:421. Type on Pennisetum

alopecuroides (L.) Spreng., China, Kiangsi Prov., Kuling, coll. F.L. Tai 2201. Tilletia ajrekari Mundkur, 1939:103. Type on Pennisetum typhoides Stapf, Indta, Ahmedabad, in experimental plots, 2.1X.1933, S.L. Ajrekar, HCIO. Neovossia macrospora Petrak, 1947:114. Type on Pennisetum japonicum Trinius ex Spreng. (= P. alopecuroides (L.) Spreng.), China, Chinkiang, X.1921, K. Kolthoff 33.

Fig. 3. Sori of Sporisorium sphacelatum Vanky in the inflorescence of

Pennisetum sphacelatum (Nees) Dur. & Schinz (type). Habit, and to the left a Bar = | cm.

healthy inflorescence. Fig. 4. Sori of Sporisorium tothii Vanky in some ovaries of Pennisetum glaucum (L.) R. Br. (type). Habit, and enlarged two spikelets, each with a sorus and a healthy seed. Bars = 1 cm for habit, 3 mm for the detail drawing.

16

Sori in some ovaries of an inflorescence, swollen, obovoid or ellipsoidal, 3-6 mm long, partly hidden by the glumes, first covered by a thick, dark brown membrane (pericarp) which ruptures from its apex disclosing the black, pulverulent mass of spores intermixed with sterile cells. Spores globose, subglobose to broadly ellipsoidal, 14-36 um long (including ornamentation), reddish-brown to subopaque, provided with densely situated, truncate, 1.5-4 um long warts ("scales"). Sterile cells variable in number, shape and size, globose, ellipsoidal or irregular, 10-30 pm in diameter, pale yellowish-brown, wall 1.5—4 um thick, laminated or not, sometimes with a mycelial appendage.

On Oryza sativa L., Panicum hians Ell., P. obtusum H.B.K., Paspalum distichum L., Pennisetum alopecuroides (L.) Spreng., Pe. orientale Rich., Pe. triflorum Nees, Pe. typhoides Stapf; Asia, Indonesia, N. America.

There are variations in the measurements and colour of the spores, in the height of spore ornamentation and number of sterile cells between different collections on the same or on different host plants. Tullis & Johnson (1952), demonstrated by artificial infections that this depends also on the infected host plant. Therefore, they concluded that Neovossia horrida, N. barclayana and Tilletia pennisetina represent the same species. This view was adopted also by Duran & Fischer (1961:35).

12. Ustilago bahuichivoensis Duran, 1970:1102. Type on Pennisetum pringlei Leeke, Mexico, Chihuahua, 1 mi. E. of Bahuichivo, 13.X.1969, J.F. Hennen. Holotype WSP 58559, isotype HUV 14508!

Ustilago beckeropsidis Zambettakis, 1979(1980):406. Type on Beckeropsis uniseta (Nees) Schum. (= Pennisetum unisetum (Nees) Benth.), Belgian Congo [= Democratic Rep. of Congo], 1913, H.J.R. Vanderyst. The holotype is probably lost, it is not in BR or in PC. Paratypes: Sierra Leone, 8.1.1951, T.S. Jones, IMI 45383!, and Sudan, 17.X.1952, J.K. Jackson, IMI 51127! (syn. here).

Sori (Fig. 11) on the inner surface of the uppermost, hypertrophied, hispid leaf sheaths, fusiform, 1-3 cm wide, 3~8 cm long, partly hidden by healthy leaf sheaths, at maturity fissure longitudinally exposing the dark chocolate-brown, powdery mass of spores. Spores ovoid, ellipsoidal, elongate, rarely subglobose, 5—7(-8) x 6.5-10.5(-11) pm, yellowish-brown; wall even, c. 0.5 ym thick, finely, densely punctate, spore profile smooth, in SEM finely verrucose.

On Pennisetum pringlei Leeke, and P. unisetum (Nees) Benth. (Beckeropsis uniseta (Nees) K. Schum.); N. America (Mexico), Africa (Congo, Sierra Leone, Sudan).

Fig. 5. Spores of Entyloma frondosa Vanky on Bidens frondosa L., in LM (type). Figs. 6, 7. Spores of Sporisorium divisum Vanky on Pennisetum divisum (Forssk. ex Gmel.) Henr., in LM and in SEM (type). Figs. 8, 9. Spore balls and spores of Sporisorium sphacelatum Vanky on Pennisetum sphacelatum (Nees) Dur. & Schinz, in LM and in SEM (type). Bars = 10 pm.

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18

Fig. 10. Sori of Sporisorium divisum Vanky, in the spikelets of Pennisetum divisum (Forssk. ex Gmel.) Henr. (type). Fig. 11. Sori of Ustilago bahuichivoensis Duran, on the inner surface of the uppermost, hypertrophied leaf sheath of Pennisetum pringlei Leeke (type). Fig. 12. Sori of Ustilago kamerunensis H. & P. Sydow, destroying all spikelets of an inflorescence of Pennisetum purpureum Schum. (Uganda, HUV 19977). Bars = 1 cm.

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Study of the paratypes of Ustilago beckeropsidis Zambett. revealed that it is identical with and a synonym of U. bahuichivoensis Duran.

13. Ustilago kamerunensis H. & P. Sydow, 1911:262. Type on Pennisetum sp., Cameroon, Dyutitsa's, XII.1908, Ledermann.

Sori (Fig. 12) destroying all the spikelets in the inflorescence, leaving intact only groups of long bristles and the inflorescence axis, 3-10 x 40-110 mm, first covered by a thin, greyish membrane which ruptures disclosing the blackish- brown, powdery mass of spores. Spores subglobose, ellipsoidal, often hemispherical with a more or less depressed side, in facial view 6.5—7 x 6.5—8 um, in lateral view 4—5 pm wide, yellowish-brown; wall c. 0.5 um, thinner on the flattened side, apparently smooth to extremely finely punctate.

On Pennisetum purpureum Schum. (P. benthamii Steud.), P. sp.; C. & E. Africa (Cameroon, Congo, Kenya, Rwanda, Uganda).

Not on Pennisetum Ustilago bonariensis Speg. (= Sporisorium panici-leucophaei (Bref.) M. Piepenbr.). Lectotype on Panicum spectabile Nees, Argentina. According to Zundel, 1953:54, Pennisetum tristachyum, one of the syntypes, seems to be a misdetermination.

Ustilago scheffleri H. & P. Sydow, 1911:262. Type on "Pennisetum inclusum Pilger", "Massaihochland, Lamuru" [?= Kenya, Kiambu Distr., Limuru], alt. 3000 m, 30.VI.1909, G. Scheffler 293. Isotypes in BPI 166232, 194468, K, M, HUV 12741!

The host plant is not a Pennisetum but a Panicum (teste K. Vanky; = P. coloratum L., det. H. Scholz, B). The fungus is a Sporisorium sp., maybe one of the about 20 Sporisorium species reported on Panicum. The sori destroy the whole inflorescence, are up to 3 cm long, first covered by a thin, greyish peridium which ruptures disclosing the dark brown, powdery mass of spores surrounding several, long, filiform columellae. Spores ellipsoidal, ovoid, more rarely globose or slightly irregular, (6.5—)7—9.5 x 8-12 um, yellowish-brown; wall even, 0.5—0.8 um thick, finely, densely punctate-verrucose, spore profile smooth to very finely wavy.

In the original description, the spores. are given as reticulate. Zundel (1953:197) studied the type and wrote that the spores are "smooth but contents with a semi-reticulate, granular pattern (not reticulate as originally given)".

Sporisorium pamparum (Speg.) Vanky, in Vanky & Guo, 1986(1987):234. Type on Setaria sp., Argentina. It was reported on Panicum, Pennisetum and Setaria species. Specimens on Pennisetum alopecuroides (L.) Spreng. represent Sporisorium penniseti-japonici (Henn.) Vanky. Those on Pennisetum flaccidum Griseb. (e.g. in Vanky & Guo, 1986(1987):234) are very similar to, if not identical with S. penniseti (Rabenh.) Ershad, which has priority. Further studies are needed to elucidate possible conspecificity of S. pamparum with S. penniseti.

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Key to the smut fungi of Pennisetum (M. = Moesziomyces, S. = Sporisorium, U. = Ustilago)

Le SONMTESTHICICR 1O.INe, OVATICS ie terrae en eae ares rh nary gee ene ere eer ge 2 ‘Sori not restricted.tovthe ovartess.. 0) eaadedeees see. csc aes Meten osc csavessatioles . 2. Spores single, 14-36 wm in diameter ............00. eee Tilletia barclayana —» spores, forming spore balls; smalleri jvc. )eie ile sateen Be et Rete 3 3. Spores in the balls mixed with sterile cells. Columella absent......... M. bullatus Spores in the balls not mixed with sterile cells. Columella present................... 4

4. Columella one, short. Spores all alike, 8—11(—12) um long, smooth.......§. tothii Columellae several, filiform. Spores dimorphic, 9.5—15 um long,

QULEIISNOTES VEITUCOSE Wier oie ee ada uy erat ean S. ehrenbergii 5(1). Sori on the inner surface of the uppermost leaf sheaths .. U. bahuichivoensis =) SOTLNOGSO.:F.. aut Ler La ne oe Ati ROD UR Nd: OR 6 6. Sori destroying the whole inflorescence 2.):.: Rate, fr... ae eeieeerereso-t tenes 7 oor restricted tO the Spikelets-au wea dein te ony Cate ec te tie conan crt eae 9 7. Spores 5.5—7.5 pam 1Ong..........ecc ce ceeeesccceeesseceeessssseecessstseesssseeee Oe Sphacelatum ah SPOTS: LAT RET: Uy etee aes OM aE 2S GY | ORT E| eR) | 8 8. Spores 8—12(—13) um long, slightly dimorphic ....................:eceee S. cenchri Spores 10—14(—16) ym long, not dimorphic.................... S. pennisetinum 9(6).Columella and spore balls absent. Spores 6.5—8 um long.... U. kKamerunensis Columella and spore balls present. Spores larger ...............ccccceeseeceerteeeeeees 10 10.Plants stunted, forming witches’ brooms. Spores 7—11 pm long ....S. dubiosum =/7Plants notsov spores laroery Minnie... cert. tees Een ere ts ee are 1] 11.Columella one, stout. Spores (9—)10—13.5(—14.5) um long............. S. penniseti “4. Columeliac several ii. isc censor ei cole Se cere te eR ee 12 IZ Spores 9-12 pi long Ne se eee ek cerca tae te eee Ba S. divisum = SPOres 12-16" LM TONE Nn are een ener tee) S. penniseti-japonici

THE SMUT FUNGI OF CYMBOPOGON (GRAMINEAE)

Cymbopogon Spreng., in the subfam. Panicoideae, tribe Andropogoneae, subtribe Andropogoninae, is a fairly homogeneous genus of c. 40 species in the Old World tropics and subtropics; some species are introduced to tropical America (Clayton & Renvoize, 1986:351). At least 29 smut fungi have been reported on it under different generic names (Sorosporium, Sphacelotheca, Sporisorium, Tolyposporium, Ustilago), which all belong to the genus Sporisorium. Several are considered to be synonyms. The 13 recognised species, including two new ones, are: 1. Sporisorium barberi (Mundkur) Vanky, comb. nov.

Basionym: Ustilago barberi Mundkur, Trans. Brit. Mycol. Soc. 23:98, 1939. Sorosporium barberi (Mundkur) L. Ling, 1951a:44. Farysia_barberi (Mundkur) Zundel, 1953:40. Type on Cymbopogon coioratus (Nees) Stapf, India, Madras, Tinnevelly Distr., Palamcotta, 11.V.1901, C.A. Barber 2677, HCIO 7398. Isotypes in BPI 170200, IMI, HUV 17331!

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Sorosporium cornutum M.S. Pavgi & Thirum., in Thirumalachar & Pavgi, 1956:282. Type on Cymbopogon confertiflorus (Steud.) Watson, India, Wynaad, Panora, 17.XI.1909, W. McRae, HCIO. Isotype IMI 44440! (syn. here).

Sori destroying the innermost floral organs transforming them into tough, horn-shaped bodies, 1-2 x 5-25 mm, first covered by a thick, brown, smooth peridium which later splits longitudinally disclosing the semiagglutinated to powdery, dark brown mass of easily disintegrating spore balls and spores surrounding several (3—7) tapering columellae. The fungus usually produces proliferation of the infected spikelets and the sori are agglomerated like witches’ brooms. Spores globose, subglobose, ovoid or subpolyhedrally slightly irregular, more or less variable in size, 7-12 x 8-13 pm, yellowish- to reddish-brown; wall evenly or unevenly thick, 0.5—-1.5 um, provided with sparsely situated, rounded, coarse warts, in SEM appearing as tubercles.

On Cymbopogon coloratus (Nees) Stapf, C. flexuosus (Nees ex Steud.) Watson, C. nardus (L.) Rendle var. confertiflorus (Steud.) Stapf ex Bor (C. confertiflorus (Steud.) Stapf); S. Asia (India).

There are rather big variations in the size of the spores between different collections, and in some collections even within the same sorus. E.g., the spore diameter of the globoid spores in the type of Sporisorium barberi varies between 6.5 and 12 um. The spores of the HUV 8903 specimen (on Cymbopogon coloratus, India, Kerala, at Lake Periyar, 5.I].1980, K. Vanky) are more uniform, the globoid spores measuring 10-13 pum. In the type of Sorosporium cornutum, the spores are also uniform but smaller, measuring (7—)8—9.5(—10.5) x 8-11 pm. However, the typical sori and the even more typical spores, provided with sparsely situated tubercles, present in all these collections, make the identification easy. Despite the differences in spore measurements, I consider Sorosporium cornutum to be a synonym of Sporisorium barberi.

2. Sporisorium bengalense (H. & P. Sydow & Butler) Vanky, comb. nov. Basionym: Ustilago bengalensis H. & P. Sydow & Butler, Ann. Mycol. 10:250, 1912. Sphacelotheca bengalensis (H. & P. Sydow & Butler) Mundkur, 1939:108. Type on Cymbopogon pendulus (Nees ex Steud.) Watson, India, Bengal, Duars, Banarhat, 8.[X.1908, I.H. Burkill, HCIO 1432. Isotypes in BPI 157972, 177259, 195064, HUV 17271!

Sori in the ovaries, probably also comprising other, inner floral organs, cylindrical, 0.5—1 x 3-10 mm, first covered by a brown peridium which ruptures irregularly disclosing the brown, powdery mass of spores and sterile cells surrounding a central, tapering columella. Spores when mature single, globose, subglobose to ellipsoidal, 10.5-13.5 x 11-15 wm, yellowish-brown; wall even, c. 0.5 um thick, finely, densely punctate, spore profile smooth to finely serrulate, in SEM finely, densely echinulate. Sterile cells small, single, globose to subglobose, 5—8 ym in diameter, hyaline; wall 0.5—1 um thick, smooth.

On Cymbopogon pendulus (Nees ex Steud.) Watson; S. Asia (India).

22

Fig. 13. Sori of Sporisorium compactum Vanky, as long, narrow, horn-shaped bodies in the flowers of Cymbopogon giganteus Chiov. (type). Habit and enlarged two sori. Bars = 1 cm for habit, and 2 mm for the detail drawing.

Fig. 14. Sori of Sporisorium cymbicum Vanky in the ovaries of Cymbopogon nardus (L.) Renle (type). To the left a healthy inflorescence. Bar = 1 cm.

a

3. Sporisorium compactum Vanky, sp. nov. Typus in matrice Cymbopogon giganteus Chiov., Senegal, Bakel, 14°54' N, 12°26' W, 3.1.1964, leg. G. Adam 19647, HUV 19818!

Sori ipsis speciei Sporisorium barberi similiter organa floralia intima in corporibus longis, corniformibus, agglomeratis, 0,5—1 x 5—15 mm transformantes, involucris floralibus proliferantibus partim occulti et peridio cinereobrunneo cooperti, quo longitudinaliter rupto massam nigram, semi-agglutinatam usque granuloso pulveream glomerulorum sporarum columellamque 1(—2) longam, filiformem ostendentes. Glomeruli sporarum globosi, ovoidei, ellipsoidales usque subpolyedrice irregulares, 40-80 x 50-120 um, atrobrunnei usque opaci, e sporis multis, firmiter agglutinatis compositi. Sporae globosae, ellipsoidales usque parum irregulares, 9,5-13 x 12-15 «wm; sporae externae atro-olivaceobrunneae; pariete 1—1,5 um crasso, in superficie libera dense echinulato, imago obliqua sporae serrulata; sporae internae colore pallidiores; pariete tenui (cca. 0,5 um), conspicue levi.

Sori (Fig. 13), similar to those of Sporisorium barberi, transforming the innermost floral organs into long, horn-shaped, agglomerated bodies, 0.5—-1 x 5—15 mm, partly hidden by proliferating floral envelopes and covered by a greyish-brown peridium which ruptures longitudinally disclosing the black, semiagglutinated to granular-powdery mass of spore balls and 1(—2) long, filiform columellae. Spore balls (Figs. 17, 18) globose, ovoid, ellipsoidal to subpolyhedrally irregular, 40-80 x 50-120 um, dark brown to opaque, composed of many, firmly agglutinated spores. Spores (Figs. 17, 18) globose, ellipsoidal to slightly irregular, 9.5—13 x 12-15 um; outer spores dark olivaceous brown; wall 1-1.5 wm thick, densely echinulate on the free surface, spore profile serrulate; inner spores lighter coloured, thin-walled (c. 0.5 um), apparently smooth.

On Cymbopogon giganteus Chiov.; W. Africa (Senegal).

The compact spore balls of firmly united spores, which break rather than separate under pressure, differentiate this species from all other smuts of Cymbopogon.

4. Sporisorium cymbicum Vanky, sp. nov. Typus in matrice Cymbopogon nardus (L.) Rendle (det. S. Kativu), Zimbabwe, Manicaland Prov., Nyanga National Park, Mtarazi Waterfalls, 18°29'07" S, 32°47'36" E, alt. cca. 1700 m.s.m., 22.11.1999, leg. C. & K. Vanky, HUV 19808! Isotypi in BPI 841911, et in Vanky, Ust. exs. no. 1150. Paratypus in matrice Cymbopogon validus (Stapf) Stapf ex Burtt Davy, South Africa, Northern Prov., prope Duiwelskloof, Ebeneser Dam, road no. 528, 23°56'33" S, 29°57'12" E, alt. cca. 1490 m.s.m., 23.1.1997, leg. C. & K. Vanky, HUV 19816. Isoparatypi in BPI 841712, et in Vanky, Ust. exs. no. 1151. Paratypus in matrice Cymbopogon plurinodis (Stapf) Stapf ex Burtt Davy, South Africa, Western Cape Prov., 55 km E Knysna, Tsitsikamma National Park, 33°57'42" S, 23°32'00" E, alt. cca. 250 m.s.m., 19.XII.1996, leg. C. & K. Vanky, HUV 198871. Isoparatypi in Vanky, Ust. exs. no. 1156. Paratypus in matrice Cymbopogon nardus (det. M. Namaganda), Uganda, Iganga Distr., 50 km NNE Iganga, 00°48'35" N,

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fs

33°48'31" E, alt. cca. 1155 m.s.m., 1.]11.2002, leg. M. Namaganda, C. & K. Vanky, HUV 19979! Isoparatypi in MHU, Kampala.

Sori in ovariis omnibus eiusdem inflorescentiae, ellipsoidales, 0,7-1 x 2—4 mm, indumentis floralibus plus-minus obtecti, primo peridio pallide brunneo cooperti, quo ab apice rupto massam sporarum atrobrunneam, pulveream, columellam centralem, flagelliformem, soros excedentem circumdantem demonstrantes. Sporae in maturitate singulares, globosae, subglobosae, ellipsoidales, raro parum irregulares, (9—)9,5—13 x (9,5—)10,5—13,5 um, flavido- brunneae; pariete aequali, cca. 1 um crasso, leniter, dense verrucoso-echinulato; imago obliqua sporae undulata usque leniter serrulata. Cellulae steriles paucae, in catervis irregularibus, cellulae singulares earum subglobosae, ellipsoidales usque irregulares, 11-17 um longae, hyalinae vel subhyalinae; pariete cca. 1 um crasso, levi.

Sori (Fig. 14) in the ovaries infecting all spikelets in an inflorescence, ellipsoidal, 0.7—-1 x 2-4 mm, more or less hidden by the floral envelopes, first covered by a pale brown peridium which ruptures from its apex disclosing the dark brown, powdery mass of spores surrounding a flagelliform central columella exceeding the sor. Spores (Figs. 19, 20) when mature single, globose, subglobose, ellipsoidal, rarely slightly irregular, (9-)9.5-13 x (9.5—)10.5—13.5 um, yellowish-brown; wall even, c. | tm thick, finely, densely verrucose- echinulate; spore profile wavy to finely serrulate. Sterile cells (Figs. 19, 20) few, in irregular groups, single cells subglobose, ellipsoidal to irregular, 11-17 um long, hyaline or subhyaline; wall c. 1 um thick, smooth.

On Cymbopogon nardus (L.) Rendle, C. plurinodis (Stapf) Stapf ex. Burtt Davy, C. validus (Stapf) Stapf ex Burtt Davy (C. afronardus Stapf); Africa (South Africa, Uganda, Zimbabwe). According to Gibbs Russel et al., 1991:95, C. validus probably represents C. nardus.

Variations of + 1 um in the spore measurements of different collections may occur. The spores of the paratype from Uganda are somewhat smaller (9.5-13 um long) and paler.

Sporisorium cymbicum is closest to S. panamense from which it differs especially by the smaller spores.

5. Sporisorium cymbopogonis (Mundkur) Vanky, comb. nov. Basionym: 7olyposporium cymbopogonis Mundkur, Indian J. Agric. Sci. 14:51, 1944. Sorosporium cymbopogonis (Mundkur) Thirumalachar & Neergaard,

Figs. 15, 16. Spore balls and spores of Sporisorium tothii Vanky on Pennisetum glaucum (L.) R. Br., in LM and in SEM (type).

Figs. 17, 18. Spore balls and spores of Sporisorium compactum Vanky on Cymbopogon giganteus Chiov., in LM and in SEM (type).

Figs. 19, 20. Spores of Sporisorium cymbicum Vanky on Cymbopogon nardus (L.) Rendle, in LM and in SEM (type). Bars = 10 um.

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1977(1978):183. Type on Cymbopogon citratus (DC.) Stapf, India, Mysore, Bangalore, 20.X11.1941, M.J. Thirumalachar, HCIO. Isotype in BPI 192617!

Tolyposporium christensenii Raghunath, 1968:120 (as "Ragunath"). Type on Cymbopogon flexuosus (Nees ex Steud.) Watson, India, Kerala, near Aruvikkara, 1.1965, T. Raghunath, AMH 264. Isotypes in HCIO 31248, HUV 5379!, 17274, 17275, IMI 137316.

Sori destroying the racemes, long cylindrical, 0.5—2 x 20-40 mm, more or less hidden by leaf sheaths and spathae, covered by a pale yellowish-brown peridium which dehisces from its apex disclosing the black, granular-powdery mass of spore balls and 1—2(—3), long, filiform columellae. Usually all racemes of an inflorescence are infected, denoting a systemic infection. Infected inflorescence more or less congested. Raghunath (1970) also observed local infection with formation of witches’ brooms. Spore bails persistent, variable in shape and size, subglobose, ellipsoidal, elongated or irregular, 30-85 x 40-100 pm, dark reddish- brown, composed of tens of spores which separate by pressure. Spores dimorphic, outer spores subglobose, ovoid, ellipsoidal to usually subpolyhedrally slightly irregular, from 6.5-9 x 7-11 ym to 8.5—-13 x 9-13 um, reddish-brown; wall uneven, 1-2 ym thick, free surface moderately densely verrucose-echinulate, contact sides punctate or apparently smooth, spore profile of the free surface serrulate; inner spores rounded, subpolyhedrally slightly irregular, about the size of the outer spores, pale yellowish-brown; wall even, c. 0.5 ym thick, apparently smooth. Sterile cells absent.

On Cymbopogon citratus (DC.) Stapf, C. coloratus Stapf, C. flexuosus (Nees ex Steud.) Watson; S. Asia (India), Indonesia (Bali).

I am considering S. cymbopogonis in a broad sense, with rather big variations in the spore measurements. These in the type of S. cymbopogonis are 6.5—-9 x 7-11 pm, in 7. christensenii 8.5-13 x 9-13 um, and in a specimen on C. flexuosus collected in Bali (HUV 19910) 7—10.5 x 8-11 jm. Other characters are identical in all collections I have seen.

6. Sporisorium cymbopogonis-distantis (L. Ling) L. Guo, 1998:1. Sorosporium cymbopogonis-distantis L. Ling, 1953b:341. Type on Cymbopogon distans (Nees ex Steud.) Watson, China, Szechwan Prov., vicinity of Chengtu, XI.1947, K.R. Lin, BPI 195143!

Sori in the ovaries (0.5—1 x 1-2 mm) or in the whole raceme (10—17 mm long), concealed by the spathae, first covered by a brown, thick pernidium which ruptures disclosing the semiagglutinated to granular-powdery mass of spore balls surrounding several filiform columellae. Spore balls globose, ovoid, ellipsoidal to irregular, 40-90 x 60-120 pm, dark reddish-brown to opaque, composed of numerous spores which separate by pressure. Spores subglobose, ellipsoidal to subpolyhedrally irregular, 10.5—13.5 x 12—17(-19) um, yellowish- to reddish- brown; wall even to slightly uneven, c. 1 wm thick, outer spores apparently smooth to finely punctate-verruculose, inner spores smooth to finely punctate.

On Cymbopogon distans (Nees ex Steud.) Watson; E. Asia (China).

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7. Sporisorium densiflorum (L. Ling) Vanky, comb. nov.

Basionym: Sorosporium densiflorum L. Ling, Lloydia 16:188, 1953a. Type on Cymbopogon densiflorus (Steud.) Stapf, Belgian Congo [= Democratic Rep. of Congo], Leverville, 6.XII.1918, H. Vanderyst, BR 1329. Isotype in BPI 179970!

Sori comprising the racemes, narrow-cylindrical, 0.5-1 x 15-20 mm, completely hidden by leaf sheaths and spathae, first covered by a thin peridium which ruptures from its apex disclosing the dark brown, semiagglutinated to powdery mass of spore balls, spores and 2-7 long, filiform columellae. Spore balls subglobose, ovoid, oblong or irregular, 40-100 x 50-140 um, dark reddish- brown to opaque, composed of numerous, easily separating spores. Spores subglobose, ovoid, ellipsoidal, mostly subpolyhedrally irregular, 9-13 x 9.5—-14(-15) um, reddish-brown; wall unevenly thick, 0.5—2 um, finely, rather densely punctate-verruculose, spore profile smooth. Sterile cells not seen.

On Cymbopogon densiflorus (Steud.) Stapf, and C. dieterlenii Stapf ex Phill.; C. & S. Africa (Congo, South Africa).

8. Sporisorium lanigeri (Magnus) Vanky, comb. nov. Basionym: Ustilago lanigeri Magnus, Verh. K. K. Zool.-Bot. Ges. Wien 49:88, 1899, Sphacelotheca lanigeri (Magnus) Maire, in Zundel, 1930:141. Type on Andropogon laniger Desf. (= Cymbopogon schoenanthus (L.) Spreng.), Iran, Kerman, Mt. Kuh-tagh-Ali, alt. 2200 m, 20.VI.1892, J. Bornmiiller 4436, S. Isotypes in BPI 194475! and HBG.

Ustilago furcata Patouillard & Hariot, 1900:236. Sphacelotheca furcata (Pat. & Har.) Pat. & Har., in Hariot & Patouillard, 1909:197. Type on Cymbopogon sp., French Sudan, Reg. Tombuctou, Sompi, VIII.1899, M.A. Chevalier, FH. (syn. by Ling, 1951a:42).

Ustilago schoenanthi H. & P. Sydow & Butler, 1906:425. Sphacelotheca schoenanthi (H. & P. Sydow & Butler) Zundel, 1930:136. Type on Andropogon schoenanthus L. (= Cymbopogon schoenanthus (L.) Spreng.), India, Madras, Tinnevelli, Alancolam, 1.VII.1901, C.A. Barber, HCIO 449. Isotypes in BPI 166233, 188944, 193952, HUV 1994, 17304! (syn. here).

Sphacelotheca moggii Zundel, 1930:130. Type on Andropogon plurinodis Stapf (= Cymbopogon plurinodis (Stapf) Stapf ex Burtt Davy), South Africa, British Bechuanaland, Armoedsvlakte, 28.VIII.1924, A.O.D. Mogg, PREM 19859. Isotypes in BPI 178080, 195089, HUV 18171! (syn. here).

Sphacelotheca concentrica Zundel, 1930:138. Type on Cymbopogon plurinodis (Stapf) Stapf ex Burtt Davy, South Africa, Transvaal, Pretoria, Kaalplaats, 1.XI.1917, A.O.D. Mogg, PREM 10708. Isotypes in BPI 195116, HUV 18020! (syn. by Ling, 1951a:43, confirmed).

Sorosporium pretoriense Zundel (as "pretoriaense"), 1930:146. Type on Andropogon dichrous Steud. (= Cymbopogon plurinodis; comp. Zundel, 1953:70), South Africa, Transvaal, Pretoria, between Ashbury and Irene, 14.11.1917, LB. Pole-Evans, PREM 10045. Isotypes in BPI 180139, HCIO 10143, HUV 15413! (syn. here).

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Sphacelotheca natalensis Zundel, 1930:139. Type on Andropogon sp. (= Cymbopogon excavatus (Hochst.) Stapf ex Burtt Davy; comp. Doidge, 1950:382), South Africa, Natal, Mooi River, 4.[X.1917, A.O.D. Mogg, PREM 11705. Isotypes in BPI 190145, 195074, 195090, HUV 18173! (syn. here).

Sphacelotheca columellifera "(Tul.) Yen", 1937:76 (not (L.-R. & C. Tul.) Ciferri, 1928:32). Sphacelotheca yenii Zundel, 1939a:584. Type on Andropogon laniger Desf. (= Cymbopogon schoenanthus (L.) Spreng.), Morocco, near Skourat, 13.V1I.1934, G.J.L. Malengon, PC. Isotypes in HMAS, HUV 12075! (syn. by Ling, 1951a:43, confirmed). |

Sphacelotheca cymbopogonis W.Y. Yen, 1938:7. Type on Cymbopogon proximus Stapf, Chad, Quadai, between Bittine and Abéché, 10.X.1935, M. Murat 34, LAM. (syn. by Ling, 1951a:43).

Sphacelotheca consueta H. Sydow, in Sydow & Ahmad, 1939:442. Type on Cymbopogon parkeri Stapf, Pakistan, Panjab, Shahkot Hills, 15.XII.1935, S. Ahmad 60, HCIO 10101. Isotypes in BPI 195065, S, HUV 16374, topotype in BPI 1110460. (syn. by Ling, 1951a:43, confirmed).

Sorosporium ladharense H. Sydow, in Sydow & Ahmad, 1939:443. Type on Cymbopogon jwarancusa (Jones) Schult., India, Punjab, Ladhar, Sheikhupura, 10.1V.1937, S. Ahmad 60a, HCIO 10104. Isotypes in BPI 195124, HUV 15618! (syn. here).

Sphacelotheca cymbopogonis-colorati Mundkur & Thirum., in Thirumalachar & Mundkur, 1951:3. Sporisorium cymbopogonis-colorati (Mundkur & Thirum.) Kakishima & Ono, 1993:189. Type on Cymbopogon coloratus (Nees) Stapf, India, Mysore, Hassan, Bellur, 28.X1I.1942, M.J. Thirumalachar, HCIO 10698. Isotypes in HUV 17280!, IMI 4954. (syn. here).

Sphacelotheca mutila Mundkur & Thirum., in Thirumalachar & Mundkur, 1951:4. Type on Cymbopogon caesius (Nees) Stapf, India, Mysore, Bangalore, 18.VIII.1944, M.J. Thirumalachar, HCIO 10774: isotype in HUV 17292! (syn. here).

Sori destroying the racemes, more or less hidden by the enveloping spathae, cylindrical, tapering at the apex, 1-2 x 5-16 mm, first covered by a pale brown, thick peridium which ruptures irregularly disclosing the dark brown, semiagglutinated to powdery mass of spore balls, spores and sterile cells surrounding a stout, tapering, rarely flagelliform, sometimes bifurcate, central columella. Spore balls globose, ellipsoidal to irregular, 30-120 x 40-140 um, dark reddish-brown to opaque, composed of numerous spores which separate by pressure. Spores globose, subglobose, ellipsoidal to slightly irregular, often slightly compressed, 5.5-8 x 6~9 um, yellowish- to reddish-brown; wall evenly thick, 0.5-1 um, apparently smooth to finely, moderately densely verruculose- echinulate; spore profile smooth to finely wavy. Sterile cells in irregular groups, single cells subglobose, ellipsoidal to irregular, 7-20 um long, hyaline to pale yellowish-brown tinted; wall 1—1.5 um thick, smooth.

On Cymbopogon ambiguus A. Camus, C. bombycinus (R. Br.) Domin, C. caesius (Nees) Stapf, C. coloratus (Nees) Stapf, C. commutatus (Steud.) Stapf, C. distans (Nees ex Steud.) Watson, C. elegans Spreng., C. excavatus (Hochst.)

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Stapf ex Burtt Davy, C. flexuosus (Nees ex Steud.) Watson, C. jwarancusa (Jones) Schult., C. marginatus (Steud.) Stapf ex Burtt Davy, C. nardus (L.) Rendle, and its var. confertiflorus (Steud.) Stapf ex Bor (C. confertiflorus (Steud.) Stapf), C. obtectus $.T. Blake, C. parkeri Stapf, C. plurinodis (Stapf) Stapf ex Burtt Davy, C. procerus (R. Br.) Domin (C. exaltatus (R. Br.) Domin), C. proximus Stapf, C. refractus (R. Br.) A. Camus, C. schoenanthus (L.) Spreng. (Andropogon schoenanthus L., Andropogon laniger Desf.), C. validus (Stapf) Stapf ex Burtt Davy; Africa (Chad, Morocco, South Africa, Sudan, Zimbabwe), Asia (India, Iran, Iraq, Pakistan), Australia. |

Study of the types of Ustilago lanigeri Magnus, Ustilago schoenanthi H. & P. Sydow & Butler, Sphacelotheca moggii Zundel, Sorosporium pretoriense Zundel, Sphacelotheca natalensis Zundel, Sorosporium ladharense H. Sydow, Sphacelotheca cymbopogonis-colorati Mundkur & Thirum. and Sphacelotheca mutila Mundkur & Thirum. revealed only insignificant differences in the colour, size and ornamentation of the spores, hence they are considered to be synonyms.

9. Sporisorium mildbraedii (H. & P. Sydow) Vanky, comb. nov. Basionym: Ustilago mildbraedii H. & P. Sydow, in Mildbraed, Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907-1908, Bd. 2, p. 95, 1914. Sphacelotheca mildbraedii (H. & P. Sydow) Zundel, 1930:135. Type on Andropogon schoenanthus L. (= Cymbopogon schoenanthus (L.) Spreng.), Deutsch Ost Africa [= Rwanda], Mpororo, at Kakitumbe Creek, 23.VII.1907, G.W.J. Mildbraed 357, S. Isotypes in BPI 163317!, 194453!

Sphacelotheca cymbopogonis-afronardi L. Ling, nom. herb. On Cymbopogon afronardus Stapf (= C. nardus (L.) Redle), Uganda, Kawanda, VII.1940, leg. C.G. Hansford 2754, BPI 177450, 177451, Herb. Mycol. Dep. Agric. Uganda 2754, HUV 1958, 6066, IMI 10956.

Sori destroying the racemes, concealed by the spathae, 1-2 x 10-15 mm, peridium lacking. Spore mass brown, powdery, surrounding a well-developed central columella with short lateral branches. Spore balls subglobose, ellipsoidal, irregular, 40-80 x 40-120 pm, dark reddish-brown to subopaque, composed of numerous, easily separating spores. Spores globose, subglobose, ovoid to ellipsoidal, 56.5 x 5.5-7 pm, yellowish-brown; wall even, c. 0.5 wm thick, finely, moderately densely punctate to verrucose-echinulate; spore profile smooth, wavy to sparsely serrulate. Sterile cells in irregular groups, single cells globose, ellipsoidal to irregular, 7-12 ym long, subhyaline to pale yellowish- brown, wall c. 0.5 um thick, smooth.

On Cymbopogon schoenanthus (L.) Spreng. (Andropogon schoenanthus L.), C. nardus (L.) Redle (C. afronardus Stapf); E. Africa (Tanzania, Uganda).

Sporisorium mildbraedii differs from S. lanigeri especially in the lack of a peridium and in the smaller, more conspicuously ornamented spores.

10. Sporisorium mutabile (H. Sydow) Vanky, comb. nov. Basionym: Sphacelotheca mutabilis H. Sydow, Ann. Mycol. 35:24, 1937. Sorosporium mutabile (H. Sydow) L. Ling, 1951b:107. Type on Cymbopogon

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refractus (R. Br.) A. Camus, Australia, New South Wales, Pennant Hills, V1.1931, L.R. Fraser 118. Isotypes in BPI 195071, HUV 18115!, IMI 37633.

Sorosporium cantonense Zundel, 1939a:584 (as "Cantonensis"). _ Sporisorium cantonense (Zundel) L. Guo, 1998:1. Type on Cymbopogon hamatulus (Nees) A. Camus (as "hematatus"), China, 80 miles north of Canton, on North River, Yinktak (Guangdong, Guangzhou), 9.1X.1921, A.S. Hitchcock 18822, BPI 179478! (syn. here).

Sorosporium terrareginalense Zundel, 1944:409. Type on Cymbopogon refractus (R. Br.) A. Camus, Australia, Queensland, Brisbane, Highway near Mt. Coot-tha, 9.111943, M.S. Clemens, BPI 180159! (syn. by Ling, 1951b:107).

Sori destroying the racemes, long cylindrical, 0.5—1 x 5-13 mm, partly hidden by leaf sheaths and spathae, first covered by a brown peridium which ruptures from its apex disclosing the dark brown, semiagglutinated to granular powdery mass of spore balls and 1-3 long, filiform columellae. Spore balls globose, ellipsoidal, irregular, 40-80 x 40-120 wm, dark reddish-brown to subopaque, composed of numerous, easily separating spores. Spores subpolyhedrally irregular to elongated, 10-15 x 13—18(—20) um, yellowish-brown; wall unevenly thick, 0.5—1.5(—2) um, evidently, rather densely verrucose, especially on the free surface, inner spores punctate-verruculose; spore profile smooth to wavy or finely serrulate.

On Cymbopogon bombycinus (R. Br.) Domin, C. coloratus (Nees) Stapf, C. densiflorus (Steud.) Stapf, C. distans (Nees ex Steud.) Watson, C. hamatulus (Nees) A. Camus, C. nardus (L.) Rendle, C. refractus (R. Br.) A. Camus; Africa (Malawi, Zimbabwe), Asia (China, India, Pakistan), Australia.

Comparison of the types revealed that Sporisorium cantonense is identical with and a synonym of Sporisorium mutabile.

11. Sporisorium nardi (H. & P. Sydow) Vanky, comb. nov. Basionym: Ustilago nardi H. & P. Sydow, in H. & P. Sydow & Butler, Ann. Mycol. 4:425, 1906. Sphacelotheca nardi (H. & P. Sydow) Zundel, 1930:137. Sorosporium nardi (H. & P. Sydow) L. Ling, 1951a:47. Type on Andropogon nardus L. (= Cymbopogon nardus (L.) Rendle), India, Malabar, Wyanaad Hills, Vayitri, 2.X1.1904, E.J. Butler, HCIO 470. Isotypes in BPI 163727-173730,, HUYV 1977, 17293,:S:

Sori in the racemes, filiform, c. 0.5 x 5-10 mm, completely hidden by the spathae, covered by a brown peridium which ruptures disclosing the dark brown, semiagglutinated to granular-powdery mass of spore balls and 1—2 long, filiform columellae usually protruding beyond the sori. Spore balls globose, ellipsoidal to irregular, 30-70 x 30-120 um, dark reddish-brown to subopaque, composed of many, easily separating spores. Spores subglobose, ellipsoidal to slightly irregular, 5.5—8 x 6-9.5 um, yellowish-brown; wall even, c. 0.5 um, apparently smooth to finely punctate-verruculose, spore profile smooth, in SEM finely, moderately densely verruculose.

On Cymbopogon nardus (L.) Rendle (Andropogon nardus L.); S. Asia (India).

3]

12, Sporisorium panamense (Zundel & Dunlap) M. Piepenbr., in BPI (2001). Sphacelotheca panamensis Zundel & Dunlap, in Zundel, 1939b:995. Type on Cymbopogon sp. (= C. citratus (DC.) Stapf), Panama, Cherique Prov., near El Bogrete, [X.-X.1911, A.S. Hitchcock, BPI 190260!

Sori in the ovaries, c. 0.5-1 x 3-4 mm, hidden by the floral envelopes, first covered by a thin, brown peridium which ruptures disclosing the dark brown, semiagglutinated to powdery mass of spores, sterile cells and a narrowing central columella. Spores when mature single, subglobose to ellipsoidal, 12-15 x 13—17.5 um, yellowish-brown; wall even, c. 1 um thick, finely, densely punctate- verruculose, spore profile smooth, wavy to finely serrulate. Sterile cells in iregular groups, single cells ellipsoidal to irregular, 12-16 um long, hyaline; wall c. 1 um thick, smooth.

On Cymbopogon citratus (DC.) Stapf); C. America (Panama).

13. Sporisorium spermoideum (Berk. & Broome) Vanky, comb. nov. Basionym: Ustilago spermoidea Berkeley & Broome, J. Linn. Soc., Bot. 14:94, 1875. Sphacelotheca spermoidea (Berk. & Broome) Mundkur, 1939:96. Sorosporium spermoideum (Berk. & Broome) L. Ling, 1951a:48. Sorosporium spermoideum (Berk. & Broome) Zundel, 1953:74 (superfl. comb., not Ling, 1951a). Type on Cymbopogon martinii (Roxb.) Watson (as "martii"), Ceylon [= Sri Lanka], Dolosbagey, V.1868, Thwaites 589, BM. Isotype in BPI 166549!

Sori destroying the racemes, long cylindrical, 0.5—1 x 5-15 mm, hidden by leaf sheaths and spathae, first covered by a pale brown peridium which ruptures irregularly disclosing the blackish-brown, semiagglutinated to granular powdery mass of spore balls and 1-5 long, filiform columellae. Spore balls ovoid, oblong, usually irregular, 50-140 x 60-210 um, opaque, composed of many spores which separate by pressure. Spores subglobose, ellipsoidal to subpolyhedrally slightly irregular, (7—)8—10.5 x 9-11(—12.5) um, yellowish-brown, paler on one side; wall unevenly thick, thinner on the pale side, 0.5—1 um, apparently smooth to finely, densely punctate-verruculose, spore profile smooth.

On Cymbopogon confertiflorus (Steud.) Stapf, C. martinii (Roxb.) Watson, C. nardus (L.) Rendle, ?Capillipedium venustum (Thw.) Bor (Andropogon venustus Thw.); S. Asia (Sri Lanka).

Zundel, 1930:142, described Sphacelotheca zilligii Zundel, on Andropogon sp., South Africa, Cape Province, Vryburg, 25.III.1921, A.O.D. Mogg, PREM 20666. Isotypes in BPI 192099, 195088, HUV 18128! Doidge, 1950:384, and also Zundel, 1953:115, considered the host plant to be a Cymbopogon sp. Unfortunately, no healthy host plant is preserved. Judged from the sori, I doubt that the host is a Cymbopogon.

Recently, Bag and Agarwal (2001:221) published a new species, Sporisorium martinii Bag & Agarwal (as "martinae"), type on Cymbopogon martinii (Roxb.) Watson, India, West Bengal, Coochbehar, III.1996, D.K. Agarwal, HCIO 42945. Unfortunately, study of the type specimen was not possible. The incomplete

D2

description of this species, reproduced below, does not permit its comparison with other smut fungi of Cymbopogon, or its inclusion into the following key. "Sori produced in the ovaries, only in parts of spikelets between the spreading - glumes which occasionally exceed in length, enclosed by peridium that soon ruptures and dehisces releasing dark brown to black dusty spore mass. Spores loosely adhere to one another forming loose balls. Sterile cells globose to subglobose, hyaline, similar or mostly larger than spores, 5.1—-11.1 um. Spores light orange yellow (Ridgeway Plate III, d-3), globose to subglobose, spherical, 5.1-7.7 um (Avg. 6.6 pm) in size. SEM showed echinulate sporewall."” ,

Key to the smut fungi of Cymbopogon (S. = Sporisorium)

1. Sori horn-shaped, forming witches' brooms in the inflorescence...................... 2 = DOTETIOUSO feet tet eerie tice ae eee Ae eR ee ne Eee, Ir een eee 3 2. Spore balls easily disintegrating. Spores tuberculate ......................... S. barberi Spore balls compact, permanent. Outer spores echinulate...........8. compactum 3. Spores (12-913-17(-20) (im long wie aes ee et eleratieecteec creates 4 MS POTES SIMAMOTE eric Satirist es COU eee TT ee or Ee Tree ete 6 4. Columella one, simple. Sterile cells present. ...........0.....eeeeees S. panamense Columella 1—several, filiform. Sterile cells absent ..........00000ccecccceecce cece eeeeeeeeeeee 5 5. Spore wall c. 1 um thick. Outer spores apparently smooth

to finely punctate-verruculose...............0:cccceeseeeeeees 5S. cymbopogonis-distanti

Spore wall 0.5—1.5(—2) wm thick. Outer spores verrucose............... S. mutabile OS) Spores between 10S1S im long nee ee ree Net res 4 —inpores smaller ee ra een ee een VOD od, Rep eet Nar fe, Pee Wee 9 Je Stenlercells small 5S en ee ets ee eit tree S. bengalense =" Sterile cellsi' lly lim Oriabsent mite es et eee Leroy Peer yn eae eee ae 8 8. Sori in the ovaries, ellipsoidal, 2-4 mm long. Columelia 1,

fiageilitomi sterilecells presefity. . fe eee S. cymbicum Sori in the racemes, 15-20 mm long. Columellae 2-7,

filiform: Sterile cells absent vee ee S. densiflorum 9(6). Spores 9—11(—12.5) pm long, lighter on one side................. S. spermoideum -- spores smaller not lighter on One SIde®..0c. eee eer me ereeseeereteterrst rarer tes 10 10. Columella one, stout, rarely flagelliform. Sterile cells present...................... 11 Columella 1-several, filiform. Sterile cells absent .....................cccceccceceeceeeeees 12 11. Columella with short branches. Peridium absent. Sterile cells

7-12 pm long. Spores 5.5—7 pam long ..............eecccsseeseceensesecenees S. mildbraedii Columella simple or bifurcate. Peridium present. Sterile cells |

10-20 wm long. Spores 6-9 pm long ....0.... eee ececeeesceneeeeseeesneees S. lanigeri

12(10). Spores 7-13 pm long, free surface verrucose-echinulate..§. cymbopogonis Spores 6—9.5 pm long, free surface finely punctate-verruculose ........... S. nardi

B3

ABOUT SOME OF SPEGAZZINI'S SMUT FUNGI

Marie L. Farr (1973) enumerated Spegazzini's fungal taxa and, based on publications by different authors, also all taxonomic and nomenclatural changes since their publication. Study of many of Spegazzini's original specimens, preserved in LPS (La Plata, Argentina), and in other herbaria, resulted in some additional changes. New name Sporisorium spegazzinii Vanky, nom. nov.

Replacing Sorosporium argentinum Spegazzini, Anales Soc. Ci. Argent. 12:65, 1881 (not Sporisorium argentinum (Hirschh.) Vanky, opus praesens). Ustilago argentina (Speg.) Spegazzini, 1925:151. Type on "“Tricholaena insularis" (= misnamed Panicum urvilleanum Kunth; comp. Spegazzini, 1925:151), Argentina, Prov. Rioja, near Vinchina, leg. G. Hieronymus.

Ustilago negeriana Dietel, 1898:(147). Type on Panicum urvilleanum Kunth, Chile, Prov. Concepcion, near Yumbel, coll. F. Neger, BPI 179441. (syn. by Spegazzini, 1925:151).

Sori in the flowers, infecting all in a panicle, inconspicuous, destroying the inner floral organs, protected by the outermost floral envelopes, 1.5-2 x 2—2.5 mm, first covered by a yellowish-brown peridium which ruptures at maturity from its distal part disclosing the semiagglutinated to powdery, dark brown mass of spore balls and spores surrounding three, short, simple or distally bifurcate columellae. Spore balls subglobose, ellipsoidal to slightly irregular, 35-90 x 45-130 um, dark reddish-brown, composed of tens to hundreds of easily separating spores. Spores subpolyhedrally irregular, more rarely subglobose or ellipsoidal, yellowish-brown, 8—10.5 x 9-12(—13) pm; wall in LM c. 0.5 um thick, in some places thinner (which is difficult to see), surface smooth to very finely and inconspicuously punctate; in SEM the spores have several (6-87), often mamillate depressions, surface very finely, densely verruculose. Sterile cells not seen.

On Gramineae: Panicum urvilleanum Kunth; S. America (Argentina, Chile).

Synonyms Ustilago cordobensis is U. syntherismae

Spegazzini (1881:64) described Ustilago cordobensis Speg. Sphacelotheca cordobensis (Speg.) H.S. Jackson, 1930:298. Sphacelotheca cordobensis (Speg.) Ciferri, 1931:16 (comb. superfl.). Sporisorium cordobense (Speg.) Vanky, 2000:208. Type on “Panicum jaboncillum" Hieron., nom. nud. (= Panicum leucophaeum H.B.K., = Trichachne insularis (L.) Nees, = Digitaria insularis (L.) Fedde; comp. Spegazzini, 1909:288), Argentina, near Cordoba, coll. G.H.E.W. Hieronymus. (syn. here).

Ustilago cacheutensis Spegazzini, 1909:293. Type on Panicum leucophaeum H.B.K. (= Digitaria insularis (L.) Fedde), Argentina, near Mendoza, Cacheuta, I.1909, C. Spegazzini. (syn. by Spegazzini, 1925:153).

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Study of several Argentinean collections of Ustilago cordobensis on Digitaria californica (HUV 19192), D. insularis (LPS 3010), "Panicum penicilligerum" (LPS 3011), "Trichachne rigida" (LPS 12361) could not reveal any essential _ difference between Ustilago cordobensis and the earlier published Caeoma (Uredo) syntherismae Schweinitz, 1834:290 (= Ustilago syntherismae (Schwein.) Peck, 1875:103, type on Syntherisma sp. (= Digitaria sp.), USA), hence they are considered synonyms.

The sori of Ustilago cordobensis and Sporisorium panici-leucophaei are similar, destroying the whole inflorescence. They are surrounded by a thin peridium which flakes away disclosing the dark brown spore masses and several filiform columellae. Probably this is the reason for the confusion between these two smuts. Zundel, 1953:86 & 104, based on spore measurements, demonstrated that S. cordobense and S. panici-leucophaei are two different species.

Ustilago syntherismae is one of the intermediate species between Ustilago and Sporisorium having some characters of both of these genera. Molecular data may be helpful for a more correct generic placement.

Entyloma phalaridis is Jamesdicksonia dactylidis s. \at. Spegazzini described Entyloma phalaridis Spegazzimi, 1912:21 (n. v.), type on Phalaris intermedia Bosc ex Poir., La Plata, X.1919, C. Spegazzini, LPS 3362!, Later, Spegazzini, 1925:155, considered it to be Ustilago striiformis (West.) Niessl. This could not be confirmed by study of the type specimen, which contain spores of /. dactylidis (Pass.) Cif., s. lat. (= Jamesdicksonia dactylidis (Pass.) R. Bauer, Begerow, Nagler & Oberw.), type on Dactylis glomerata L.

Ustilago abortifera is Sporisorium reilianum Ustilago abortifera Spegazzini, 1899:208, type on Zea mays L., Argentina, Salta, VIL1881, LPS 3195! is Sporisorium reilianum (Kihn) Langdon & Full., as shown by Ciferri, 1938:403 (as Sphacelotheca holci-sorghi (Riv.) Cif.), and by Hirschhorn, 1941:350, and others (as Sorosporium or Sphacelotheca reiliana); comp. Farr, 1973:1582. (synonymy confirmed).

Ustilago sorghicola is Sporisorium sorghi Ustilago sorghicola Spegazzini, 1902:58, type on Sorghum vulgare Pers. (Sorghum bicolor (L.) Moench), Argentina, La Plata, IIl.1902, C. Spegazzini, LPS 3047! is Sporisorium sorghi Ehrenberg ex Link., as demonstrated by Ling, 1953b:333 (as Sphacelotheca sorghi (Link) Clinton). (synonymy confirmed).

Excluded species Entyloma nectrioides is a Plasmodiophoromycota Spegazzini, 1891:171, described Entyloma nectrioides Speg. (as "nectrioide"), type "Ad folia Leguminosae? arboreae cujusdam in montanis inter Peribebuy et Mbatobi, Jun. 1883 (sub n. 3783)", Paraguay, Mt. Cordillera de Peribebuy, between Peribebuy and Mbatoby, V1.1883, B. Balansa 3783. Isotypes in Balansa, Pl. du Paraguay 1878-1884, no. 3783, PC, HUV 15939!

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According to the original description, the fungus produces amphigenous, rounded, somewhat swollen, reddish-brown leaf spots, a few mm in diameter. Spores globose, ovoid or ellipsoidal, densely packed, often subangular or sinuous due to mutual pressure, large (50 x 50-80 um in diameter); wall very thick (10 pum), hyaline, multilayered, with densely situated, concentric striae; endoplasm coarsely, densely granular-guttulate, orange-yellow.

Study of this fungus was possible through the kindness of the late Professor Ch. Zambettakis (Paris) who gave me a small fragment of the isotype, preserved in PC as "B. Balansa, Pl. du Paraguay 1878-1884. No. 3783. Entyloma nectrioide Speg. (n. sp.). Paraguay, Cordillére de Péribebuy, au-dessus de Mbatoby, VI.1883".

Spores of 50-80 um diameter, with hyaline, smooth, 4-10 um_ thick, multilayered wall, which collapses when the "spores" are emptied, certainly do not belong to a smut fungus. It turned out also that the content of these "spores" is composed of tightly packed, ovoid or ellipsoidal, small, hyaline cells measuring c. 1 x 1.5 um. These are probably developing zoospores of a Plasmodiophoromycota, and the giant "spores" the sporangia.

Entyloma ameghinoi is a Physoderma sp. Spegazzini, 1902:57, published Entyloma ameghinoi Speg., type on Ranunculus cymbalaria Pursh, Argentina, Patagonia, in the marshes of Chonkenk-aik, along Rio Chico, autumn 1899, C. Ameghino, LPS 3355!

Sori on the leaves and petioles as minute, pale, not or barely swollen spots. Spores globose or ellipsoidal, 15—27 x 16-34 um, pale yellowish-brown; wall c. 1 um thick, smooth.

Study of the type specimen revealed large spores, typical for a Physoderma sp., not for an Entyloma.

Ciferri, 1963:338, considered E. ameghinoi to be a synonym of "Tuburcinia ranunculi-gouani (DC.) Liro", a name which was never published by Liro.

Ustilago dubiosa is insect work

Spegazzini, 1881:64, described Ustilago dubiosa Speg. Type on Conyza graminifolia Spreng. (= Aster divaricatus Nutt.), Argentina "in herbosis campestribus secus el Rio de la Plata, Recoleta”, VI.1881, C. Spegazzini, LPS 3208!. Spegazzini, 1881:65, remarked: "Species valde dubia, corolla bulloso- inflatam, . . . . ; cavum internum a larva dipteri speciei cujusdem semper habitatum!". No spores of a smut fungus could be detected in the scanty type specimen, only insect galls. It is certainly not a smut fungus. For its description see Spegazzini (I.c.) and Zundel, 1953:157, who considered it a doubtful smut.

Ustilago lycoperdospora is a Ustilaginoidea Spegazzini, 1884a:88, described Ustilago lycoperdospora Speg. from immature panicles of Vilfa sp., with the remarks "Species perquam dubiosa". Zundel, 1953, does not mention it. Study of the type (LPS 16382, also under the name of Ustilago peridermiospora Speg.) showed that it is a Ustilaginoidea.

36

Lectotypes proposed Spegazzini, 1925:147, described Entyloma chilense Speg. Type on Conyza chilensis Spreng., La Plata and Buenos Aires, 1919-1923. No type was indicated by Spegazzini. I am designating as a lectotype the specimen of Buenos Aires, 1X.1919)-LPS'3359! For its description see Spegazzini (op. cit.), and Zundel, 1953:242.

Spegazzini, 1909:291, described Tilletia eremophila Speg. Type "Ad spiculas Sporoboli asperifolii vulgatissima circa Mendoza, Aest. 1902-1908", without indicating a type. In LPS there are two collections: LPS 3675 "Tipo", which is extremely scanty, and LPS 3676 "Co-tipo", a rich collection. As lectotype, I am designating the specimen from Argentina, Mendoza, 5.XII.1902, C. Spegazzini, LPS 3676!

Duran & Fischer, 1961:30, demonstrated that 7. eremophila is a synonym of T. asperifolia Ellis & Everhart, type on Sporobolus asperifolius Nees & Mey. (= Muhlenbergia asperifolia (Nees & Mey.) Parodi), USA. (syn. confirmed).

Spegazzini, 1884a:88, published Ustilago paraguayensis Speg. Type "ad paniculas foemineas Graminaceae monoicae elatioris (an bambusaceae?)", Paraguay, Cordillera de Peribebuy, 2.1V.1883, B. Balansa 3776.

In LPS there are two envelopes of this collection: LPS 3704 and LPS 3705. Of these, | am proposing as lectotype the richer one, that of LPS 3704, which is also associated with measurements and drawings of the spores, made by Spegazzini.

THE SMUT FUNGI OF SCHIZACHYRIUM (GRAMINEAE)

Schizachyrium Nees, with c. 60 species throughout the tropics, belongs to the subfam. Panicoideae, tribe Andropogoneae, subtribe Anropogoninae (Clayton & Renvoize, 1986:352). On Schizachyrium two smut fungi are known. Three additional species are described below.

1. Sporisorium absconditum Vanky, sp. nov.

Typus in matrice Schizachyrium fragile (R. Br.) A. Camus, Australia, Queensland, 56 km SW oppid. Mount Garnet, Forty Mile Scrub National Park, 18°03'21" S, 144°51'58" E, alt. cca. 770 m.s.m., 6.1V.1998, leg. C. & K. Vanky. Holotypus in BRIP, isotypi in Herbario Ustil. Vanky, HUV 19960, et in Vanky, Ust. exs. no. 1167. Paratypus in matrice Schizachyrium sanguineum (Retz.) Alston, Venezuela, Estado Aragua, cca. 23 km N urbe La Victoria, 10°26' N, 67°19' W, alt. cca. 2020 m.s.m., 14.XII.1993, leg. R. Berndt, C. & K. Vanky, HUV 19966, isoparatypus in BPI.

Sori racemos destruentes, anguste cylindrici, 0,5—-1 x 15-30 mm, spatheolo tubuliformi paene complete obtecti, apice tantum protrudentes, peridio tenui, pallide brunneo cooperti, quo a parte distali rupto massam nigram, granuloso- pulveream glomerulorum sporarum columellamque longam, filiformem

Bi

ostendentes. Saepe surculi omnes eiusdem plantae infecti, sed non raro planta infecta cum surculo unico in flore, vel surculis nonnullis florentibus proveniens. Glomeruli sporarum persistentes, subglobosi, ovoidei, ellipsoidales usque irregulares, 50-80 x 50-100 pm, atro-rubrobrunnei, opaci, e sporis aliquoties decem, pressu valido tantum separabilibus compositi. Sporae dimorphae, externae earum subgloboae, ellipsoidales usque subpolyedrice irregulares, 8-12 x 10,5—-13,5 ym, atro-flavidobrunneae; pariete inaequali, alternatim tenue vel incrassato, 0,5-2 pm, areas atriores et pallidiores sporarum provocantes; superficies libera sporarum externarum dense verrucosa; imago obliqua sporarum extemarum in superficie libera serrulata, in lateribus contactis levis; sporae internae subpolyedrice irregulares, sporis externis magnitudine circiter aequales, subhyalinae usque pallide flavidobrunneae; pariete tenui, usque cca. 0,5 pm, levi. Cellulae steriles absentes.

Sori (Fig. 21) destroying the racemes, narrow-cylindrical, 0.5-1 x 15-30 mm, almost completely hidden by the tubuliform spatheole, with only their tip protruding, covered by a thin, pale brown peridium which ruptures from its distal part, disclosing the black, granular powdery mass of spore balls and a long, filiform columella. Often all shoots of a plant are infected, but not rarely, an infected plant may have one or several flowering shoots. Spore balls (Figs. 23, 24) persistent, subglobose, ovoid, ellipsoidal or irregular, 50-80 x 50-100 um, dark reddish-brown, opaque, composed of tens of spores which separate only by hard pressure. Spores (Figs. 23, 24) dimorphic, outer spores subglobose, ellipsoidal to subpolyhedrally irregular, 8-12 x 10.5—13.5 ym, dark yellowish- brown; wall uneven, alternatively thin and thick, 0.5-2 um, causing darker and lighter areas on the spores; free surface of the outer spores coarsely verrucose; spore profile serrulate on the free surface, smooth on the contact sides; inner spores subpolyhedrally irregular, about the size of the outer spores, subhyaline to pale yellowish-brown; wall thin, up to c. 0.5 ym, smooth. Sterile cells absent.

On Schizachyrium fragile (R. Br.) A. Camus, S. sanguineum (Retz.) Alston; S. America (Venezuela), Australasia (Australia).

Etymology: from the Latin absconditus, -a, -um = hidden, concealed, referring to the hidden position of the sori.

2. Sporisorium berndtii Vanky, sp. nov. Typus in matrice Schizachyrium sanguineum (Retz.) Alston (det. H. Scholz, B), Sumatra, Samosir Island, Toba Lake, Pangururan Hot Springs, 20. VIII.1997, leg. V. Faust-Berndt & R. Berndt. Holotypus in Herbario Ustil. Vanky, HUV 19957, isotypus in BPI.

Sori in ovariis et organis internis floralibus spiculorum nonnullorum sessilium inflorescentiae eiusdem, cylindrici, arcuati vel torti, cca. 1 x 5-20 mm, inter glumis protrudens, primum peridio pallide cinerescenti cooperti, quo in medio soris longitudinaliter fisso massam atrobrunneam, pulveream glomerulorum sporarum, sporarum et columellarum (1—)2—4(—-5), filiformium ostendentes. Glomeruli sporarum \axi, forma et magnitudine varii, ellipsoidales, elongati vel

38

bY \ Vy FP ! | / \ ; / Nin | is \ | \) /\\ | Gen W MALY 22 Hf hiked | \

Fig. 21. Sori of Sporisorium absconditum Vanky in the racemes of Schizachyrium fragile (R. Br.) A. Camus (type), almost

SSS *,

NV) yy Me, completely hidden by the spatheolae. Me Some shoots escaped infection and are ° flowering. Habit. Bar = 1 cm. oy Fig. 22. Sori of Sporisorium berndtii Vanky in some flowers of Schiza-

chyrium sanguineum (Retz.) A. Camus (type). Habit. Bar = 1 cm.

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irregulares, 35-90 x 50-130 ym, atro-rubrobrunnei, e sporis pluribus decem vel pluribus centum, valde leviter separabilibus compositi. Sporae globosae, ovoideae, ellipsoidales usque subpolyedrice parum irregulares, 8-10,5 x 9-12 um, pallide usque atro-flavidobrunneae; pariete aequali usque inaequaliter crasso, 0,5—1(—1,5) um, ad sporas externas crassiore et simul parum obscuriore, surficie leniter, dense punctato-verrucoso; imago obliqua sporarum levis, usque leniter undulata. Cellulae steriles nullae.

Sori (Fig. 22) in ovaries and inner floral organs of some sessile spikelets of an inflorescence, cylindrical, bent or twisted, c. 1 x 5-20 mm, protruding between the glumes, first covered by a pale greyish peridium which splits longitudinally in the middle of the sorus disclosing the dark brown, powdery mass of spore balls, spores, and (1—)2—4(—5), filiform columellae. Spore balls (Figs. 25, 26) loose, variable in shape and size, ellipsoidal, elongated or irregular, 35-90 x 50—130 um, dark reddish-brown, composed of tens or hundreds of spores which separate very easily. Spores (Figs. 25, 26) globose, ovoid, ellipsoidal to subpolyhedrally slightly irregular, 8—-10.5 x 9-12 um, pale to dark yellowish- brown; wall evenly to unevenly thick, 0.5—1(—1.5) um, thicker on the outer spores which are also somewhat darker, surface finely, densely punctate- verrucose; spore profile smooth to finely wavy. Sterile cells absent.

On Schizachyrium sanguineum (Retz.) Alston, Indonesia (Sumatra).

Etymology: This fungus is named in honour of Dr. Reinhard Berndt (Tibingen, Germany), excellent rust fungus specialist and friend, who also collected many interesting smut fungi, including this species.

3. Sporisorium guaraniticum (Speg.) Vanky, 1989:155. Ustilago guaranitica Spegazzini, 1884a:87. Sphacelotheca guaranitica (Speg.) Zundel, 1930:135. Sorosporium guaraniticum (Speg.) L. Ling, 1953a:190. Type on Andropogon condensatus Kunth (= Schizachyrium condensatum (Kunth) Nees), Paraguay, between Paraguari and Valenzuela, V.1883, B. Balansa 3739, LPS 3050. Isotypes HMAS, HUV 12065!

Sori destroying the racemes, linear, 2-3 cm long, partly hidden by the spathae, first covered by a thin peridium which ruptures irregularly disclosing the blackish-brown, granular-powdery mass of spore balls and a long, slender, filiform columella. Spore balls loose, subglobose, ellipsoidal, elongated or irregular, 40-80 x 50-120 um, light to medium reddish-brown, composed of 20 to hundreds of spores which separate easily. Spores globose, subglobose or slightly irregular, usually ellipsoidal, 12-16 x 13-20 um, yellowish-brown, free surface of the outer spores finely, densely verruculose which just affects the spore profile, inner spores apparently smooth, in SEM very finely verruculose; wall typically unevenly thickened, with (0—)2—4(-©) protuberances or thicker (0.8—2.5 ym) and darker areas, alternating with thin-walled (0.5—1 pm) and light areas. Sterile cells absent.

On Schizachyrium condensatum (Kunth) Nees, S. sanguineum (Retz.) Alston; S. America (Argentina, Ecuador, Paraguay, Venezuela), Australasia (Australia).

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41

Fig. 29. Sori of Sporisorium zambianum Vanky in the racemes of Schizachyrium exile (Hochst.) Pilger, almost completely hidden by the spatheolae (type). To the left a healthy inflorescence. Habit.

Fig. 30. Sori of Sporisorium schizachyrii Vanky in the ovaries of Schizachyrium exile (Hochst.) Pilger (type). Habit. Bar = | cm.

Figs. 23, 24. Spore balls and spores of Sporisorium absconditum Vanky on Schizachyrium fragile (R. Br.) A. Camus, in LM and in SEM (type).

Figs. 25, 26. Spore balls and spores of Sporisorium berndtii Vanky on Schizachyrium sanguineum (Retz.) A. Camus (type).

Figs. 27, 28. Spore balls and spores of Sporisorium zambianum Vanky on Schizachyrium exile (Hochst.) Pilger, in LM and in SEM (type). Bars = 10 pm.

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4. Sporisorium schizachyrii Vanky, 2002a:418.

Type on Schizachyrium exile (Hochst.) Pilger, Zambia, Southern Prov., 75 km _ESE Kafue, Chirundu Fossil Forest, alt. 490 m, 28.1V.2001, C. & K. Vanky, HUV 19654. Isotypes in Vanky, Ust. exs. no. 1128.

For description and illustrations see Vanky, 2002a:418-421. See also Fig. 30.

On Schizachyrium exile (Hochst.) Pilger, and S. sanguineum (Retz.) Alston; C. Africa (Zambia), S. America (Venezuela).

The South American specimens on Schizachyrium sanguineum have 1-5 columellae, whereas in the African specimens on S. exile there is only one columella in a sorus. Other characters are identical.

5. Sporisorium zambianum Vanky, sp. nov. Typus in matrice Schizachyrium exile (Hochst.) Pilger, Zambia, Lusaka Prov., 169 km ENE Lusaka, 15°04'10" S, 29°45'22" E, alt. cca. 900 m.s.m., 17.1V.2001, C., T. & K. Vanky, HUV 19963. Isotypes in BPI, IMI et K.

Sori racemos destruentes, longe cylindrici, 0,5—-1 x 20-40 mm, spatheolo tubuliformi obtecti excepta parte distali eorum, primum peridio tenue, pallide brunneo cooperti, quo rupto et ab apice disjuncto, massam nigrobrunneam, granuloso-pulveream glomerulorum sporarum columellamque longam, filiformem ostendentes. Glomeruli sporarum globosi, subglobosi, ovoidei, usque late ellipsoidales, 30-50 x 40-60 ym, atro-rubrobrunnei usque opaci, e sporis nonnullis decem, pressu leviter separabilibus compositi. Sporae forma variae, subpolyedrice vel polyedrice irregulares, 9-15 x 10,5—17 tm, atro-flavido- usque atro-rubrobrunneae; pariete tmaequaliter crasso, 0,5—-2,5(-3) pm, superficies libera sporarum externarum prominenter echinulata, spinis 1-2 ym altis; sporae internae pallidiores, leniter, disperse punctatae-verrucosae; pariete tenui, cca. 0,5 um, saepe ad angulos crassiore. Cellulae steriles nullae. |

Sori (Fig. 29) destroying the racemes, long-cylindrical, 0.5—-1 x 20-40 mm, hidden by the tubuliform spatheole except its distal part, first covered by a thin, pale brown peridium which ruptures and dehisces from its apex, disclosing the blackish-brown, granular powdery mass of spore balls and a long, filiform columella. Spore balls (Figs. 27, 28) globose, subglobose, ovoid, to broadly ellipsoidal, 30-50 x 40-60 um, dark reddish-brown to opaque, composed of a few tens of spores which separate easily by pressure. Spores (Figs. 27, 28) variable in shape, subpolyhedrally or polyhedrally irregular, 9-15 x 10.5—17 pm, dark yellowish- to dark reddish-brown; wall unevenly thick, 0.5—2.5(—3) um, free surface of the outer spores prominently echinulate, spines 1-2 um high; inner spores lighter, finely, sparsely punctate-verrucose; wall thin, c. 0.5 um, often thicker at the angles. Sterile cells absent.

On Schizachyrium exile (Hochst.) Pilger; C. Africa (Zambia).

43

Key to the smut fungi of Schizachyrium (S. = Sporisorium)

TR OOLl Gest OVitip, Mic TACCINES 11.2.0: 4 te eum seen, haat ts Vere aL 2 =) SOM UE SPIKCLOIS ye nee: esai seek ete es Mee a Ties UUs ately eet gS 4 2. Spores 13-20 um long, free surface finely verruculose............ S. guaraniticum Spores smaller, free surface coarsely ornamented ...........0..0.cccceceeeeeeeeeeeeeeees 8 3. Spores 10.5—17 um long, free surface coarsely echinulate .......... S. zambianum

Spores 10.5—13.5 um long, free surface coarsely verrucose ......8. absconditum 4. Spores dimorphic, i.e. outer and inner spores in a ball very

different, 8-10.5 um long, free surface echinulate ..................... S$. schizachyrii Spores not dimorphic, 9-12 um long, punctate-verrucose................. S. berndtii

A NEW SMUT FUNGUS OF ELIONURUS (GRAMINEAE)

Two smut fungi are known on Elionurus: Sporisorium elionuri (Henn. & Pole- Evans) Vanky, on Elionurus muticus (Spreng.) Kunth, in Africa and South America, and S. elionuri-tristis Vanky, on Elionurus tristis Hack., in Madagascar (comp. Vanky, 1999:155-158). An additional smut fungus, collected in Zambia is: Macalpinomyces elionuri-tripsacoidis Vanky, sp. nov.

Typus in matrice Elionurus tripsacoides Willd., Zambia, Southern Prov., 70 km SW Lusaka, 15°55'29" S, 28°10'00" E, alt. cca. 1060 m.s.m., 16.[V.2001, leg. T., C. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19956. Isotypi in BPI, IMI et K. Paratypi in matrice Elionurus tripsacoides Willd., Zambia, Southern Prov., 22 km NE Pemba, 16°23'03" S, 27°2924" E, alt. cca. 1060 m.s.m., 12.IV.2001, leg. C., T. & K. Vanky, HUV 19959, isoparatypus in BPI; Southern Prov., 87 km NNE Livingstone, 17°16'19" S, 26°15'01" E, alt. cca. 1210 m.s.m., 14.IV.2001, leg. C., T. & K. Vanky, HUV 19958.

Sori in ovariis spiculorum nonnulorum sessilium, hermaphroditorum, longe cylindrici, deinde torti, 0,7-1 x 5-30 mm, peridio brunneo cooperti, qua longitudinaliter rupta et lacerata massam olivaceobrunneam, semiagglutinatam usque pulveream sporarum et cellularum sterilium ostendentes. Sporae globosae, subglobosae usque ellipsoidales, magnitudine variae, 5,5—9,5(-10,5) x 6,5—10,5(-12) wm, flavidobrunneae; pariete aequali, 1-1,5 ym crasso, spinis obtusis, brevibus inclusis; imago obliqua sporarum serrata. Cellulae steriles in catervis magnis, laxis, cellulae singulae globosae usque ovoideae, 4-95 x 6,5—10,5 um, hyalinae; pariete tenui, 0,3—0,4 um, levi.

Sori (Fig. 31) in the ovaries of some of the sessile, hermaphrodite spikelets, long cylindrical, later twisted, 0.7-1 x 5-30 mm, covered by a brown peridium which ruptures and shreds longitudinally disclosing the olivaceous-brown, semiagglutinate to powdery mass of spores and sterile cells. Spores (Figs. 32, 33) globose, subglobose to ellipsoidal, variable in size, 5.5—-9.5(-10.5) x

44

31 TVayT

Fig. 31. Sori of Macalpinomyces elionuri-tripsacoidis Vanky in some ovaries of Elionurus tripsacoides Willd. (type). Habit. Enlarged two healthy spikelets and a sorus. Bars = 1 cm for habit, 3 mm for the detail drawing.

Figs. 32, 33. Spores and sterile cells of Macalpinomyces elionuri-tripsacoidis Vanky on Elionurus tripsacoides Willd., in LM and in SEM (type).

Figs. 34, 35. Spore balls and spores of Sporisorium tristachyae-nodiglumis Vanky on 7ristachya nodiglumis K. Schum., in LM and in SEM (type).

Figs. 36, 37. Spore balls and spores of Restiosporium dapsilanthi Vanky on Dapsilanthus elatior (R. Braun) B.G. Briggs & L.A. Johnson, in LM and in SEM (type). Bars = 10 pm.

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6.5—10.5(—12) pm, yellowish-brown; wall even, 1—1.5 ym thick, including the short, conical spines; spore profile serrate. Sterile cells (Figs. 32, 33) in large, loose groups, single cells globose to ovoid, 4-9.5 x 6.5-10.5 pm, hyaline; wall thin, 0.3—-0.4 um, smooth. |

On Elionurus tripsacoides Willd.; C. Africa (Zambia).

Key to the smut fungi of Elionurus . Spores with conical spines..................... Macalpinomyces elionuri-tripsacoidis Spores finely verruculose or finely echinulate.....................ccccsccceeseeeenreceeeneeeeeed . Spores 6.5—9.5(—10.5) pm long. Sterile cells absent......... Sporisorium elionuri Spores 10—13 pm long. Sterile cells present........... Sporisorium elionuri-tristis

a

NO

A NEW SMUT FUNGUS OF TRISTACHYA (GRAMINEAE)

I revised the smut fungi of the closely related Loudetia, Trichopteryx, Tristachya and Zonotriche (Vanky, 1997:160-171) and recognised ten species, of which six are Sporisorium. A different species was collected in Zambia:

Sporisorium tristachyae-nodiglumis Vanky, sp. nov. Typus in matrice 7ristachya nodiglumis K. Schum., Zambia, Lusaka Province, 169 km ENE urbe Lusaka, 15°04'10" S, 29°45'22” E, alt. cca. 900 m.s.m., 17.1V.2001, leg. C., T. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19719, isotypi in Vanky, Ust. exs. no. 1144.

Sori in flosculis, organa floralia intima destruentes, cylindrici, cca. 1 x 15-25 mm, involucris floralibus partim cooperti, primo peridio crasso, pallide flavidobrunneo obtecti, quo wregulariter rupto massam nigram, granuloso- pulveream glomerulorum sporarum, columellas 3-4, filiformes circumdantem ostendentes. Infectio systemica: plerumque spiculae omnes inflorescentiae eiusdem affectae. Glomeruli sporarum permanentes, forma et magnitudine varii, subglobosi, ellipsoidales, elongati, plerumque parum irregulares, 30-90 x 40—110(—140) ym, rufobrunnei usque subopaci, e sporis 15 usque pluries decem vel etiam centum(?) compositi. Sporae dimorphae. Sporae externae ellipsoidales, ovoideae, plerumque subpolyedrice irregulares, 9,5-13 x 10—15(—16) pm, atro- rufobrunneae; pariete inaequali, in latere interno cca. 0,5 um, in latere libero, externo usque 2,5 um crasso, verrucis vel spinis dense dispositis, cca. 1 pm altis inclusis; imago obliqua sporae dense serrulata. Sporae internae magis rotundae, subglobosae, ellipsoidales vel parum subpolyedricae, 9,5-12 x 10-13,5 um, subhyalinae; pariete aequaliter crasso (cca. 0,5 pm), levi. Cellulae steriles nullae.

Sori (Fig. 38) in the florets, destroying the innermost floral organs, cylindrical, c. 1 x 15-25 mm, partly hidden by the floral envelopes, first covered by a thick, pale yellowish-brown peridium which ruptures irregularly disclosing the black, granular-powdery mass of spore bails surrounding 3-4 filiform columellae.

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38

Fig. 38. Sori of Sporisorium tristachyae-nodiglumis Vanky in the florets of Tristachya nodiglumis K. Schum. (type). Habit, a healthy inflorescence (to the left) and enlarged a triplet of spikelets, each with two sori in the two florets.

Bars = 1 cm for habit, 4 mm for the detail drawing.

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Infection systemic; usually all spikelets of an inflorescence are affected. Spore balls (Figs. 34, 35) permanent, variable in shape and size, subglobose, ellipsoidal, elongated, usually slightly irregular, 30-90 x 40—110(—140) um, reddish-brown to subopaque, composed of 15 to tens or a hundred(?) spores. Spores (Figs. 34, 35) dimorphic. Outer spores ellipsoidal, ovoid, usually subpolyhedrally irregular, 9.5—-13 x 10—-15(—16) pm, dark reddish-brown; wall uneven, on the inner side c. 0.5 um, on the outer, free side up to 2.5 um thick, including the c. 1 um high, densely situated warts or spines; spore profile serrulate. Inner spores more rounded, subglobose, ellipsoidal or slightly subpolyhedral, 9.5—12 x 10—13.5 um, subhyaline; wall evenly thick (c. 0.5 um), smooth. Sterile cells absent. On Tristachya nodiglumis K. Schum., C. Africa (Zambia).

Key to the Sporisorium species of Loudetia, T Sie Hi Sang Tristachya and Zonotriche

1. Sori comprise the whole inflorescence........................ S. loudetiae-pedicellatae Sori restricted to the spikelets (ovaries or florets) .............0.cccccccsseeceeeteeeeeneeees 2 2. Columellae more than 8. Spore balls evanescent. Spores all alike,

GO Lun Tong) yee ei ee tty eas a ieee S. catinatum Columellae less than 5. Spore balls more or less permanent.

Sporesamore:or less dimorphicwiarcer., cetera. eee net eta seear aes 3 SA Sporessl 1222. tim lonpyenct sy ee ieee tee cit See S. loudetiae-superbae SS Sporesmsmialler meh is, Wie aeee Ehe e NEN MSM yo caa a at vss eacugseses 4 4, Spores 12-17 um long. Contact wall of the outer spores thicker

(Cc stim) than thetreewwall (c? lium) y..2.etee eee es S. tristachydis Spores smaller. Contact wall of the outer spores thinner than or

equalling the thicknessofthe free waller is 00 | nye an aed d eestor ae o 5. Columellae 1-3. Spore balls 25-65 wm long............ eee eeeteeees S. decorsei 1 Columellaei3=5/iSpore ‘balls laroercay) Wun sens eee en Aas Pe eet ah 6 6. Spore balls 35-90 um long. Free surface of the spores finely

verruculose. Spore profile nearly smooth...............0...... S. tristachyae-hispidae Spore balls 40—110(—140) um long. Free surface of the spores

verrucose-echinulate. Spore profile serrulate ............ S. tristachyae-nodiglumis

Fig. 39. Sori of Restiosporium dapsilanthi Vanky in the seeds of Dapsilanthus elatior (R. Braun) B.G. Briggs & L.A. Johnson (type). To the left a healthy female inflorescence, to the right a healthy male inflorescence, in the middle an infected male or female inflorescence (habit). Bar = 1 cm.

39

49

50

A NEW SMUT FUNGUS ON DAPSILANTHUS (RESTIONACEAE)

Restiosporium dapsilanthi Vanky, sp. nov.

‘Typus in matrice Dapsilanthus elatior (R. Braun) B.G. Briggs & L.A. Johnson (Leptocarpus elatior R. Brown; det. B. Waterhouse), Australia, Queensland, Cape York Peninsula, Bamaga, near the Air Port, 11°51'07" S, 146°29'18" E, alt. cca. 10 m.s.m., 8.III.2000, leg. R.G. Shivas, C. & K. Vanky, BRIP. Isotypi in BPI 841710, IMI, in Herbario Ustil. Vanky, HUV 19260, et in Vanky, Ust. exs. no. 1149. 3

Sori in nucibus, ovoidei, apice acuti, cca. 0,5 x 1 mm, semina massa glomerulorum sporarum granulosa-pulverea, nigra substituentes. Fungus proliferationem inflorescentiae et simul transformationem eius producens; per soros plantae masculinae infectae in femineis transformatae. Glomeruli sporarum subglobosi, ellipsoidales usque wregulares, 20-50 x 25-60 wm, atro olivaceobrunnei, e sporis 4—40 (vel pluribus?), faciliter separabilibus compositi. Sporae subglobosae, ovoideae, ellipsoidales usque subpolyedrice irregulares, 9-13 x 11—-16(-17) um, olivaceobrunneae; pariete aequaliter crasso, 1-2 pm, conspicue levi usque leniter, dense punctato, in SEM leniter, dense verruculoso.

Sori (Fig. 39) in the nuts, ovoid with an acute tip, c. 0.5 x 1 mm, replacing the seeds with a black, granular-powdery mass of spore balls. The fungus produces proliferation of the inflorescence and also transvestism; infected male plants are transformed into female ones with sori. Spore balls (Figs. 36, 37) subglobose, ellipsoidal to irregular, 20-50 x 25—60 pm, dark olivaceous-brown, composed of 4—40 (or more?), easily separating spores. Spores (Figs. 36, 37) subglobose, ovoid, ellipsoidal to subpolyhedrally irregular, 9-13 x 11-16(-17) pm, olivaceous-brown; wall evenly thick, 1-2 pm, apparently smooth to finely, densely punctate, in SEM finely, densely verruculose.

On Restionaceae: Dapsilanthus elatior (R. Braun) B.G. Briggs & L.A. Johnson (Leptocarpus elatior R. Brown); Australia.

A NEW SPECIES OF MACALPINOMYCES

Of the 25 known Macalpinomyces species six form sori on the top of sterile shoots of Gramineae (Capillipedium, Chrysopogon, Loudetia, Trichopteryx, Tristachya, Vetiveria), Recently, a different Macalpinomyces species was collected in Uganda, on both Sorghastrum and Loudetia:

Macalpinomyces ugandensis Vanky, sp. nov.

Typus in matrice Sorghastrum stipoides (Kunth) Nash (det. M. Namaganda, MHU), Uganda, Rakai Distr., 40 km SW urbe Masaka, 6 km NE pag. Kyotera, 00°35'37" S,.31°35'13” E,-alt’"cca, 1190 m.s.m:, 27.11-2002; leg. T., C. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19992, isotypi in MHU, BPI et in Vanky, Ust. exs. no. 1177. Paratypus in matrice Loudetia phragmitoides (Peter) C.E. Hubb. (det. M. Namaganda); the same place, date and collectors as the type, HUV 19993. Isoparatypes in MHU, BPI et K.

Fig. 40. A sorus. of Macalpinomyces ugan- densis Vanky on the top of a sterile shoot of Sorghastrum stipoides (Kunth) Nash, as a tube filled with spores, which splits at maturity (type). To the left a healthy inflorescence. Bar = 1 cm.

40

rl

3

Sori in apice surculorum, tubos flagelliformes, usque 1 m longos et 1-3 mm latos telarum hostilium massis nigrobrunneis, agglutinatis sporarum et cellularum relative paucarum intermixtarum completos formantes. Maturitate tubi e medio vel a parte distali eorum longitudinaliter fissi, massae sporarum dispersae et tubi aperti ligamenta typica, torta, persistentes formantes. Infectio systemica. Plerumque surculi omnes plantae eiusdem infecti et steriles. Sporae maturae singulae, globosae, subglobosae usque late ellipsoidales, 10,5-13,5 x 12-14,5 um, saepe lateraliter parum deplanatae (8,5-10,5 jum lata), flavido- usque olivaceobrunneae; pariete aequali, cca. 1 um crasso, dense, prominenter echinulato; imago obliqua sporarum serrulata. Cellulae steriles globosae usque ellipsoidales, in catervis laxis vel singulae, magnitudine extreme variae, 6-26 pm longae, hyalinae; pariete aequali, 0,5—2,5 um crasso, levi.

Sori (Fig. 40) on the top of shoots forming flagelliform tubes of host tissue, up to 1 m long, 1-3 mm wide, filled with blackish-brown, agglutinated masses of spores intermixed with relatively few sterile cells. At maturity, the tubes split longitudinally, starting from their middle or distal part, the spore masses are scattered and the opened tubes form typical, persistent, twisted bands. Infection systemic. Usually all shoots of a plant are affected and remain sterile. Spores (Figs. 43, 44) single when mature, globose, subglobose to broadly ellipsoidal, 10.5-13.5 x 12-14.5 um, often laterally slightly flattened (8.5—10.5 um wide), yellowish- to olivaceous-brown; wall even, c. 1 wm thick, densely, prominently echinulate; spore profile serrulate. Sterile cells (Figs. 43, 44) globose to ellipsoidal, in small, loose groups or single, extremely variable in size, 6-26 ym long, hyaline; wall even, 0.5—2.5 um thick, smooth.

On Sorghastrum stipoides (Kunth) Nash and Loudetia phragmitoides (Peter) C.E. Hubb.; C. Africa (Uganda). Known from the type collections only.

Key to the Macalpinomyces species with sori on the top of sterile shoots

1. Spores 4—6.5 pum long, Verrucose ............cccccccsesscccesecsseecsssneeceesenes M. effusus POLES LAT PET, Bees, ce Rt sens s CMe Ah ae renal Pune art, Nets ater Oe ee 2 2. Spores 7—9 um long, wall unevenly thick, 0.5—1.5 um................ M. tristachyae Spores larger, wall evenly or unevenly thick ........0........ccccccceescccesseceesteeeesseeeens : Sspores’8—12. (rm Lona Re a aaa Pe ER a Ce ae tie Yea, eee, 4 = Spores slarpertcGe.:, Page ae, yee na EE Meat mnt, 6 4. Sterile cells 12-35 ym long, wall thick (2.5—3 pum)................ M. trichopterygis ~— Sterile cells 5-15 pm long, wall thin (c. 0.5 WM)... eee eescceteeereeseenteeeees 5 5. Spores coarsely, moderately densely, low echinulate ....... M. chrysopogonicola ~— Spores finely, densely verruculose......1.........cc:cccssccssscesenesseaceserees M. nigritanae 6(3). Spores 9-15 pm long, often acuminate, finely verrucose;

spore: profile wavy sen caked ee ee eh eae 2 We nae M. simplex

Spores 12-14.5 um long, never acuminate, densely echinulate; spore proileserrulate i. ot ee he Gs coc reo ae M. ugandensis

53

41 42

Fig. 41. Sori of Sporisorium rhytachnes-rottboellioidis Vanky in the ovaries of Rhytachne_ rottboellioides Desv. (type).. An infected and a_ healthy inflorescence (habit). Enlarged, two healthy and two infected spikelets (below). Bars = 1 cm for habit, 2 mm for the detail drawings.

Fig. 42. Sori of Sporisorium eriochrysis Vanky in both sessile and pedicelled spikelets of Eriochrysis brachypogon (Stapf) Stapf (type). An infected and a healthy inflorescence (habit). Enlarged, some healthy and some infected spikelets. Bars = 1 cm for habit, 3 mm for the detail drawings.

54

A NEW SPORISORIUM SPECIES ON RHYTACHNE (GRAMINEAE)

I revised (Vanky, 2000:168-172) the smut fungi of Rhytachne and similar grass

genera. On Rhytachne two smut fungi have been recognised: Sporisorium rhytachnes (H. Sydow) Vanky, destroying the whole inflorescence of R. triaristata Stapf (Sierra Leone), and Tolyposporella rhytachnes Viennot-Bourgin, on the leaves of R. minor Pilger (Guinea). A different smut fungus, collected in Uganda, 1s:

Sporisorium rhytachnes-rottboellioidis Vanky, sp. nov. Typus in matrice Rhytachne rottboellioides Desv. (det. M. Namaganda, MHU), Uganda, Masaka Distr., 13 km E Masaka, on Bukakata road, 00°18'19" S, 31°55'52" E, alt. cca. 1140 m.s.m., 17.11.2002, leg. C., T. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19994, isotypi in MHU, BPI, IMI et K.

Sori in ovariis, ovoidei vel cylindrici, 0,5—-1 x 3-5 mm, involucris floralibus partim obtecti, primum peridio cinerescentibrunneo cooperti, quo irregulariter rupto massam atrobrunneam, pulveream glomerulorum sporarum laxorum sporarumque et columellam centralem flagelliformem ostendentes. Glomeruli sporarum globoidei, ellipsoidales, elongati vel irregulares, 20-45 x 25-60 um, rubrobrunnei, e sporis 15 usque pluries decem, leviter separabilibus compositi. Sporae forma et magnitudine variae, rotundatae usque elongatae, plerumque subpolyedrice irregulares, 6,5—9 x 7.5—13,5 um, flavidobrunneae; pariete parum inaequali, 0,5—1 zm crasso, a conspicue levi usque leniter, dense punctato; imago obliqua sporarum levis. Cellulae steriles non observatae.

Sori (Fig. 41) in ovaries, ovoid or cylindrical, 0.5—1 x 3—5 mm, partly hidden by the floral envelopes, first covered by a greyish-brown peridium which ruptures irregularly disclosing the dark brown, powdery mass of loose spore balls and spores, and a flagelliform, central columella. Spore balls (Figs. 45, 46) globoid, ellipsoidal, elongated or irregular, 20-45 x 25-60 ym, reddish-brown, composed of 15 to tens of easily separating spores. Spores (Figs. 45, 46) variable in shape and size, rounded to elongated, usually subpolyhedrally irregular, 6.5-9 x 7.5-13.5 tum, yellowish-brown; wall slightly uneven, 0.5—-1 pm thick, from apparently smooth to finely, densely punctate; spore profile smooth. Sterile cells not seen.

On Rhytachne rottboellioides Desv.; C. Africa (Uganda).

Figs. 43, 44. Spores and sterile cells of Macalpinomyces ugandensis Vanky on Sorghastrum stipoides (Kunth) Nash, in LM and in SEM (type). Figs. 45, 46. Spore balls and spores of Sporisorium rhytachnes-rottboellioidis Vanky on Rhytachne rottboellioides Desv., in LM and in SEM (type). Figs. 47, 48. Spore balls and spores of Sporisorium eriochrysis Vanky on Eriochrysis brachypogon (Stapf) Stapf, in LM and in SEM (type). Bars = 10 um.

56

Key to the smut fungi of Rhytachne 1. Sori on the leaves. Spore wall 2-8 um thick............ Tolyposporella rhytachnes Sori not on the leaves. Spore wall 0.5—1.5 yam thick......0.....00eccceeeseeeesceeeeteees 2 2. Sori in the whole inflorescence. Spores 11—15(—16) um long, dimorphic, outer spores prominently verrucose-echinulate ........... Sporisorium rhytachnes Sori in the ovaries. Spores 7.5—-13.5 um long, not dimorphic, finely, densely punctate ...............0....... Sporisorium rhytachnes-rottboellioidis

A SMUT FUNGUS ON ERIOCHRYSIS (GRAMINEAE)

Eriochrysis P. Beauv, in the subfam. Panicoideae, tribe Andropogoneae, subtribe Saccharinae, is a small, homogeneous genus of 7 species in Africa, tropical America and one species in India. It is allied to Saccharum (Clayton & Renvoize, 1986:331). No smut fungus has been previously reported on Eriochrysis.

Sporisorium eriochrysis Vanky, sp. nov. Typus in matrice Eriochrysis brachypogon (Stapf) Stapf (det. M. Namaganda, MHU), Uganda, Masaka Distr., 20 km E Masaka, Lake Nabugabo pr. Bale, 00°20'22" 8, 31°52'25" E, alt. cca. 1130 m.s.m., 26,]1.2002, leg. C., T. & K. Vanky. Holotypus in Herbario Ustil. Vanky, HUV 19995, isotypi in MHU, BPI, K et in Vanky, Ust. exs. no. 1178.

Sori in spiculis et sessilibus et pedicellatis, organa floralia intima destruentes, ovoidei usque cylindrici, 0,7~1 x 3-6 mm, glumis partim obtecti, primum peridio flavidobrunneo cooperti, quo itegulariter rupto massam nigrobrunneam, semiagglutinatam usque pulveream glomerulorum § sporarum laxorum sporarumque, columellam centralem flagelliformem circumdantium ostendentes. Glomeruli sporarum globosi, elongati usque itregulares, 30—70 x 40-80 um, atro- flavidobrunnei usque subopaci, e sporis pluribus decem, leviter separabilibus compositi. Sporae globosae, subglobosae, ovoideae, ellipsoideae usque parum irregulares, raro elongatae, 9-11 x 10-12(-13) pm, flavidobrunneae; pariete inaequaliter crasso, 0,5—1(—1,5) um, leniter, dense punctato-verruculoso; imago obliqua sporarum levis. Cellulae steriles non observatae.

Sori (Fig. 42) in both sessile and pedicelled spikelets, destroying the innermost floral organs, ovoid to cylindrical, 0.7—1 x 3-6 mm, partly hidden by the glumes, first covered by a yellowish-brown peridium which splits irregularly disclosing the blackish-brown, semiagglutinated to powdery mass of loose spore balls and spores, surrounding a flagelliform, central columella. Spore balls (Figs. 47, 48) globose, elongate to irregular, 30-70 x 40-80 ym, dark yellowish-brown to subopaque, composed of tens of spores which separate easily. Spores (Figs. 47, 49) globose, subglobose, ovoid, ellipsoidal to slightly irregular, rarely elongated, 9-11 x 10-12(—-13) um, yellowish-brown; wall unevenly thick, 0.5—-1(—1.5) pm, finely, densely punctate-verruculose; spore profile smooth. Sterile cells not seen.

On Eriochrysis brachypogon (Stapf) Stapf; C. Africa (Uganda).

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NEW COMBINATIONS

Macalpinomyces pretoriensis (Pole-Evans) Vanky, comb. nov. Basionym: Ustilago pretoriensis Pole-Evans, in Sydow, H. & P., Ann. Mycol. 12:263, 1914. Sphacelotheca pretoriensis (Pole-Evans) Zundel, 1938:301. Sporisorium pretoriense (Pole-Evans) Vanky, 1998, nom. herb. Type on Panicum helopus Trin. var. glabrescens (K. Schum.) Stapf (Urochloa panicoides Beauv.), South Africa, Transvaal, Pretoria, 20.11.1914, A.O. D. Mogg, PREM 7408. Isotype in HUV 18004!

Sori in some flowers of an inflorescence transforming the inner floral organs into one or several, horn-shaped, cylindrical or twisted bodies, 0.3-2.5 x 5—40 mm, first covered by a greyish-brown peridium which shreds into numerous, long, filaments disclosing the semiagglutinated to powdery mass of spores and sterile cells. Spores single when mature, globose, subglobose to broadly ellipsoidal, 9-12.5 x 9.5-13 pm long, yellowish-brown; wall even, c. 1 wm thick, prominently, moderately densely echinulate, spore profile serrulate. Sterile cells in irregular groups, single cells subglobose, ovoid, ellipsoidal or subpolyhedrally slightly irregular, smaller than the spores, 5S—8 x 6—11 um, hyaline; wall c. 0.5 pm thick, smooth, content homogeneous.

On Panicum maximum Jacq., Urochloa panicoides Beauv. (U. helopus (Trin.) Stapf, Panicum helopus Trin. var. glabrescens (K. Schum.) Stapf), U. trichopus (Hohst.) Stapf; C. & S. Africa (South Africa, Zambia, Zimbabwe).

Savile, 1953:664, described Thecaphora apicis, forming spore balls in the ovaries of Carex pyrenaica Wahlenb. Species of Thecaphora are restricted to dicotyledonous host plants. Those on Cyperaceae belong to the genus Moreaua

Moreaua apicis (Savile) Vanky, comb. nov. Basionym: Thecaphora apicis Savile, Canad. J. Bot. 31:664, 1953. Type on Carex pyrenaica Wahlenb., Canada, British Columbia, 15 miles NW of Keremeos, Apex Mountain, 6.[X.1951, H.A. Senn, C. Frankton & J.M. Gillett, DAOM 34034!

Sori destroying the nutlets, globoid, c. 1 mm in diameter, blackish-brown, first covered by a thin layer of host epidermis which early dehisces exposing the powdery mass of spore balls. Spore balls globose, ellipsoidal to irregular, 9-25 x 12-30 um, yellowish-brown, composed of (—1)2—20 spores. Spores variable in shape and size, globoid, ellipsoidal to irregular, with slightly flattened sides, sometimes subcuneiform, 5.5—9.5 x 6.5-14.5 pm, yellowish-brown; wall even, 0.5-0.8 ym thick (in lactophenol, with time, much swollen, up to 3 ym), free surface rough, in SEM finely, densely verruculose.

On Cyperaceae: Carex pyrenaica Wahlenb.; Europe (Mt. Pyrénées), N America (Canada).

58

Viennot-Bourgin, 1958:170, described Sphacelotheca elymandrae in the ovaries of Elymandra androphila. Species of Sphacelotheca are restricted to members of the Polygonaceae family. Study of the type of S. elymandrae showed that it belongs to the genus Macalpinomyces.

Macalpinomyces elymandrae (Viennot-Bourgin) Vanky, comb. nov. Basionym: Sphacelotheca elymandrae Viennot-Bourgin, Rev. Pathol. Vég. Entomol. Agric. France 37:170, 1958. Type on Elymandra androphila Stapf, Guinea, near Boké, XI.1956, leg. Jacques-Félix. Holotype in Herb. Viennot- Bourgin, PC, isotype in HUV 15801!

Sori in some ovaries of an inflorescence, swollen, ovoid, ellipsoidal, often apically narrowing into an acute tip, 8-12 x 10-15 mm, covered by a thin, _ yellowish-brown peridium of host and fungal origin which ruptures at maturity disclosing the dark brown, powdery mass of spores intermixed with groups of sterile cells. No conspicuous columella only a short prolongation of the floral axis mixed with fungal elements. Spores yellowish-brown, globose, subglobose to ellipsoidal, 9.5—13.5 x 10.5—15 um, including the 1-1.5 pm high, conical spines. Sterile cells in irregular groups, single cells variable in shape and size, globose, ellipsoidal to irregular, 5-16 pm long, subhyaline, collapsed in old specimen; wall thin (c. 0.5 um), even, smooth.

On Gramineae: Elymandra androphila Stapf; W. Africa (Guinea). Known only from the type collection.

Sporisorium argentinum (Hirschh.) Vanky, comb. nov. Basionym: Crozalsiella argentina Hirschhorn, Notas Mus. La Plata, Bot. 5:236, 1940. Sphacelotheca argentina (Hirschh.) Zundel, 1953:81. Type on Panicum demissum Trin. (= P. sabulosum Lamark), Argentina, Buenos Aires, Hudson, Villa Elisa, X.1924, J.B. Marchionatto, LPS 3649!

Sori at the base of the last upper internode, probably comprising the young inflorescence, globose or ovoid, 3—5 x 3-5 mm, with a long, apiculate host plant remnant on its distal part, covered by host tissues and a brown peridium which ruptures irregularly disclosing the dark brown, semiagglutinated mass of spores and sterile cells surrounding several short columellae. Spores single when mature, subglobose, broadly ellipsoidal or often kidney-shaped, 5.5-9.5 x 8—-10.5 um, yellowish-brown; wall evenly thick (c. 0.5 wm), finely, densely punctate-verruculose, spore profile smooth. Sterile cells 8-15 um long, in small groups, collapsed in old specimen, hyaline; wall thin, c. 0.5 tm, smooth.

On Gramineae: Panicum sabulosum Lamarck (P. demissum Trin.), and P. urvilleanum Kunth; Argentina, Uruguay.

Sporisorium anadelphiae (Viennot-Bourgin) Vanky, comb. nov. Basionym: Sorosporium anadelphiae Viennot-Bourgin, Bull. Soc. Bot. France 104:266, 1957. Type on Anadelphia pumila Jacques-Félix, Guinée francaise [= Guinea], Foulaya near Kindia, 1.1957, G. Viennot-Bourgin. Holotype in PC, isotype in HUV 15800!

39

Sori in some spikelets of an inflorescence, destroying the innermost floral organs, long cylindrical, 0.5—-1.5 x 2-14 mm, covered by a thick, yellowish- brown peridium which ruptures longitudinally from its apex into 2-3 bands, disclosing the blackish-brown, granular-powdery mass of spore balls surrounding a stout central columella. Spore balls variable in shape and size, globoid, ellipsoidal, elongated to irregular, subpolyangular, 30-125 x 40-160 um, dark reddish-brown, composed of tens to hundreds of spores which separate by pressure. Spores dimorphic. Outer spores globoid, ellipsoidal to subpolyhedrally irregular, 9.5—13(—15) x 10.5—16 ym, dark reddish-brown; wall uneven, 0.8-2.5 um thick, densely, prominently verrucose on the free surface, warts up to 1 um high, spore profile serrulate. Inner spores subglobose, ellipsoidal, usually rounded, subpolyhedrally irregular, 7-11.5 x 8-11(-12) um, pale yellowish- brown; wall even, c. 0.5 um thick, minutely, densely punctate or apparently smooth.

On Gramineae: Anadelphia pumila Jacques-Félix, Africa. Known only from the type locality.

Sporisorium panici-carthaginense (Spegazzini) Vanky, comb. nov. Basionym: Ustilago panici-carthaginenis Spegazzini, Anales Mus. Nac. Buenos Aires, Ser. 2, 6:207, 1899 (as "panici-carthagenenis"). Sorosporium panici- carthaginense (Speg.) Zundel, 1953:68 (as "panici-carthaginenis"). Type on Panicum carthaginense Arechav. (= P. fultum Hack.), Uruguay, Cerro de Montevideo, summer 1882-91, leg. J. Arechavaleta & C. Spegazzini, LPS (?)3038!; topotype on P. fultum Hack., V1.1927, G. Herter, HUV 5216!

Sori destroying the whole inflorescence and also the basal part of the terminal leaf sheaths, swollen, globoid, 2-5 mm in diameter, with an acute tip bearing a more or less well-developed leaf blade, compact, covered by a peridium of fungal and host tissues which ruptures usually transversally at its base disclosing the powdery, brown mass of spore balls, spores, sterile cells and several, short, stout, bent columellae. Spore balls composed of many, easily separating spores. Spores single when mature, variable in shape and size, ovoid, ellipsoidal, irregular, often with a flattened side, sometimes elongated into a subacute tip, (6.5—)7—9.5(—11) x 8.5—13 um, yellowish-brown; wall even, c. 0.5 ym thick, finely, densely punctate to finely verrucose-echinulate, spore profile smooth to finely wavy. Sterile cells in irregular groups, single cells subglobose, ellipsoidal to irregular, 8-15 pm long, hyaline; wall 0.5—1 ym thick, smooth, collapsed in old specimens.

On Gramineae: Panicum fultum Hack. (P. carthaginense Arechav., not Swartz, P. fasciculatum Swartz var. carthaginense Arechav.);, Uruguay.

Sporisorium parodii (Hirschh.) Vanky, comb. nov. Basionym: Ustilago parodii Hirschhorn, Darwiniana 3:401, 1939 Type on Muhlenbergia diffusa Willd. (= M. schreberi Gmel.), Argentina, La Plata, Bosque, X.1936, L. R. Parodi, Herb. Hirschhorn 35, isotype in LPS 3055! Sori comprising the distal part of the shoots (inflorescence? and some terminal leaves), long cylindrical, 3-4 x 10-12 mm, first covered by a whitish-brown

60

peridium with the tips of 1-2 leaves. The peridium ruptures longitudinally disclosing the brown, granular to powdery mass of spores surrounding 8-12 filiform columellae of the length of the sorus. Spores single when mature, - globose, subglobose, ellipsoidal to slightly irregular, 8-12 x 8-15 pm, globose spores 8-12 um in diameter, yellowish-brown; wall even, 1—2 um thick, finely, densely verruculose, spore profile wavy to finely serrulate. Sterile cells not seen. Spore germination (Hirschhorn, 1939:405 & Fig. 4c) results in 4-celled basidia producing mycelia and also a basidiospore(?).

On Gramineae: Muhlenbergia schreberi Gmel. (M. diffusa Willd.); Argentina.

This fungus differs from all 18 smut fungi described on Muhlenbergia. Unfortunately, the host plant identity could not be verified from the material at hand.

Urelytrum Hack., in the subtribe Rottboelliinae, tribe Andropogoneae, has 7 species in tropical Africa (Clayton & Renvoize, 1986:361). Only one smut fungus was so far described from it:

Sporisorium urelytri (L. Ling) Vanky, comb. nov.

Basionym: Sorosporium urelytri L. Ling, Sydowia 7:156, 1953c. Type on Urelytrum giganteum Pilger, Sudan, Equatoria Prov., Meridi-Yei boundary, near Olo River, 4.V.1938, J.G. Myers 9003. Holotype in IMI 25276! Paratypes on Urelytrum giganteum, Congo, a la Grelo section, VII.1931, P. Quarré 2593, IMI 36152, BPI 180815; on Urelytrum stapfianum C.E. Hubb. (= U. digitatum K. Schum.), Angola, no date, coll. Gossweiler 9518, IMI 25277; U. stapfianum (=U. digitatum), Angola, Lalange Plateau, Kela, 3.1.1931, Gossweiler, IMI 36153.

Sori destroying the inner floral organs of both sessile and pedicelled spikelets, long ellipsoidal or cylindrical, with one or several, short, acute tips, c. 1 x 3-6 mm, partially exposed between the glumes, first covered by a thick, pale brown peridium which dehisces irregularly from its apex, disclosing the dark brown, semiagglutinated to powdery mass of spore balls surrounding a stout, central columella with short apical branches. Spore balls subglobose, ellipsoidal, elongated to usually irregular, 40-80 x 45-130 wm, dark reddish-brown, composed of tens of spores which separate by pressure. Spores subglobose, ellipsoidal, usually rounded subpolyhedrally irregular, 9-14 x 10—15(—16) pm, medium yellowish-brown; wall even, 0.5—0.8 um thick, finely, densely punctate- verruculose, spore profile smooth.

On Gramineae: Urelytrum giganteum Pilger, U. digitatum K. Schum. (U. stapfianum C.E. Hubb.), Africa (Angola, Congo, Sudan).

ACKNOWLEDGEMENTS I am grateful to Dr. S. Toth (Gédéll6, Hungary) for providing the Latin descriptions, to Dr. E. McKenzie (Auckland, New Zealand), and Dr. R. Berndt (Tubingen, Germany) for reading the manuscript and serving as pre-submission reviewers. Thanks are also due to the Directors and Curators of the Herbaria AMH, BPI, BR, BRIP, FH, GH, HCIO, IMI, LE for loans of specimens.

6]

LITERATURE CITED

Ahmad, S. 1956. Ustilaginales of West Pakistan. Mycol. Pap. 64:1-17.

Arthur, J.C. 1884. Memorandum of Iowa Ustilagineae. Iowa Agr. Coll. Dept. Bot. Bull. 1884:172-174.

Baccarini, P. 1906. Funghi dell’ Eritrea. Ann. Bot.(Rome) 4:269-277 + Pl. X.

1917. Funghi etiopici. - Ann. Bot. (Rome) 14:117-140.

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MY COTAXON

Volume LXXXV, pp. 67-75 January-March 2003

THREE INTERESTING THERMOPHILIC TAXA OF GYMNOPUS (BASIDIOMYCETES, TRICHOLOMATACEAE): G. PUBIPES SP. NOV., G. PUBIPES VAR. PALLIDOPILEATUS VAR. NOV. AND G. DRYOPHILUS VAR. LANIPES COMB. NOV.

ANTONIO ORTEGA Dpto. Biologia Vegetal, Facultad de Ciencias. E-18071 Granada, Spain; e-mail: aortegad@ugr.es

VLADIMIR ANTONIN Moravian Museum, Dept. of Botany, Zeln_ trh 6, CZ-659 37 Brno, Czech Republic; e-mail: vantonin@mzm.cz

FERNANDO ESTEVE-RAVENTOS Dpto. Biologia Vegetal, Facultad de Biologia. E-28871 Alcala de Henares, Spain; e-mail: fernando.esteve@uah.es

Abstract Three Gymnopus-taxa (Basidiomycetes, Tricholomataceae) were studied. One new species, G. pubipes, is described for a thermophilic fungus usually erroneously identified as G. benoistii. This study is based on studies of the original description of the latter species and recently collected material from Spain. A macroscopically different but microscopically identical fungus recorded in the Czech Republic, Romania, Slovakia and Sweden is described as G. pubipes var. pallidopileatus. A new combination, G. dryophilus var. lanipes. is also proposed and its neotype from a recent collection from Spain is designated here.

Key words: Agaricales, new taxa, type-revision, taxonomy, nomenclature.

Introduction

During the past years, we collected three taxa of the genus Gymnopus (Pers.) Roussel in Mediterranean and Central-European thermophilic ecosystems. They have been named Gymnopus (or Collybia) benoistii or G. (C.) dryophilus var. lanipes, respectively, in most recent mycological literature (MALENCON & BERTAULT 1975;

68

MORENO ET AL. 1990; ORTEGA ET AL. 1991; ANTONIN & NOORDELOOS 1997; BON 1999; HAUSKNECHT & KRISAI-GREILHUBER 2000). To clarify the taxonomic position of these taxa, we made a comparative study of collections from different European countries, as well as of the type material.

The type material of C. benoistii Boud. is apparently not preserved, at least not in Boudier’s herbarium deposited in the herbarium of the Museum of Natural History in Paris (PC), so this species has to be interpreted only according to the original description (BOUDIER 1900). Most fungi identified in recent years as C. benoistii (see references above) agree only partially with Boudier’s species, and his original diagnosis does not include data on some essential characters in Gymnopus taxonomy, e.g. the type of pileipellis or its reaction with KOH. Regarding those circumstances, we decided to describe the new species Gymnopus pubipes, which includes the concept of C. benoistii of many European mycologists.

The type collection of Marasmius dryophilus var. lanipes Malencon et Bertault preserved in the herbarium in Montpellier (MPU) was studied and compared with recent Italian and Spanish samples. Unfortunately, the holotype does not correspond to the original diagnosis (MALENCON & BERTAULT 1975: 362). Therefore, a neotype was designated from Spanish material, as no paratypic collections exist in Montpellier Herbarium.

Material and methods

Gymnopus pubipes and G. dryophilus var. lanipes have a wide distribution in the Mediterranean countries (southern Europe and northern Africa) and one variety of the first one (see below) also in thermophilous associations (oak and hornbeam-oak stands) in Central and Northern Europe (Austria, Czech Republic, Romania, Slovakia, Sweden). Most Spanish localities of the latter species were recorded in Mediterranean ecosystems in Andalucia (Granada, Malaga) and Castilla-La Mancha (Guadalajara), whereas the material of Gymnopus dryophilus var. lanipes comes from southern Spain (Andalucia, provinces of Granada, Malaga and Sevilla), as well as from Italy (material previously studied and published by LONATI 1986).

Microscopic slides of dried material were prepared with 5% NH,OH, Melzer’s reagent and Congo Red in 10% ammonia. Macroscopic reactions on the pileus of the samples were tested with a 10% KOH solution. Drawings were made with the aid of Zeiss (A.O. and F.E.-R.) and Olympus (V.A.) drawing tubes under an oil-immersion lens. Colours of basidiomata were compared with KORNERUP & WANSCHER (1973). Spore measurements are given according to HEINEMANN & RAMMELOO (1985). Herbarium acronyms are according to HOLMGREN ET AL. (1990). Abbreviations of authors of fungal names follow KIRK & ANSELL (1992).

Apart from other references cited in the text, we have followed the general taxonomic criteria proposed in the European Collybia s. 1. monograph by ANTONIN & NOORDELOOS (1997), and the essential treatise for Mediterranean agarics by MALENCON & BERTAULT (1975).

69

Gymnopus pubipes Antonin, A. Ortega et Esteve-Rav., sp. nov.

Misapplied names: Collybia benoistii Boud., Bull. Soc. mycol. France 16: 102, 1900; Gymnopus benoistii (Boud.) Antonin & Noordel., Mycotaxon 63: 363. 1997 s. auct. p. part., non s. Boud.

Pileo 10-40 mm lato, hemisphaerico vel convexo, dein applanato, centro applanato vel depresso, obscure brunneo vel purpureo-brunneo, pallescente pallide brunneo vel cremeo. Lamellis adnexis vel emarginatis, cremeis vel ochraceis. Stipite 20-60 = 2-7 mm, cylindraceo, cum pileo concolore, tomentoso. Sporis (5.0-)5.5-8.0 x 2.5-4.0 um, oblongis-ellipsoideis, sublacrimoideis. Cheilocystidiis 20-40 x 5.0-20 um, clavatis, cylindraceis, subcapitatis, fusiformibus, frequenter irregularibus. Pileipellis e cellulis similibus sicut in Gymnopus dryophilus constructa. Caulocystidiis cylindraceis, clavatis, obtusis, 3.0-8.0 um latis.

Holotypus: Hispania, Huétor-Santillan, area protecta Sierra de Huétor, sub Quercus ilex subsp. ballota, 18. X. 1977 leg. G. Malencon, X. Llimona & A. Ortega (holotypus in herbario GDA 10332 asservatur, isotypi in herbaria AH 26979 et BRNM 670685 asservantur).

Selected descriptions and iconography: Malencon & Bertault (1975: 385-387, fig. 81, pl. 15, as Marasmius terginus f. benoistii); Moreno et al. (1990: 118-120, as Collybia benoistii); Antonin & Noordeloos (1997: 97-98, fig. 29, pl. 24, as Gymnopus benoistii

psp3):

Pileus 10-40 mm, hemisphaerical to convex, expanding to applanate or slightly depressed with age, hardly or not umbonate, with involute then straight or slightly reflexed margin, hygrophanous and striate at the margin when moist, chestnut-brown (9E8) or purplish-brown (10E8) when moist, pallescent to pallid brown (6C8) or cream (5A3-4) on drying, smooth to slightly radially rugulose. Lamellae rather distant, L = 20- 25, 1 = 1-3, deeply emarginate to adnexed, subventricose, pale cream (SA3-4) to buff (6C6-8), often showing a pale reddish reflection, with entire to slightly sinuose edge. Stipe 20-60 x 2-7 mm, cylindrical, often compressed, tenacious, whitish to pale cream when young, becoming concolorous with pileus with age, entirely pubescent, densely tomentose towards the base, with fine white to cream coloured hairs; white rhizoids often present at the base. Context concolorous with surface. Smell and taste indistinct. Spores (5.0-)5.5-8.0 x 2.5-4.0 um, X = 6.5 x 3.4 um, E = 1.6-2.3, Q = 1.9, oblong- ellipsoid to sublacrymoid, smooth, hyaline, non-amyloid. Basidia 18-25 x 5.0-7.5 um, 4-spored, clavate. Basidioles 12-30 x 3.0-8.0 ym, cylindrical to clavate. Lamella edge sterile. Cheilocystidia 20-40 x 5.0-20 um, very variable in form, mostly clavate to subcapitate, cylindrical to fusiform, sometimes irregular, lobate or with several wide projections. Pileipellis with typical dryophila-structure, composed by cylindrical to inflated up to 15 um wide hyphae, with lobate to coralloid terminal elements up to 20 uum wide, showing incrusting pigment. Stipitipellis a cutis, of cylindrical, parallel, slightly thick-walled, up to 5.0 um wide hyphae. Caulocystidia very numerous, 3.0-8.0 uum wide, polymorphic, sinuose, usually cylindrical to clavate, with obtuse apex, slightly thick-walled, sometimes septate, sometimes with obtuse projections at the apex. Clamp-connections present in all tissues. Chemical reactions: KOH reaction on pileus green.

Gymnopus pubipes (holotype): pileipellis, cheilocystidia, spores and caulocystidia. Scale bar = 20 um.

Ecology: Often gregarious, saprotrophic on vegetal debris (leaves, branches), especially of Quercus or Pinus, or in mixed stands, sometimes under Juniperus, in thermophilous forests on calcareous soils.

Distribution and phenology: Widespread in southern Europe and northern Africa. Autumn to early winter in the Mediterranean region.

Material studied: SPAIN: Granada: Huétor-Santillan, Parque natural de la Sierra de Huétor, road to campamento de la Alfaguara, under Quercus ilex subsp. ballota, 18 Oct. 1977 leg. G. Malencgon, X. Llimona and A. Ortega (holotype GDA 10332, isotypes AH 26979 and BRNM 670685); Baza, Parque natural de la sierra de Baza, pinar de Cala, under Juniperus communis, 3 Nov. 1992 leg. A. Ortega and D. Sousa (GDA 36231); Lanteira, Parque natural de Sierra Nevada, under Salix atrocinerea in Pinus forest, 13 Nov. 1999 leg. A. Capilla, F. Esteve-Raventos and A. Ortega (GDA 44742); Juviles, Parque natural de Sierra Nevada, under Pinus sylvestris, 28 Oct. 2000 leg. L. Alcoba and A. Ortega (GDA 44743); Guadalajara: Albalate de Zorita, under Quercus ilex

A

subsp. ballota, 15 Oct. 1999 leg. V. Bandala, F. Esteve-Raventés, L. Montoya and M. Villarreal (AH 26931).

The fungus described above represents a Mediterranean type” of this species. A fungus with totally identical microscopic features grows in thermophilous stands of Central Europe and southern parts of northern Europe. However, it differs by macroscopic features, especially the colour of carpophores. Therefore, we decided to describe it as a new variety of G. pubipes.

Gymnopus pubipes var. pallidopileatus Antonin, A. Ortega et Esteve-Ray., var. nov.

A varietate typica carpophoris parvis et pallide coloratis differt.

Holotypus: Cechia, Moravia, Ochoz prope Brno, area protecta Hornek, ad folia deiecta Quercus, 20. VI. 2001 leg. A. Vagner (holotypus in herbario BRNM 667673 asservatur).

Selected descriptions and iconography: Antonin & Noordeloos (1997: 97-98, fig. 29, pl. 24, as Gymnopus benoistii p. p.).

It differs from the typical variety in having a smaller, 15-30 mm broad pileus, coloured brown (6C5) at centre, paler (pale cream, beige or whitish) towards margin, paler lamellae (paler than 4A2) and paler stipe, almost whitish above and brown (6C4-5) towards base. Its microscopic features fully agree with var. pubipes.

Ecology: It grows on fallen Quercus-leaves, rarely on leaves of other broad-leaved trees (Acer campestre, Carpinus).

Distribution and phenology: It is known from Central Europe (Austria, Czech Republic, Romania and Slovakia) and southern Sweden. It has been recorded in spring and summer (May to August).

Material studied: CZECH REPUBLIC: Hlasna Tieban, 27 May 1945 leg. V. Vacek (PRM); Ochoz near Brno, Udoli Riéky nature reserve, Lysa hora, on fallen leaves and twigs in a Carpinus-Quercus stand, 19 July 2000 leg. V. Antonin and A. Vagner (Antonin 00.25, BRNM 652808); Ochoz near Brno, Hornek nature reserve, on fallen leaves of Carpinus, Quercus and Acer campestre, 29 July 2000 leg. A. Vagner (BRNM 652989) and 20 June 2001 leg. A. Vagner (BRNM 667673, holotype); Moravsky Krumlov, Krumlovsko-rokytenské slepence nature reserve, on fallen leaves of Quercus, 23 Aug. 2001 leg. V. Antonin and Z. Bieberova (Antonin 01.210, BRNM 666476); Kobyli na Mor., Ochozy, on Quercus leaves, 12 June 1994 leg. V. Antonin 94.53 (BRNM 599017); Valtice, Rendezvous, in Quercus leaves, 18 June 1993 leg. H Deckerova (Antonin 93.30 and 31, BRNM 576463 and 576464). ROMANIA: Bucuresti, destination Pitesti, 14 July 1976 leg. J. Kubi¢ka (PRM). SLOVAKIA: Nitra, Zobor, 3 June 1972 leg. L. Opold (BRA). SWEDEN: VéAstergétland, Létene, Medelplana, Munkangarnas naturreservat, on Quercus leaves, on limestone, 29 July 1985 leg. L. & A. Stridvall 83/003 (herb. Stridvall, L, BRNM 603950).

Gymnopus pubipes is characterised by a more or less uniformly reddish-brown pileus (var. pubipes) or a pileus brown at centre and beige, pale cream to whitish

72

towards margin (var. pallidopileatus), a strongly pubescent to velutinous stipe, a dryophila-type pileipellis, clavate to subcapitate, cylindrical to fusiform, sometimes irregular cheilocystidia and a green reaction of the pileus surface and pileipellis with KOH. Gymnopus fuscopurpureus (Pers.: Fr.) Antonin et al. (= Collybia obscura J. Favre) represents a close taxon. It can sometimes be found in the same habitats and shows the same green reaction on the pileus with KOH. The species differ in the colours of the basidiomata, stipe covering and size and shape of cheilocystidia. Gymnopus pubipes belongs to the subsect. A/kalivirentes Antonin et Noordel. having a dryophila- type pileipellis and green chemical reaction on the pileus (ANTONIN & NOORDELOOS loc. cit.). |

As previously stated in the introductory chapter, G. pubipes has regularly been identified as Collybia (or Gymnopus) benoistii (MORENO ET AL. 1990; ANTONIN & NOORDELOOS 1997) or Marasmius terginus f. benoistii (Boud.) Malencon et Bertault (MALENCON & BERTAULT 1975). All these authors indicate a spore size ranging (5.5-)6- 9(-11) x 3-4(-5) pm, although information about the KOH reaction on the pileus is lacking or said to be absent (ANTONIN & NOORDELOOS loc. cit.). The most recent record of this fungus was published by HAUSKNECHT & KRISAI-GREILHUBER (2000) with a sporal range of 6-8 < 3-3.5 um, which agrees with our observations (it represents var. pallidopileatus). BON’s (1999) concept of C. benoistii is rather similar to that of BOUDIER (1900) because he refers to a species with much larger spores [(8-)9-10(-11) x (3.5-)4-5(-6) pm].

MALENCON & BERTAULT (loc. cit.) and ORTEGA ET AL. (1991) subordinated C. benoistii to Marasmius terginus (Fr.) Fr. or Collybia tergina (Fr.) S. Lundell as a form or variety, respectively. However, the pileipellis of the tergina group 1s rather different from our former concept of C. benoistii (= Gymnopus pubipes), as already remarked by ANTONIN & NOORDELOOS (loc. cit.).

Unfortunately, the absence of type material in Boudier’s herbarium does not allow to have a correct picture of what C. benoistii might represent mainly regarding the reaction of the pileus with KOH, the type of pileipellis and the spore size, which are very important characters in Gymnopus taxonomy. Boudier’s species could also be related to some other taxon of Gymnopus sect. Vestipedes (Fr.) Antonin et al. or sect. Levipedes (Fr.) Halling and, in our opinion, should be best regarded as a nomen dubium.

Gymnopus dryophilus var. lanipes (Malencon et Bertault) A. Ortega, Antonin et Esteve-Rav., comb. nov.

Basionym: Marasmius dryophilus var. lanipes Malencon et Bertault, Trav. Inst. Sci. Cherif. Fac. Sci. Rabat 33 (Fl. Champ. Supér. Maroc 2): 362, 1975.

= Collybia dryophila var. lanipes (Malen¢on et Bertault) A. Ortega et Vizoso, Doc. Mycol. 21(82): 23, 1991.

Neotype (selected here): Spain, Malaga, Road Malaga-Colmenar, venta de Garvey, in Pinus forest, 10 Nov. 2000, leg. L. Alcoba and A. Ortega, GDA 44739 (isoneotypes AH 26980 and BRNM 670686).

Selected descriptions and iconography: Malencon & Bertault (1975: 361-364, fig. 77, as Marasmius dryophilus var. lanipes); Lonati (1986: 15-17, as Marasmius dryophilus var. lanipes).

74)

Pileus 12-45 mm, convex to plano-convex, later applanate or slightly depressed around the centre, with inflexed, then straight margin, hygrophanous, translucently striate at the margin (sometimes reaching half of the pileus) when moist, entirely orange-brown to ochraceous-brown when humid (7-8E8), strongly pallescent on drying, becoming yellow-ochraceous to cream (5A3-4), smooth, glabrous. Lamellae moderately crowded, adnate to emarginate, narrow, segmentiform or slightly ventricose, whitish (5A1), with entire to subentire concolorous edge. Stipe 30-65 x 3-8.5 mm, cylindrical to broadened towards the base, subconcolorous to paler than the pileus, brown-orange to ochraceous- yellow, whitish at the apex, surface finely pubescent, especially near the base, never tomentose, whitish rhizoids present at the base. Context concolorous with surface. Smell and taste indistinct.

Spores (3.8-)4.5-7.2(-8.0) x 2.2-3.5 um, Xm = 5.2 x 2.8 um, E = 1.6-2.3, Q = 1.85, oblong-ellipsoid to (sub-)lacrimoid, smooth, hyaline, non-amyloid. Basidia 23-31 x 5.0- 6.0 um, 4-spored, clavate. Basidioles 15-32 x 2.5-7.0 um, cylindrical to clavate, less frequently subfusoid. Cheilocystidia rather inconspicuous, 18-25 x 4.0-7.0 um, cylindrical, more rarely subclavate, mostly furcate to lobate, irregular. or diverticulate. Trama hyphae cylindrical, non-dextrinoid, up to 12 um wide. Pileipellis typically “Sigsaw puzzle-like”(dryophila-type), with strongly diverticulate elements, smooth or incrusted. Stipitipellis a cutis, of cylindrical, parallel, slightly thick-walled, up to 5.0 um wide hyphae. Hairs of the stipe covering up to 50 x 1.0-5.0 um, filiform or vermiform, thin- to moderately thick-walled, arising from clusters of diverticulate terminal elements, similar to cheilocystidia. Clamp-connections present in all tissues.

Chemical reactions: KOH on pileus none.

Ecology: Gregarious in Mediterranean thermophilous forests among vegetal debris, especially of Quercus ilex, Pinus, or more rarely Cistus.

Distribution and phenology: Known from northern Africa (Mauritania), and southern Europe (France, Italy and Spain).

Material studied: ITALY: Roma: Villa Ada, under Robinia pseudoacacia, Laurus nobilis and Quercus ilex, 8 Nov. 1983 leg. G. Lonati (herb. Lonati, duplicate AH). SPAIN: Granada: Sierra Elvira, under Quercus ilex subsp. ballota, 28 Oct. 1979 leg. R. Galan (GDA 10328); Bolones, Parque natural de la Sierra de Huétor, under Pinus halepensis, 2 Jan. 1996 leg. L. Alcoba and A. Ortega (GDA 42754, AH 25268); Malaga: Yunquera, parque natural de la Sierra de las Nieves, puerto de la Caina, under Cistus albidus, 30 Nov. 1995 leg. F. Esteve-Raventos, E. Horak, G. Moreno and A. Ortega (GDA 44974); Road Malaga-Colmenar, venta de Garvey, in Pinus forest, 10 Nov. 2000 leg. L. Alcoba and A. Ortega (neotype GDA 44739, isoneotypes AH 26980 and BRNM 670686); Sevilla: Aznalcazar, pinar de Aznalcazar, under Pinus pinea, 18 Nov. 2000 (GDA 44740) and 31 March 2001 (GDA 44741) leg. L. Alcoba and A. Ortega.

This poorly known taxon was described in detail by MALENCON & BERTAULT (1975), putting special emphasis to the presence of a finely velutinous stipe, the covering formed by cylindrical, thin-walled hairs or caulocystidia. There is no doubt that the original description represents the well-known G. dryophilus, except for the pubescent stipe. The material studied by us differs from the typical G. dryophilus also in having smaller cheilocystidia and longer basidia.

74

fs Sei WO |

Gymnopus dryophilus var. lanipes (neotype): pileipellis cells (upper left), cheilocystidia (lower left), spores and hairs of the stipe cover (right). Scale bar = 20 um.

The material deposited as a holotype (n° 4508) in Malencon’s herbarium at Montpellier (MPU) was studied by us, and it does not correspond to the taxon described in the protologue. Although it has a pubescent stipe, its spores are larger (9-11 =< 4-5 um), the pileipellis is a cutis of typical filamentous hyphae without a dryophila- structure, and the basidiomata are brown-reddish. These characters do not agree with G. dryophilus, but it may represent a taxon from the G. terginus group. Because Malencgon knew G. terginus very well, a designation of such material was surely an unintentional error. Therefore, because of the absence of any paratypic collection, we selected a neotypus from Iberian material which fits the original description in all aspects, and keeps the originally used name.

Gymnopus dryophilus var. lanipes might be confused with pale or dehydrated forms of G. pubipes, both sharing a pubescent stipe. Gymnopus dryophilus var. lanipes can be distinguished macroscopically by the whitish to cream lamellae and the absence of rhizoids at the stipe base, and microscopically by its smaller spores and the inconspicuous cylindrical, often diverticulate cheilocystidia. Moreover, the chemical reaction with KOH on the pileus is a good character to distinguish both species.

Acknowledgements

A. Ortega wishes to thank the CICYT (Ministerio de Educacién y Cultura) for granting the research project PB 98-1316, in which these results are included. F. Esteve- Raventos thanks the Consejo Social of Alcala de Henares University for finantial support concerning a visit to Montpellier Herbarium (MPU), and also to the director and staff at MPU for the facilities provided during his stay to study Malencgon’s collections. All authors also thank Jan W. Jongepier (Veseli nad Moravou, Czech Republic) for correcting the English manuscript.

75

References

Antonin V. & Noordeloos M. E. (1997): A monograph of Marasmius, Collybia and related genera in Europe. Part 2: Collybia, Gymmopus, Rhodocollybia, Crinipellis, Chaetocalathus and additions to Marasmiellus. Libri Botanici, vol. 17. IHW-Verlag. 256 pp.

Bon M. (1999): Flore Mycologique d’Europe. 5. Les Collybio-Marasmioides. Doc. Mycol., Mém. hors sér. 5: 1-171.

Boudier E. (1900): Champignons nouveaux de France. Bull. Soc. mycol. Fr. 16: 193-200.

Hausknecht A. & Krisai-Greilhuber I. (2000): Riiblinge, Schwindlinge und verwandte Taxa in Ostésterreich. Osterr. Z. Pilzk. 9: 31-66.

Heinemann P. & Rammeloo J. (1985): De la mesure des spores et de son expression. Agarica 6: 366-380.

Holmgren P. K., Holmgren N. H. & Barnett L. C. (1990): Index herbariorum. Part I. The herbaria of the world. 8" ed. New York Botanical Garden, New York.

Kirk P. M. & Ansell A. E. (1992): Authors of fungal names. Index of Fungi Suppl. C. A. B. International, Oxon.

Kornerup A. & Wanscher J. H. (1973): Petit lexique des couleurs (ed. frang.). - Musterschmidt- Verlag. Ziirich-G6ttingen. 245 pp.

Lonati G. (1986): Altri due miceti maroccani Laziali. Marasmius dryophilus var. lanipes Malencon et Bertault. Hebeloma pallidum Malencgon. Boll. Assoc. Micol. Ecol. Romana 5: 15-18.

Malencon G. & Bertault R. (1975): Flore des champignons superieurs du Maroc, vol. II. Fac. Sci. Rabat. 540 pp.

Moreno G., Esteve-Raventos F. & Ilana C. (1990): Estudios micoldgicos en el parque natural de Monfragiie y otras zonas de Extremadura (Espafia), IV. Agaricales. Bol. Soc. Micol. Madrid 14: 115-142.

Ortega A., Vizoso M. T. & Contu M. (1991): Notas sobre la micoflora xero-termofila y sabulicola de Andalucia (primera parte). - Doc. Mycol. 16 (82): 14-42.

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MYCOTAXON

Volume LXXXV, pp. 77-79 January-March 2003

PHALLUS PYGMAEUS, A NEW MINUTE SPECIES FROM A BRAZILIAN TROPICAL RAIN FOREST

IURI GOULART BASEIA!, 2TATIANA BAPTISTA GIBERTONI & 3LEONOR COSTA MAIA

Universidade Federal de Pernambuco, Centro de Ciéncias Biolégicas Departamento de Micologia, Av. Prof. Moraes Rego, 1235 Cidade Universitaria, 50670-901, Recife - PE, Brazil libaseia@bol.com.br, *tbgibertoni@hotmail.com, *leonoremaia@hotmail.com

ABSTRACT - Phallus pygmaeus, a remarkable new species fundamentally characterized by its basidiomata not exceeding 10 mm high, smooth surface of the dark pileus and lignicolous habitat, is described and illustrated.

KEYWORDS - Neotropics, Phallales, gasteromycetes, taxonomy INTRODUCTION

Recent collecting activities in Northeastern Brazil have revealed a rich macromycete biota (Gibertoni & Cavalcanti 2000; Baseia & Galvao 2002; Baseia & Milanez 2002). Ongoing taxonomic studies in this poorly explored neotropical region allowed the discovering of a new and interesting phalloid.

According to Kreisel (1996), in a preliminary survey of the genus Phallus Pers. sensu lato, 31 species were accepted; from those 3 were limited to the New World, 18 to the Old World, and 10 were known from both hemispheres, although in a few cases the apparent wide distribution may be due to introduction by man.

The most important morphological criteria for the infrageneric taxonomy of Phallus are the shape and surface configuration of the pileus, coloration of the receptacle and volva, and the size of the basidiomata (Ventenat 1798, Lloyd 1909, Dring & Rose 1976, Kreisel 1996).

MATERIALS AND METHODS

Collecting expeditions to the Gurjau Ecological Station, a remnant of the Atlantic rain forest, located in the State of Pernambuco (8°21'30°’- 8°12'00” S and 31°56'30’’- 35°45’30”’ W), were conducted during June 2002.

Basidiomata were examined in the field for their fresh characteristics. Colors were coded according to Kornerup & Wanscher (1978; KW’ code noted in parenthesis) and described subjectively. Basidiomata were dried slowly over a stove and placed in containers with naphthalene to prevent spoilage in the excessively humid conditions. In addition, some basidiomata were pickled fresh in FAA.

Microscopic anatomical details were determined from dried and pickled specimens with a Zeiss microscope with bright field and phase contrast optics.

78

Features were observed in 3% KOH and Melzer’s reagent. A total of 30 spores were measured from the holotype and subsequent collection. The material was preserved at the Herbarium Father Camille Torrend (URM).

TAXONOMIC SECTION

Phallus pygmaeus Baseia sp. nov. Figs 1-2 Ovo subgloboso, 2-3 mm diametro, epigeae, colore superius olivacea brunnea, inferius aureo albo, mycelli filamentis robustis; stipite cylindrico, reticulato, colore albo, 5-10 mm alto, 1-2 mm lato; pileo campanulato, apice cum orificio, superficie gleba gelatinosa, laevi, olivacea brunnea; sporis ellipsoideis, laevis, 3-4 x 1-1,5 um, hyalinis. Carnis putridae odore.

Ad lignum putridum. Holotypus: Brasil, Pernambuco, Gurjau, 28/V1I/2002, leg. I. G. Baseia and T. B. Gibertoni, in Herb. URM ne 77078 conservatus est.; isotypus: URM ne 77079.

Egg subglobose, 2-3 mm diam., epigeal, olive brown (KW 4F4) at its apex, yellowish white (KW 4A2) at the lower part, mycelial cord strong; stipe cylindrical, reticulate, not pigmented, 5-10 mm high, 1-2 mm wide; pileus campanulate, apex perforate, glebal surface gelatinose, smooth, olive (KW IF5); spores ellipsoid, smooth, 3-4 x 1-1.5 um, hyaline. Beef cattle smell.

The epithet ‘pygmaeus’ refers to the very small basidiomata of this species.

Material studied: the holotype and isotype.

Habitat: groups on rotten woody.

Distribution: Brazil, State of Pernambuco.

Remarks: Gaudichaud (1827) described Sophronia brasiliensis Gaudich., which subsequently was considered a synonym of Phallus indusiatus Vent. Pers. Later, Moller (1895) described four new Brazilian phalloid species: Aporophallus subtilis A. Moeller (=Phallus subtilis (A. Moller) Lloyd), [tajahya glutinolens A. Moller (=P. glutinolens (A. Moller) O. Kuntze), /. galericulata A. Moller (=P. galericulatus (A. Moller) Kreisel) and Dictyophora callichroa today considered as P. callichrous (A. Moller) Lloyd. Since then, no new species of Phallus have been described from Brazil.

Phallus pygmaeus is usually easy to distinguish on account of its small basidiomata, nature of the dark pileus and lignicolous habitat. This new species recalls P. impudicus L.: Pers. in shape, color and spore size, differing, however, in the following characteristics: basidiomata not exceeding 10 mm high, and smooth surface of the pileus.

According to Kreisel (1996), the genus Phallus is divided in 5 subgenera: Aporophallus (A. Moller) Kreisel, Jtajahya (A. Moller) Kreisel, Endophallus (M. Zang & R. H. Petersen) Kreisel, Satyrus (Bosc) Kreisel, and Phallus. Members of the subgenus Phallus have a campanulate pileus with perforate apex and yellow pigments, characteristics observed in P. pygmaeus. However, this taxon also possesses some common characteristics with subgenus Aporophallus, such as a campanulate pileus with a smooth surface; indusium absent and receptacle without pigments. It differs from subgenus Aporophallus by having an apical pore in the pileus. Following the

19

subgeneric concepts of Kreisel (1996), P. pygmaeus forms an interesting link between subgenus Phallus and subgenus Aporophallus.

Figs 1-2 Phallus pygmaeus: |. mature and immature basidiomata; 2. basidiospores

ACKNOWLEDGMENTS

We acknowledge the support from the Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico (CNPq). Thanks to Dr. Francisco Calonge and Dr. Paul Kirk for critically reviewing the manuscript.

REFERENCES

Baseia, I. G. & Galvao, T. C. O. 2002. Some interesting Gasteromycetes (Basidiomycota) in dry areas from northeastern Brazil. Acta Bot. Bras. 16(1): 1-8.

Baseia, I. G. & Milanez, A. I. 2002. Geastrum setiferum (Gasteromycetes): a new species with a setose endoperidium. Mycotaxon 84: (in press).

Dring, D. M. & Rose, A. C. 1976. Additions to West African Phalloid Fungi. Kew Bull. 31 (3): 741-751.

Gaudichaud, C. B. 1827. Voyage Monde, Uranie Physicienne, Bot.: 179.

Gibertoni, T. B. & Cavalcanti, M. A. Q. 2000. Novos registros de Aphyllophorales para o Brasil. Acta Bot. Bras. 14(3): 267-271.

Komerup, A. & Wanscher, J. 1978. Methuen Handbook of Colour, 3" ed., Methuen, London,

252 pp.

Kreisel, H. 1996. A preliminary survey of the genus Phallus sensu lato. Czech Mycol. 48: 273- 281.

Lloyd, C. G. 1909. Synopsis of the known Phalloids. Bull. Lloyd Library no. 13: 1-96.

Moller, A. 1895. Brasilische Pilzblumen. Jena. 152 pp.

Ventenat, E. P. 1798. Dissertation sur le genre Phallus. Mém. Inst. Nat. Sci. Arts. Sci. Math. Phys. 1: 503-52

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MYCOTAXON

Volume LXXXV, pp. 81-84 January-March 2003

PARACERATOCLADIUM POLYSETOSUM, A NEW RECORD FROM BRAZIL

LUIS FERNANDO PASCHOLATI GUSMAO Universidade Estadual de Feira de Santana —- UEFS Departamento de Ciéncias Bioldgicas KMO03 BR116, 44.031-460

Feira de Santana, BA, Brazil (lgusmao@uefs.br) and

FABIO FERNANDES BARBOSA Universidade Federal da Bahia - UFBA Instituto de Biologia Rua Baréo de Geremoabo, s/n, Ondina, 40.170-290 Salvador, BA, Brazil

ABSTRACT

Paraceratocladium polysetosum has dark brown setae, monophialidic conidiogenous cells and hyaline, septate conidia. Brazilian material of this species differs in having polyphialidic conidiogenous cells. P. polysetosum from Brazil is illustrated, described, and compared with other Paraceratocladium species. Geographical distribution data is included.

Key words: Dematiaceous Hyphomycetes, mitosporic fungus, taxonomy

INTRODUCTION

During a survey of mitosporic fungi associated with leaf litter of native plants of “restinga” vegetation at the “Area de Prote¢ao Ambiental (APA) das Lagoas e Dunas do Abaeté”, Salvador, State of Bahia, the fungus Paraceratocladium polysetosum Castafieda was found.

The genus Paraceratocladium was erected by Castafieda (1987) and to date five species have been described: P. silvestre Castafieda (type species) and P. polysetosum Castafieda (Castafieda 1987), P. triseptata Dulymamode, W. Wu & Peerally (Dulymamode et ai. 1998), P. malaysianum Goh & Hyde (Goh & Hyde 2000), and P. seychellarum Whitton, McKenzie & K.D. Hyde (Whitton et al. 2001). The criteria used to delimit the genus are presence of simple or branched setae, monophialidic or polyphialidic conidiogenous cells, and ovoid, ellipsoid or narrowly ellipsoid, cylindrical, acicular or falcate, 0-3-septate, hyaline conidia. Both P. triseptata and P. seychellarum have been described with polyphialidic conidiogenous cells. The

82

Brazilian fungus also-has polyphialidic conidiogenous cells and thus differs from P. polysetosum as described by Castafieda (1987). A morphological comparison and distribution data for all species are presented in table 1.

Paraceratocladium polysetosum Castafieda, Fungi Cubenses II. INIFAT, Habana, Cuba: 9. 1987. (Fig. 1-6)

Setae solitary, erect, straight or flexuous, acerose, branched, septate, smooth, dark brown at the base and brown at apex, 185-300 x 5-8um; branches 29-86 x 2-3um; conidiophores irregulary branched, micronematous, arising on the surface of the setae and branches, smooth, pale brown; conidiogenous cells monophialidic or polyphialidic, usually intercalary, discrete, lageniform, smooth, pale brown 5-12 x 4.5-6.5um; collarette 1.5-2.5 x 2-2.5um; conidia solitary, aggregating in slimy masses, cylindric, base obtuse, apex slightly swollen, 1-septate, smooth, hyaline, 13.5-18 x 1.5-2.5um.

Material examined: on dead leaves of Manilkara salzmanii (DC.) H.J. Lam. (Sapotaceae), BRAZIL, Bahia, APA das Lagoas e Dunas do Abaeté, F.F. Barbosa, 3. V. 2002, HUEFS 56647. (permanent slides).

The Brazilian material has smaller setae and slightly longer conidia than those described in the literature. Castafieda (1987) described setae of P. polysetosum as up to 450um in length x 9-1241m wide at the base, and conidia as 14-17 x 2m. However, this variation is probably influenced by the substrate or environmental conditions.

The Brazilian material was collected from fragments of leaf litter put in damp chambers. P. polysetosum was unique species present in all fragments suggesting that this fungus is the most important species colonizing and decomposing leaf litter of Manilkara salzmanii.

Table 1. Morphological comparison and geographical distribution of Paraceratocladium species.

Species Setae Conidiogenous Conidia Distribution cells shape size (tm) P. malaysianum _ simple monophialidic acerose 40-46 x 1.5-2 Malaysia P. polysetosum branched Hoy Maen et cylindric 13.5-18 x 1.5-2.5 Cuba, Brazil polyphialidic P. seychellarum branched On op Eu icig, ovoid 3-512. x 12-15" Seychelles polyphialidic China, Cuba, P. silvestre simple monophialidic acicular 21-30 x 1-2 Hone Kone. Malaysia, Mauritius

P. triseptatum simple polyphialidic acerose 28-34x 1.5-2.5 Mauritius

Fig. 1-6. Paraceratocladium polysetosum. Fig.l. Branched setae with entwined conidiophore. Fig. 2. Monophialidic conidiogenous cell. Figs 3-5. Polyphialidic conidiogenous cells. Fig. 6. Conidia. Bars: Fig. 1 = 50um, Figs 2-5 = 10pm.

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ACKNOWLEDGEMENTS

The authors wish to thank Dr Eric McKenzie, Landcare Research, New Zealand for critical rewiew of this paper.

REFERENCES

CASTANEDA RUIZ, R.F. 1987. Fungi Cubenses II. Instituto de Investigaciones Fundamentales en Agricultura Tropical “Alejandro de Humboldt“. Habana. 22pp. . DULYMAMODE, R., WU, W. & PEERALLY, A. 1998. Three new Hyphomycetes on Pandanus leaves from Mauritius. Mycoscience 39: 285-291.

GOH, T.K. & HYDE, K.D. 2000. Paraceratocladium malaysianum sp. nov. from submerged wood in Malaysia, and a key to the genus. Nova Hedwigia 71: 95-100. WHITTON, S.R., McKENZIE, E.H.C. & HYDE, K.D. 2001. Microfungi on the Pandanaceae: Paraceratocladium seychellarum sp. nov. and a review of the genus. Fungal Diversity 7: 175-180.

MY COTAXON

Volume LXXXV, pp. 85-89 January-March 2003

BIOSYSTEMATICS OF THE MYXOMYCETES DIDYMIUM SQUAMULOSUM, PHYSARUM COMPRESSUM, AND PHYSARUM MELLEUM: ADDITIONAL ISOLATES

JIM CLARK Department of Biology, University of Kentucky, Lexington, KY 40506 STEVEN L. STEPHENSON Department of Biology, Fairmont State College, Fairmont, WV 26554

Abstract: Twenty-four isolates of Didymium squamulosum, 18 isolates of Physarum compressum, and two isolates of Physarum melleum were cultured and their reproductive system determined. These isolates are additions to the 85 isolates (33 D. squamulosum, 33 P. compressum, and 19 P. melleum) that have been studied previously. Eleven of the 24 D. squamu/osum isolates were nonheterothallic and presumptive apomictics, while the remaining 13 isolates belonged to three new biological species: A4 (three isolates from Costa Rica, sympatric with the Al and A2 biological species), A5 (one isolate from Mexico), and Aé6 (one isolate from Thailand, two isolates from New Zealand, and six isolates from Australia). Seventeen of the P. compressum isolates were nonheterothallic, with the remaining isolate from Costa Rica (CR 18) being compatible with the previously determined A1 biological species isolated from the Canary Islands. Both of the P. me//eum isolates were nonheterothallic. These results confirm and extend the conclusions derived from earlier studies. The D. sguamulosum morphospecies, which consists of two overlapping morphotypes, appears to be an extremely complicated species complex consisting of numerous geographically restricted sympatric and allopatric biological species and many apomictic lines. The biological species are usually, but not always, confined to one of the morphotypes. On the other hand, the P. compressum morphospecies seems to be a species complex consisting of a single uncommon but widespread biological species and countless apomictic lines with relatively minor morphological variations. The Physarum melleum morphospecies is probably similar to P. compressum, although to date no heterothallic isolates have been reported.

Key Words: apomixis, biological species, heterothallism INTRODUCTION

The taxonomy of the myxomycetes, at present, is based almost entirely on the morphology of the fruiting body and the use of the typological species concept (i.e., a particular species consists of individuals conforming to a constant morphological form that varies only within narrow boundaries). However, myxomycetes have a fairly restricted suite of morphological characters, and the discrete gaps between species necessary for the typological concept to be applied are often not present (Clark 2000). This, combined with the fact that many named species are variable and cosmopolitan and most of the rest are rare and of restricted distribution, has resulted in a rather muddled taxonomic situation that needs to be improved by introducing a more natural system based on experimental studies.

The myxomycete morphospecies Didymium squamulosum (Alb. & Schwein.) Fr., Physarum compressum Alb. & Schwein., and Physarum melleum (Berk. & Broome) Massee recently have been subjected to biosystematic studies (El Hage et al. 2000, Irawan et al. 2000, Clark and Stephenson 2000); however, the availability of additional isolates allows us to confirm and extend these previous studies. Didymium squamulosum has been reported to be a morphologically variable complex

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consisting of several biological species and numerous variable apomictic lines (El Hage et al. 2000), P. compressum has been reported to have little morphological variability and apparently consists of a single biological species with many closely related apomictic lines (Irawan et al. 2000), and P. melleum has been reported as consisting of numerous apomictic lines with relatively minor morphological variations (Clark and Stephenson 2000). Therefore, in an attempt to confirm these earlier observations and to produce a more solid foundation for an experimentally based taxonomy, additional isolates of these species were obtained and studied. These and other similar experimental studies may provide the necessary basis for a reevaluation of myxomycete taxonomy so that the present system can be replaced with a more natural system.

MATERIALS AND METHODS

The twenty-four isolates of Didymium squamulosum, 18 isolates of Physarum compressum and two isolates of Physarum melleum examined in this study were derived from moist chamber cultures, soil cultures or from field collections (TABLE I). Standard cultural and genetic procedures were used throughout the study (Clark 1995). In brief, plasmodia were grown on water agar (16 g plain agar per L of distilled water) and fed sterilized rolled oat flakes, and amoebae were grown on CM/2 agar (8 g corn meal agar and 8 g of plain agar per L of distilled water) with a film of sterile distilled water. Clonal amoebal populations were derived from isolated single spores or single amoebae. When the clonal amoebal populations produced plasmodia by themselves, they were classified as nonheterothallic, but when they required clonal mixing to produce plasmodia, they were considered heterothallic. Tester clones representing the two mating types from each heterothallic isolate were then crossed in all possible pair-wise combinations to determine multiple alleles (plasmodia produced in cross combinations with several different mating types) and biological species (plasmodia not produced in crosses between mating types of different isolates).

TABLE I Isolates and their origin

Isolates’ Location Collection? Substrate

Didymium squamulosum:

Aust 1 Cape Tribulation, Queensland* Stephenson 14962 aerial litter

Aust 2 Cape Tribulation, Queensland Stephenson 14963 aerial litter

Aust 3 Cape Tribulation, Queensland Stephenson 14964 aerial litter

Aust 4 Cape Tribulation, Queensland Stephenson 14965 aerial litter

Aust 5 Cape Tribulation, Queensland Stephenson 14966 _ aerial litter

Aust 6 Cape Tribulation, Queensland Stephenson 14967 __ aerial litter

Aust 7 Cape Tribulation, Queensland | Stephenson 14968 aerial litter

Aust 8 Cape Tribulation, Queensland Stephenson 14972 __ aerial litter

Aust 9 Cape Tribulation, Queensland Stephenson 14982 _ aerial litter

Aust 10 Cape Tribulation, Queensland Stephenson 14989A aerial litter

GRe29 Monteverde Reserve* Clark mc Heliconia leaf litter CR 30 Monteverde Reserve Clark mc Heliconia \eaf litter GR Monteverde Reserve Clark mc Heliconia \eaf litter CR 32 Monteverde Reserve Clark me Heliconia leaf litter Chos5 Monteverde Reserve Stephenson 12723 Heliconia \eaf litter CR 34 Culture contaminant Stephenson 14628

Ha 1 Kipuka Puaulu Bird Park’ Landolt soil culture

Mex 1 E] Edén, Quintana Rod Schnittler 16289 epiphyllic liter

NZ 1 Mill Bay, Auckland® Stephenson 14480 palm frond

Naz Mill Bay, Auckland Stephenson 14487 __ palm frond

Ore 1 Oregon Landolt epiphytic soil culture Tenn1 Cosby CG, GSMNP* Stephenson 11706 __ old straw bale

Thai | Kanchanaburi Falls Stephenson 9149 forest floor litter

87

Trin | Trinidad Clark mc Heliconia \eaf litter Physarum compresum:

Aust | Daintree National Park’ Stephenson 14516 _ aerial inflorescence Aust 2 Daintree National Park Stephenson 14625 aerial inflorescence Aust 3 Cape Tribulation, Queensland Stephenson 14973 _aerial litter Aust 4 Cape Tribulation, Queensland Stephenson 14984 _ aerial litter Aust 5 Cape Tribulation, Queensland Stephenson 14989B aerial litter

Aust 6 Daintree National Park Stephenson 14990 _aerial inflorescence CR 3 Costa Rica Clark mc DelMonte banana peel CR 4 Costa Rica Clark mc Chiquita banana peel CR 18 Monteverde Reserve Clark mc Heliconia leaf liter Col | Columbia Clark mc Turbana banana peel ECUBC Ecuador Clark mc Bonita banana peel Ecu 6 Ecuador Clark mc Bonita banana peel Gua | Guatemala Clark me DelMonte banana peel Guy 2 Guyana Landolt soil culture

Idn 3 Donayan/Kediri East Java Clark mc banana leaf litter

PR4 Puerto Rico Clark mc forest floor litter

Tenn | Cosby CG, GSMNP Stephenson 11704 litter

WV | Harrison Co., West Virginia Stephenson 11731 old field litter Physarum melleum:

Mex 1 El Edén, Quinitana Ro6é Stephenson 12626 palm aerial liter SC 1 Bishopville, South Carolina Stephenson 12289 dead leaf

* Isolates = Letters indicate country or state of origin: Aust for Australia, CR for Costa Rica, Col for Columbia, Ecu for Ecuador, Gua for Guatemala, Guy for Guyana, Ha for Hawaii, Idn for Indonesia, Mex for Mexico, NZ for New Zealand, PR for Puerto Rico, SC for South Carolina, Tenn for Tennessee, Thai for Thailand, Trin for Trinidad, and WV for West Virginia.

» Collection = Clark mc for moist chamber culture by Clark, Landolt for soil culture by Landolt, and herbarium specimens supplied by Stephenson or Schnittler.

° Cape Tribulation, Northern Queensland, Australia, aerial litter collected with a canopy crane.

4 Monteverde Cloud Forest Reserve in Costa Rica.

© Sporangia developed on litter collected in Australia, but the biological species is typical of Costa Rica and is most likely the results of cross contamination during moist chamber development. "Kipuka Puaulu Bird Park in Hawaii Vocanoes National Park.

® Mill Bay west of Auckland, New Zealand.

" Cosby Campground, Great Smoky Mountains National Park.

'‘ Ecolodge near Daintree National Park, Northern Queensland, Australia.

RESULTS

Reproductive systems Didymium squamulosum isolates Aust 3, Aust 6, Aust 7, Aust 9, CR 31,CR 32, CR 33, Ha 1, Ore 1, Tenn 1, and Trin 1 were found to be nonheterothallic. The 13 remaining D. squamulosum isolates were heterothallic and members of three different biological species. The CR 29, CR 30 and CR 34 isolates defined the A4 biological species and displayed six mating type alleles (A4! and A4?in CR 29, A4° and A4‘in CR 30, A4’ and A4° in CR 34). The lone Mex 1 isolate defined the AS biological species and displayed two mating types (A5'and AS’). The Thailand isolate, two New Zealand isolates, and six Australia isolates defined the A6 biological species, which displayed 12 mating types (Thai 1 = A6'and A6?, NZ 1 = A6*and A6*, NZ 2= A6’, Aust 1 = A6’and A6*, Aust 2 = A6’and A6"”, Aust 4 = A6°and A6°, Aust 5 = A6’, Aust 8 = A6® and A6'', Aust 10 = A6* and

88

A6'”) with the NZ 2 and Aust 5 isolates containing only a single mating type. Of the 18 Physarum compressum isolates, seventeen were nonheterothallic (Aust 1, Aust 2, Aust 3, Aust 4, Aust 5, Aust 6, CR 3, CR 4, Col 1, Ecu3c, Ecu 6, Gua 1, Guy 1, Idn 3, PR 4, Tenn 1, and WV 1), with CR 18 the lone heterothallic isolate. CR 18 displayed two mating types (A1°and A1*) that were compatible with the previously identified Al biological species defined by the CI 1 isolate from the Canary Islands, which carried the Al'and Al’ mating type alleles. Both Physarum melleum isolates (Mex 1, SC 1) were nonheterothallic.

Morphology The morphology of the Didymium squamulosum isolates in this study conforms to the results of the earlier study (El Hage et al. 2000) in that they fall into two overlapping morphotypes: type 1 = fairly tall limy stalks and rugose peridial lime; type 2 = short limeless (in culture) stalks and fairly smooth thin peridial lime. Isolates Aust 1, Aust 6, CR 29, CR 30, and Ore 1 are type 1; isolates Aust 2, Aust 3, Aust 4, Aust 5, Aust 6, Aust 7, Aust 8. Aust 9, Aust 10, NZ 1, Thai 1, and Trin 1 are intermediate but with type | tendencies (low levels of lime on the stalk); isolates Mex 1, NZ 2, and CR 33 are intermediate but with type 2 tendencies; and isolates CR 31 CR 32, and CR 34 are type 2. The variations in the Physarum compressum isolates also matched those noted in earlier studies (Irawan et al. 2000), with some isolates displaying short stalks (CR 18 and Col 1), sporangial clustering (CR 3 and WV 1), and a sporotheca shape varying from the description (long-stalked sporangia with a single napiform sporotheca) provided in Martin and Alexopolous (1969). Sporothecae were mostly reniform in this group of isolates, with a few allantoid (Col 1, Ecu 3c, and Idn 3) or napiform (CR 3, Gua 1, and PR 4) isolates. The Guy 2 isolate, which was tentatively placed in this species, had very poorly developed sporangia and its morphology could not be determined adequately. The Physarum melleum isolates were typical of the morphospecies, with the exception of the Mex | isolate, which had yellowish-orange instead of the normal orange colored peridial lime deposits.

DISCUSSION

The study of these additional isolates supports and extends the results of previous papers (El Hage et al. 2000, Irawan et al. 2000, Clark and Stephenson 2000). Additional sympatric and allopatric biological species were added to the already intricate complex known to exist for Didymium squamulosum (El Hage et al. 2000). The recovery of one and the same biological species from Australia, New Zealand, and Thailand is very different from the situation in Central America, where there seem to be a number of small sympatric biological species. This morphospecies apparently consists of two genetically and morphologically overlapping complexes, each of which consists of several biological species and numerous apomictic lines. One of these complexes generally conforms to the central core of the D. sguamulosum description (Martin and Alexopoulos 1969), while the other resembles Didymium dictyopodium Nann.-Brem. & Yamam., as delimitated by Matsumoto (1999). Although these two complexes are not completely separated from each other, they are probably different enough to be considered distinct taxa from a practical point of view. However, the ten or so rare, recently described (see El Hage et al. 2000) species that differ from D. sgumulosum in one or two characters (e.g., spore ornamentation and stalk color or lime) are likely to be local variants unworthy of species rank unless they can verified by cultural studies.

The discovery of a second heterothallic isolate in the generally low genetically and morphologically variable Physarum compressum complex supports the conclusion that this morphospecies is a coherent taxon consisting ofa biological species core and numerous similar apomictic lines. However, the consistent occurrence of isolates with clustered sporangia indicates that the Physarum nicaraguense Macbr. morphospecies is not a supportable taxon. The Guy 2 isolate, which had been tentatively placed in the P. compressum complex, is probably related to Physarum gyrosum Rost., since it is very similar to the Guy | that was found to be a specimen of P. gyrosum on the basis of isozyme studies (Irawan et al. 2000).

The two new Physarum melleum isolates support the idea that this morphospecies is a complex of similar (peridial lime color is the major observed variation) apomictic lines. However, further studies may find a basal biological species, from which the lines are derived, a situation comparable to that found in P. compressum.

89

This paper, other studies on these three taxa (El Hage et al. 2000, Irawan et al. 2000, Clark and Stephenson 2000), and an examination of the literature (Clark 1995, Clark 2000) indicate that most morphospecies, examined to date, consist ofa core of sexually reproducing heterothallic strains and a swarm of nonheterothallic clonal lines. Also, the sexual strains of a species may belong to a single biological species (Haskins et al. 2000), or they may be divided into a number of reproductively isolated sibling species that may be allopatrically (Clark and Stephenson 1990) or sympatrically (El Hage et al. 2000) distributed. Together, these variations produce a complex set of reproductive possibilities that, when combined with morphological variations, produce extremely messy species complexes. Therefore, the delimitation of true taxa in this group, on the basis of morphology alone, is rather problematic, and the description of a new species should not be undertaken without extensive studies. Special care should be taken to avoid the naming of rare species that differ from a widespread taxon in one or only a few characters, since these will very likely turn out to be variant local apomictic lines not deserving of a separate taxonomic description.

ACKNOWLEDGMENTS

Weare grateful to Mr. B. Irawan, Ms. D. Black, and Drs. J. Landolt, M. Schnittler and N. McLetchie for providing material that produced some of the cultures used in these studies. This work was supported in part by a grant (DEB-9705464) from the National Science Foundation to SLS..

LITERATURE CITED

Clark J. 1995. Myxomycete reproductive systems: additional information. Mycologia 87: 779-786.

Clark J. 2000. The species problem in the Myxomycetes. Stapfia 73: 39-53.

Clark J, Stephenson SL. 1990. Didymium iridis reproductive systems: new additions. Mycologia 82: 274-276.

Clark J, Stephenson SL. 2000. Biosystematics of the myxomycete Physarum melleum. Nova Hedwigia 71: 161-164.

El Hage N, Little C, Clark J, Stephenson SL. 2000. Biosytematics of the Didymium squamulosum complex. Mycologia 92: 54-64.

Haskins E, McGuinness MD, Clark J. 2000. Heterothallic mating systems in the Echinosteliales II. Echinostelium coelocephalum. Mycologia 92: 1080-1084.

Irawan B, Clark J, Stephenson SL. 2000. Biosystematics of the Physarum compressum morphospecies. Mycologia 92: 884-893.

Martin GW, Alexopoulos CJ. 1969. The Myxomycetes. Univ. Iowa Press, Iowa City, lowa. 556 pp.

Matsumoto J. 1999. Taxonomic study of the genus Didymium (Physarales, Myxomycetes). Ph.D. Dissertation, Hiroshima University, Hiroshima, Japan.

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MYCOTAXON

Volume LXXXV, pp. 91-99 January-March 2003

NOTES ON PESTALOTIOPSIS FROM SOUTHERN CHINA

Jiaxiang Zhang Tong Xu" Qixin Ge

(Department of Plant Protection, College of Agriculture and Biotechnology, Zhejiang University, Hangzhou 310029, China )

ABSTRACT

More than 800 specimens of Pestalotiopsis and allied genera collected from southern China (Zhejiang, Guangxi, and Yunnan provinces) in 2000-2002 were examined. Among the fifty-five identified species of Pestalotiopsis, nine species formerly disposed in Pestalotia sensu Guba are transferred to Pestalotiopsis. They are Pestalotiopsis aloes, P. clusia, P. elasticola, P. eusora, P. laughtonae, P. macrochaeta, P. mangifolia, P. scirpina, and P. vismiae. In addition, ten species of Pestalotiopsis are new records in China.

Key words: new combinations; new records.

As part of our floral studies on Pestalotiopsis in China, more than 800 specimens of Pestalotiopsis and allied genera collected from southern China (Zhejiang, Guangxi, and Yunnan provinces) in 2000-2002 were examined and more than 150 isolates were obtained by single conidium culture method. The fifty-five Pestalotiopsis species and a few in allied genera including Bartalinia robillardoides Yassi, Monochaetia camelliae (Desm.) Allescher, Seiridium anceps (Saccardo) Sutton, S. ceratosporum (De Notaris) Nag Raj, S. breviaristatum (Tracy & Earle) Sutton, and S. mali (Ellis & Everhart) Nag Raj were identified according to the treatments of Steyaert (1949), Sutton (1980), and Nag Raj (1993). Among the identified species of Pestalotiopsis, nine species formerly disposed in Pestalotia sensu Guba (1961) are transferred to Pestalotiopsis and re-described. In addition, ten species of Pestalotiopsis are new records in China.

New Combinations of Pestalotiopsis 1. Pestalotiopsis aldes (Trinch.) J.X. Zhang et T. Xu, comb. nov. Fig. 1 F*3e4ait S Ef

Corresponding author, xutong@zju.edu.cn

92

= Pestalotia alées Trinch., Rend. Napoli Soc. Reale accad. Sc. Fisiche Maten. Napoli 48: 266, 1909.

Fruitbodies scattered, conidiomata acervuloid, subepidermal in origin and erumpent at maturity; conidia fusiform, 4-septate, usually without constriction at the septa, straight or slightly curved, 18.9~23.6 X5.0~7.1 um; three intermediate cells almost concolorous, dark brown, the lowest one slightly lighter, 11.8~15.3 um long; apical and basal cell conic, hyaline, with 2~3 appendages at apex, long-filiform, diverging from the acute apex, 13.0~26.0 tm long; basal appendage single, 2.4~5.9 um long.

Colony on PDA white, cottony, conidiomata aggregate, ink-like, reverse of the culture Ochraceous-Buff(Ridgway, 1912), with circular bands.

Specimens examined: On leaves of Persea americana Mill., (PSH2000 | -093), 13 XI 2000, Jinghong Yunnan (2 fa set); on leaves of Roystonea regia (HBK.) O.F. Cook, (PSH2000 I -094), 12 XI 2000, Jinghong, Yunnan.

According to Trinchieri’s original fig. 1 Conidia of Pestalotiopsis al6es (Trinch.) description, conidia on dry scapes of Aloe iXezhane een Xu virens How. are 19~25 X 5~7 wm and unusually narrow.

2. Pestalotiopsis clusiae (Griffon & Maubl.)J.X. Zhang et T. Xu, comb. nov. Fig. 2 FES Fe LAER & BP

= Pestalotia clusiae Griffon & Maubl., Bull. Soc. Mycol. Fr. 25: 239, 1909.

Spots large, of variable shape, gray white with gray brown margin. Fruitbodies punctiform, scattered, conidiomata acervuloid, subepidermal in origin and erumpent at maturity; conidia ovate-oblong often curved and _ unequilateral, 4-septate, usually without constriction at the septa, 18.9~24 X 7.7~10.0um, intermediate three cells almost concolorous, dark brown, the lowest one

slightly lighter, 13.0~16.5 tum, apical and basal cell conic, hyaline, with 2~3 appendages at apex,

Fig. 2 Conidia of Pestalotiopsis clusiae

(Griffon & Maubl.) J.X. Zhang et T. Xu

acrogenous or acropleurogenous, 18.9~28.3 um long, basal appendage single, 2.4~7.5 um long.

Colony on PDA white, cottony, fruitbodies ink-like, more or less gregarious, reverse of the culture Light Ochraceous-Salmon, without circular bands.

Specimen examined: On the leaves of Areca triandra Roxbg., (PSH2000 I -145), 12 XI 2000, Jinghong, Yunnan.

According to Griffon & Maublance’s original description, conidia on leaves of Clusia sp. are corpulent, 20~25 X 8~10 um and with long flexuous appendages.

93

3. Pestalotiopsis elasticola (P. Henn.) J.X. Zhang et T. Xu, comb. nov. Fig. 3 El Se PRI tk Ef

= Pestalotia elasticola P. Henn., Hedwigia 48: 16, 1908.

Spots pale, dried out with dark brown margin. Fruitbodies punctiform, Fig. 3 Conidia of Pestalotiopsis elasticola (P. scattered, conidiomata acervuloid, Henn.) J.X. Zhang et T, Xu subepidermal in origin and erumpent at maturity; conidia long-fusiform, tapering towards the base, slightly constricted at the septa, 4-septate, straight or slightly curved, 23.6~28.3 X5.9~8.3 um, three intermediate cells almost concolorous or versicolour, the upper two umber, the lower one olivaceous, 14.2~18.9 um long, apical cell conic, hyaline, basal cell acute, hyaline, with 2~3 appendages at apex, divergent, slender, 7.1~23.1 um long, basal appendage single, 1.2~5.9 um long.

Colony on PDA white, cottony, fruitbodies gregarious, ink-like, reverse of the culture colourless.

Specimen examined: On leaves of Phoenix roeberenii Brien, (PSH2000 I -109), 12 XI 2000, Jinghong, Yunnan.

According to Henning’s original description, conidia on leaves of Ficus elastica are 14~20 X 7~10 uum with appendages 10~18 pm long.

This species differs from Pestalotiopsis elasticae (Koord.) P.L. Zhu, Q.X. Ge et T. Xu described on the same host plant in appendages without knob at the end and wider conidia.

4. Pestalotiopsis eusora (Sacc.) J.X. Zhang et T. Xu, comb. nov. Fig.4 SAH 4 in & Bf

= Pestalotia funerea Desm. var. eusora Sacc., Ann. Mycol. 6: 568, 1908.

Fruitbodies scattered or gregarious, erumpent, black. Conidiomata acervuloid, conidia equilateral, narrow-fusiform, straight, 4-septate, usually

without constrictions at the septa, 20.1~23.6 x

Fig. 4 Conidia of Pestalotiopsis eusora (Sacc.) 5.2~7.1 um , three intermediate cells almost TX ohane et 1 XG

concolorous, the upper two brown to dark brown, the lower one olivaceous with melanin disposed at septa, 11.8~14.2 tum long, apical cell narrow-cylindric to acute, basal cell triangle, hyaline, with 2~3 appendages

at apex, acrogenous or acropleurogenous, 11.8~21.2 ttm long, basal appendage single, 3.5~7.1 um long.

Specimens examined: On leaves of Caryota mitis Loureiro, (PSH2000 | -050), 18 VIII 2000, Qingxiu Mountain, Guangxi (J” PUFF LL); on leaves of Roystonea regia (HBK.) O.F. Cook, (PSH2000 I -010),7 VIIT 2000, Nanning, Guangxi (J 75 Fd +); on leaves of Osmanthus pragrans (PSH2000 | -070), 27 VIII 2000, Guilin, Guangxi (J Pa#EAK).

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According to Guba’s description, conidia on withering leaves of Heptapeurum barteri Hiern. are 20~22 X 5. 5~6 um (Guba, 1961).

5. Pestalotiopsis laughtonae (Doidge) J.X. Zhang et T. Xu, comb. nov. Fig.5 3A tings 4 Efe

= Pestalotia laughtonae Doidge, Bothalia 4: 824, 1948.

Leaf spots irregular with dark brown margins. Fruitbodies punctiform, scattered or gregarious, conidiomata acervuloid, subepidermal then erumpent; conidia ovoid or broadly fusiform, 4-septate, usually not constricted at the septa, straight or unequilateral, 18.9~23.6 X 8.3~10.6 jum, three intermediate cells slightly swollen, versicolour, the upper two dark

brown with deposition of melanin at septa, the Fig. 5 Conidia of Pestalotiopsis laughtonae

(Doidge) J.X. Zhang et T. Xu

lower one brown, 13.0~16.5 um, apical cell cylindric or conic, basal cell turbinate, hyaline, with 2~3 appendages at apex, widely divergent, 15.3~28.3 um long, basal appendage single, 2.4~5.9 um long.

Colony on PDA white, cottony, fruitbodies aggregate, ink-like, with circular bands, reverse of the culture Mars Yellow.

Specimen examined: On leaves of Caryota ochlandra Hance, (PSH2000 | -126), 9 XI 2000, Menglun, Yunnan (zs Fae) .

According to the original description, conidia on leaves of Cassine sphaerophylla O. Kize are 2235225 X9~ 11, um.

Steyaert (1949) placed Pestalotia laughtonae in synonymy with Pestalotiopsis glandicola (Cast.) Stey. [= Pestalotia glandicola (Cast.) Guba]: conidia 21~28 X 8~11 um, three intermediate cells 16~20 um, with 2~4 appendages at apex, 10~37 um long. The later differs from Pestalotiopsis laughtonae in larger size of conidia and more appendages.

6. Pestalotiopsis macrochaeta (Speg.) J.X. Zhang et T. Xu, comb. nov. Fig.6 444] BW th & EF

= Pestalotia funerea Desm. var. macrochaeta Speg., Anal. Mus. Nac. Buenos Aires, Ser. III, 13: 412, 1911.

= Pestalotia macrochaeta (Speg.) Guba, Mycologia 24: 369, 1932.

Fruitbodies punctiform, scattered, conidiomata acervuloid, subepidermal in origin and erumpent at maturity; conidia long-fusiform, tapering towards the base, 4-septate, usually without constriction at the septa, straight, 24.8~33.0 X 6.6~7.6 um, three intermediate cells almost concolorous, upper two gray brown, light on the lowest one, 13.0~21.2 um, apical cell conic to cylindric, and basal cell conic, hyaline, with 2~3 (most 3) appendages at apex, acrogenous or acropleurogenous, 15.3~34.2 um long, basal appendage single, longer, 7.1~11.8 um long.

Colony on PDA white, cottony, fructifications slowly, fruitbodies scattered, ink-like, reverse of the culture Cinnamon-Rufous without circular bands.

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Specimen examined: On leaves of Ravenala madagascariensis Adans., (PSH2000 I -107), 10 XI 2000, Menglun, Yunnan. According to Spegazzini’s original description, conidia on decaying needles of Pinus sylvertris L. are 28~30 X 8~9 um with appendages, 30 um long.

7. Pestalotiopsis mangifolia (Guba) J.X. Zhang et T. Xu, comb. nov. Fig. 7 Tee itt & Bf

= Pestalotia mangifolia Guba, Monograph of Monochaetia & Pestalotia p. 149, 1961.

Fruitbodies punctiform, gregarious or scattered, conidiomata acervuloid, subepidermal then erumpent; conidia fusiform or long-cylindric, 4-septate, slightly constricted at the septa, straight or slightly curved, 23.6~29.5 X 5.9~9.0 tm, three intermediate cells concolorous, upper two gray brown or olivaceous, light on the lowest one, 14.2~20.1 um long, apical cell cylindric, and basal cell conic, hyaline, with 2~3 (mostly 3) appendages at apex, acrogenous or

Joan,

acropleurogenous, 20.1~28.3 um, basal appendage single, 4.2~8.3 um.

Fig. 6 Conidia of Pestalotiopsis macrochaeta

(Speg.) J.X. Zhang et T. Xu

Colony on PDA white to yellowish, \ cottony, fruitbodies scattered, ink-like, | reverse of the culture Capucine Buff, | without circular bands.

Specimens examined: On leaves of Bixa orellana L., (PSH2000 [ -122), 9 XI 2000, Menglun, Yunnan; on leaves of Mayodendron igneum (Kurz.) Kurz,

(PSH2000 I -103), 10 XI 2000, Menglun, Fig. 7 Conidia of Pestalotiopsis mangifolia (Guba) J.X. Zhang et T. Xu

Yunnan; on leaves of Crataegus pinnatifida Bge., (PSH2000 I -123), 9 XI 2000, Menglun, Yunnan. According to Guba’s original description, conidia on dead brown areas in leaves of Magnifera indica L. are 22~28 X 6~8.5 tum and with 3 appendages 8~26 tm long. This species differs from Pestalotiopsis magniferae P. Henn. broader conidia and longer appendages.

8. Pestalotiopsis scirpina (Ell. & G. Martin) J.X. Zhang et T. Xu, comb. nov. Fig. 8 FR BLAU tt 2 Ef

= Pestalotia scirpina Ell. & G. Martin, Am. Naturalist 19: 76, 1885. = Pestalotia versicolor Speg. var. americana Speg., Anal. Soc. Ci. Argent. 13: 21, 1882.

96

= Pestalotia linearis Sacc., Nuovo Gior. Bot. Ital. (n .s.) 22: 67-68, 1915.

Fruitbodies punctiform, scattered, conidiomata acervuloid, subepidermal then erumpent; conidia clavate or broadly fusiform, 4-septate, constricted at the septa, usually curved and unequilateral, 20.1~26.0 X 9.0~11.8 jum, three intermediate cells versicolor, the upper two cells dark brown, the lower one brown, 15.3~18.9(~16.9) um long, apical cell conic to columnar, and basal cell conic, hyaline, with 2~3 apical appendages, acrogenous or acropleurogenous, 22.4~34.2 um long, basal appendage single, 2.8~5.9 um long.

Colony on PDA white, cottony, fruitbodies scattered, ink-like, reverse of the culture Apricot Buff, without circular bands. Fig .8 Conidia of Pestalotiopsis scirpina (Ell. &

Specimen examined: On leaves of Gi Marini) xoZhaneeey, Xu Myristica yunnanensis Y.H. Li, (PSH2000 I -108), 11 XI 2000, Menglun, Yunnan.

According to Ellis’ description, conidia on decaying culms of Scirpus maritimus L. are 20~ 25 X 7~ 9 um.

This species is different from Pestalotiopsis versicolor (Speg.) Stey. in the broader conidia.

9. Pestalotiopsis vismiae (Petr.) J.X. Zhang et T. Xu, comb. nov. Fig.9 45 a] fi

= Pestalotia vismiae Petr., Sydowia 4: 561, 1950.

Spots irregular. Fruitbodies punctiform, loosely distributed, conidiomata acervuloid, subepidermal then erumpent; conidia long-fusiform, tapered towards ends, 4-septate, slightly constricted at the septa, straight or weakly curved, 20.1~23.6 X 5.2~6.6 wm, three intermediate cells almost concolorous, the upper two slightly brown, the lower one yellowish brown, 11.8~15.3 um _ long, apical cell conic to columnar, and basal cell conic, hyaline, with 2~3 (mostly 2) apical appendages, acrogenous or acropleurogenous, 9.4~17.7 um long, basal appendage single, 2.8~5.2 um long.

Colony on PDA white, cottony, fruitbodies scattered, ink-like, reverse of the culture Pale Orange- Yellow, without circular bands.

Specimens examined: On leaves of Aucuba chinensis Benth, (PSH2000 I -063), 27 VIN 2000, Guilin, Guangxi; on leaves of Cycas micholitzii Dyer, (PSH2000 | -039), 18 VHI 2000, Qingxiu Mountain, Guangxi; on leaves of Phoenix canariensis Chabaud, (PSH2000 I -044), 18 VIII 2000, Qingxiu Mountain, Guangxi; on leaves of

Fig. 9 Conidia of Pestalotiopsis vismiae (Petr.) J.X. Zhang et T. Xu

i,

Quercus variabilis Bl., (PSH2000 I -138), 7 XI 2000, Kunming Botanical Garden, Yunnan (Lé 844 bil); on needles of Cedrus deodara (Roxb.) Loud., (PSH2000 | -154), 6 XI 2000, Cuihu park Kunming, Yunnan (522 3|2ZS bal); on leaves of Amygdalus persica L., (PSH2000 I -081), 29 X 2000, Hangzhou, jiuxi, Zhejiang (Hf YN FUR).

According to the original description, conidia on living leaves of Visimia obtuse Spruce are 18~22 X 5~6.5 um and with 2 appendages 10~23 um long.

New records of Pestalotiopsis from China 1. Pestalotiopsis calabae (West.) Stey., Bull. Jard. Bot. Etat Bruxelles 19: 308, 1949. sf A OL EH

= Pestalotia calabae West., Bull. Acad. Roy. Belg., Bruxelles, Ser. [I], 2: 556, 1857.

On leaves of Dracontomelon duperreanum Pierre, (PSH2000 [| -006), 7 VIII 2000, Guilin, Guangxi; on leaves of Acacia confusa Merr., (PSH2000 | -009), 7 VII 2000, Nanlin, Guangxi, on leaves of Dimocarpus longan Lour, (PSH2000 | -029), 16 VIII 2000, Guangxi University campus.

2. Pestalotiopsis ixorae (Rangel) Batista & Peres apud Batista et al., Atas Inst. Micol. 3:99, 1966. FEMTTEU & Efe

= Pestalotia ixorae Rangel, Bol. Agr., Sao Poulo, 16: 321, 1915.

On leaves of Ixora chinensis Lam., (PSH2000 I -140), 14 XI 2000, Jinghong, Yunnan.

3. Pestalotiopsis leprogena (Speg.) Steyaert, Darwiniana 12(2): 159, 1961. 'RYF at ABW tt & Efe |

= Pestalotia leprogena Speg., An. Mus. Nac. Hist. Nat. Buenos Aires 23: 109, 1912; Syl. Fung. 25: 604, 1931. =Pestalotia rhipsalidis Grillo, Inst. Biol. Veg., Rio de Janeiro 1: 63-65, 1934.

On leaves of Cycas revoluta, (PSH2000 [ -038), 18 VIII 2000, Qingxiu Mountain, Guangxi.

4. Pestalotiopsis maculiformans (Guba & Zeller) Stey., Bull. Jard. Bot. Etat Bruxelles 19: 329, 1949. KKH th & Bi

= Pestalotia maculiformans Guba & Zeller, Mycologia 24: 370, 1932.

On leaves of Taxus chinensis (Pilg.) Rehd., (PSH2000 I -137), 2000.11.7. Kunming Botanical Garden, Yunnan. 5. Pestalotiopsis menezesiana (Bres. & Torr.)Bissett, Can. J. Bot. 60: 2570, 1982. PEAS tik & BFE

= Pestalotia menezesiana Bres. &Torr., Broteria ser. Bot, 8: 142, 1909.

On leaves of Ravenala madagascariensis Adans, (PSH2000 I -111), 2000.11.12. Jinghong, Yunnan; on leaves of Rhododendron stamineum Franch., (PSH2000 I 072),

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27 VI 2000, Guilin, Guangxi; on leaves of Ravenala madagascariensis Adans, (PSH2000 I -113), 9 XI 2000, Menglun, Yunnan.

6. Pestalotiopsis oxyanthi (Thuem.) Stey., Bull. Jard. Bot. Etat Bruxelles 19: 329, 1949. Rete ae s =B fl

= Pestalotia oxyanthi Thuem. Inst. Rev. Sci. Litt. Coimbra 28: 420, 1880.

On leaves of Zingiber officinale Rosc., (PSH2000 I -139), 10 XI 2000, Menglun, Yunnan.

7. Pestalotiopsis populi-nigrae (Sawada & Ito) Morelet, Bull. Sci. Nat. Archeol. Toulon Var. 205: 9 1973. $i Kimi s Bry

= Pestalotia populi-nigrae Sawada & Ito, Rep. For. Exp. Sta. Tokyo 45: 135, 1950. =Pestalotia lignorum Gambogi. Publ. Inst. Pat. Veg. Microbiol. Agr. Univ. Pisa no. 3322-15, 19506.

On leaves of Homalium luoticum, (PSH2000 I -133), 9 XI 2000, Menglun, Yunnan.

8. Pestalotiopsis sonsensis (P. Henn.) Stey., Bull. Jard. Bot Etat Bruxelles 19: 336, 1949. Se pRUih & Efe |

= Pestalotia sonsensis P. Henn., Ann. Mus. du Congo, Bot., Ser. 5, 2(3): 229, 1908.

On leaves of Mesua ferrea L., (PSH2000 I -127), 9 XI 2000, Menglun, Yunnan.

9. Pestalotiopsis triseta Stey., Bull. Jard. Bot. Etat Bruxelles 19: 337-338, 1949, = EWU hs 2 ETA

= Pestalotia funerea Desm. var. macrochaeta Speg. f. triseta, Rev. Mycol. 6: 49, 1941.

= Pestalotia triseta Guba, Monograph of Monochaetia and Pestalotia p. 244, 1961.

On leaves of Caryota mitis Cour., (PSH2000 I -088), 10 XI 2000, Menglun, Yunnan. |

10. Pestalotiopsis uvicola (Speg.) Bissett., Can. J. Bot. 60: 2572, 1982. Js Ayes 2 Bf = Pestalotia uvicola Speg., in Thuem., Pilze Weinst, Vienna p. 13, 1878. On leaves of Mahonia acanthifolia Wall. ex G. Don, (PSH2000 I -001), 7 VIII 2000, Nanning, Guangxi ()” 4 fd 3*) ; on leaves of Cycas revoluta Thunb., (PSH2000 I -035), 8 VIII 2000, Qingxiu Mountain, Guangxi; on leaves of Macadamia integrifolia Maiden, (PSH2000 I -002), 7 VIII 2000, Nanning, Guangxi.

ACKNOWLEDGEMENTS The authors would like to thank Dr. W. Y. Zhuang for pre-submission review. Kind help of Prof. Y. X Chen and Prof. G. Wei of Guangxi University to collect

99

specimens is also appreciated. This project was supported by National Natural Science Foundation of China.

LITERATURE CITED Guba, E. F. 1961. Monograph of Monochaetia and Pestalotia. Harvard Univ. Press. Cambridge, Mass., U.S.A. 342 pp

Nag Raj. T. R. 1993. Coelomycetous Anamorphs with Appendage-bearing Conidia. Mycol. Publ. Waterloo, Ontario, Canada. 1101 pp

Robert Ridgways 1912. Color Standards and Color Nomenclature. United States National Museum, Washington D. C.

Steyaert, R. L. 1949. Contribution a l’etude monographique Pestalotia de Not. et Monochaetia Sacc. (Truncatella gen. nov. et Pestalotiopsis gen. nov.). Bull. Jard. Bot. Bruxelles 19: 285

Sutton, B. C. 1980. The Coelomycetes. Fungi Imperfecti with Pycnidia, Acervuli and Stromata. Comm. Mycol. Inst. England. 696 pp

Zhu, P. L., Q.X. Ge, and T. Xu. 1991. Seven new combination of Pestalotiopsis from China. Acta Mycol. Sinica 9: 273-279

A ER] ps 7 TU tt & =E fl J Pestalotiopsis WMS

KAR FR lel* EC HH WILKS RV SEWRAL GRA =A 310029)

fi: 2000 7% 2002 *FHIIA), WYRE. HR, BSH KES 800 SIA She (25 ERA i 49 NBL), FRYE Steyaert. Sutton All Nag Raj 4} 2A, ene J Wik & Bf) (Pestalotiopsis) 55 SF, HH ARO SAA, 11 SEA Pritt. PHZAAA Pestalotiopsis aldes. P. clusiae, P. elasticola., P. eusora.,

P. laughtonae, P. macrochaeta, P. mangifolia, P. scirpina #\ P. vismiae.. ABE]: PAG Erie

* WHE, xutong@zju.edu.cn

ao

MY COTAXON

Volume LXXXV, pp. 101-108 January-March 2003

POLYPORE FUNGI NEWLY RECORDED FROM TAIWAN Sheng-Hua Wu! and Leif Ryvarden’

‘Department of Botany, National Museum of Natural Science, Taichung, Taiwan 404, Republic of China * Department of Botany, University of Oslo, P.O. Box 1045, N-0316 Oslo, Norway

ABSTRACT

Twelve species of polypore fungi are newly recorded from Taiwan, viz., Abundisporus roseoalbus, Antrodiella hydrophila, Ceriporiopsis lowei, Echinochaete brachyporus, Hexagonia hirta, Megasporoporia cavernulosa, Oxyporus corticola, Perenniporia isabellina, Polyporus dictyopus, Skeletocutis sensitiva, Theleporus calcicolor and Trametes pocas. Cultural studies are provided for EF. brachyporus, H. hirta,

Theleporus calcicolor, and Trametes pocas.

Key words: Aphyllophorales, Basidiomycota, cultural studies, new records, Taxonomy.

Specimens of twelve polypore species newly recorded from Taiwan were collected from subtropical and tropical Taiwan during 1993-2001. All specimens are deposited in the herbarium of the National Museum of Natural Science of ROC (TNM). Some duplicates have been distributed to the herbarium of the University of Oslo (O). The cultural description and species code system are basically from Nobles (1965) with amendments by Boidin and Lanquetin (1983). Nobles' code as detailed by Nakasone (1990) was adopted in this study. General methods of cultural studies have been previously described (Wu, 1996).

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1. Abundisporus roseoalbus (Jungh.) Ryvarden Belg. Journ. Bot. 131: 154. 1998. Specimens examined. Taiwan. Taipei: Yangminshan National Park, Luchiaokenghsi Protected Area, on rotten wood of angiosperm, 350 m, coll. S.H. Wu et al., 21-X-2000, Wu 0008-9 (TNM F12759). Taitung: Orchid Island, Hungtoushan, alt. 450 m, on rotten trunk of angiosperm, coll. $.Z. Chen, 24-IV-1997, Chen 665 (TNM F8831, O). Distribution. Paleotropical (Ryvarden and Johansen, 1980).

2. Antrodiella hydrophila (Berk. & Curt.) Ryvarden, Mycotaxon 20: 343. 1984. Specimen examined. Taiwan. Taitung: Chihpen Forest Recreation Area, alt. 150 m, on wood, coll. S.H. Wu et al., 25-VI-1999, Wu 9906-17 (TNM F10549).

Distribution. Central America, Africa (Ryvarden and Johansen, 1980), Taiwan.

3. Ceriporiopsis lowei Rajchenberg, Nordic J. Bot. 7: 564. 1987.

Specimens examined. Taiwan. Taitung: Orchid Island, between nuclear waste storage and Tasenshan, alt. 100 m, on branch of angiosperm, coll. SH. Wu & J.Y. Tseng, 29-IV-1997, Wu 9704-55 (TNM F8628, O), Wu 9704-75 (TNM F8646, O). Distribution. Australia, New Zealand (Rajchenberg, 1992), Brazil (type locality),

Taiwan.

4. Echinochaete brachyporus (Mont.) Ryvarden, Bull. Jard. Bot. Nat. Belg. 48: 101. 197s!

Specimen examined. Taiwan. Taitung: Taimali, Chinjenshan, alt. 500 m, on rotten trunk of angiosperm, coll. S.C. Wu, 7-[V-2001, Wu 0104-8 (TNM F12760). Distribution. Pantropical (Nufiez and Ryvarden, 1995).

Cultural description (PS-Mycelium of Wu 0104-8). Growth rate of 1-6 wk: Colony radius respectively 5-10, 16-23, 24-31, 31-42, 35-63, and 42-72 mm. 1 wk growth: Mat white, slightly brownish tinted. Advancing zone fairly even. Aerial mycelium slightly pellicular. 2 wk growth: Mat whitish, but inoculum and zonate area brown. Advancing zone uneven. Aerial mycelium absent or slightly pellicular, slightly irregularly zonate. 6 wk growth: Mat generally brownish tinted. Advancing zone uneven. Aerial mycelium absent or slightly pellicular, occasionally with slightly crustose patches, slightly irregularly zonate. Agar unchanged. No distinct odor. Not fruiting. Hyphal system monomitic. Advancing hyphae colorless, 2-4.5 um diam., thin-walled, nodose-septate. Other hyphae colorless or brownish, 2-5 pm diam., thin-walled, simple- or nodose-septate, moderately branched, occasionally with brown oily contents. Setoid elements present (Fig. 1A). Thick-walled hyphal ends (Fig. 1B) present in crustose patches. Crystals present.

Oxidase reactions. GAA: —, 0; -, 0. TAA: —, 0; +, 5-8. TyA: +, 0; +, tr.

103 A B Fig. 1. Cultural features of Echinochaete brachyporus (Wu 0104-8). A. Setoid

elements. B. Thick-walled hyphal ends from crustose patches. Culture after 6 wk of growth on 1.5% MEA at 25'C. Scale bar=10 wm.

Nuclear behavior. Normal. Spores generally uninucleate, rarely binucleate, MS-mycelium monokaryotic, PS-mycelium dikaryotic.

Sexuality. Heterothallic tetrapolar (AiBi: 1; AiB2: 5, 8; A2Bi1: 3, 6, 7, 9; A2Bz: 2, 4). Species code. 2, 31, 13, 32, 37, 38, 47, 54, 60, 61.

5. Hexagonia hirta (Fr.) Fr., Epicr. Syst. Mycol.: 497. 1838.

Specimen examined. Taiwan. Taitung: Taimali, alt. 100 m, on branch of angiosperm, coll. S.H. Wu et al., 27-V1-1999, Wu 9906-35 (TNM F10567).

Distribution. Africa (Ryvarden and Johansen, 1980), the Philippines (Ryvarden, 1989), Taiwan.

Cultural description (FB-mycelium of Wu 9906-35). Growth rate of 1-4 wk: Colony radius respectively 13-16, 32-40, 52-67, (90+) mm. 1 wk growth: Mat white. Advancing zone even. Aerial mycelium slightly pellicular. 2 wk growth: Mat white. Advancing zone even. Aerial mycelium slightly pellicular. 6 wk growth: Mat white to brown at pellicular region, brown at crustose region. Aerial mycelium generally pellicular, occasionally crustose. Agar turning brown. No distinct odor. Not fruiting. Hyphal system dimitic. Advancing hyphae colorless, 2-5 um diam., thin-walled, nodose-septate. Aerial skeletal hyphae colorless or yellow, branched, 1-3 um diam.,

slightly thick-walled, with distinct lumen. Aerial generative hyphae colorless to

104

brown, 1.5-3 pm diam., thin-walled, nodose-septate, moderately branched. Submerged hyphae colorless, 1.5-4 gm diam., thin-walled, nodose-septate, _ moderately branched. Crystals present.

Oxidase reactions. GAA: +++, 0; +++, 0. TAA: +++, 0; +++, 0. TYA: —; 21-28;--, 65-66.

Species code. 2a, 3c, 8, 32, 37, 39, 45, 54.

Remarks. Three species of Hexagonia bear long (> 7 mm long), branched, dark pileus hairs (Ryvarden and Johansen, 1980). Hexagonia hirta differs from H. apiaria in having smaller pores on the hymenial surface, and is distinct from H. hydnoides in

having larger pores.

6. Megasporoporia cavernulosa (Berk.) Ryvarden, Mycotaxon 16: 174. 1982. Specimens examined. Taiwan. Taitung: Orchid Island, between nuclear waste storage and Tasenshan, alt. 100 m, on branch of angiosperm, coll. S.H. Wu, 29-IV-1997, Wu 9704-48 (TINM F8622). Green Island, Sleeping Beauty Rock, alt. 150 m, on branch of angiosperm, coll. S.Z. Chen, 9-IV-1998, Chen 783 (TNM F8884), Chen 784 (TNM F8885).

Distribution. Widespread in tropical America and Africa (Gilbertson and Ryvarden, 1987), Taiwan.

7. Oxyporus corticola (Fr.) Ryvarden, Persoonia 7: 19. 1972.

Specimen examined. Taiwan.Taichung: Kukuan, Malin, alt. 660 m, on bark of angiosperm, coll. C.Y. Chien, 8-VII-1996, Wu 9607-1 (TNM F5336, O).

Distribution. Circumglobal (Gilbertson and Ryvarden, 1987).

8.