By Meike Piepenbring, Botanisches Institut, J. W. Goethe-Universität Frankfurt,

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Meike Piepenbring - 1 - KEY FOR SMUT FUNGI KNOWN IN COSTA RICA BASED ON FAMILIES OF THE HOST PLANTS By Meike Piepenbring, Botanisches Institut, J. W. Goethe-Universität Frankfurt, Senckenberganlage 31-33, 60054 Frankfurt/Main, Germany E-mail: piepenbring@em.uni-frankfurt.de homepage: http://www.uni-frankfurt.de/fb15/botanik/piepenbring.html On Amaranthaceae: Sori as galls on different organs; spores in balls. On Iresine diffusa Humb. & Bonpl. ex Willd. Thecaphora haumani Speg. On Asteraceae: 1 Sori as galls on stems and leaves, spore balls. 2 Sori as galls mainly in the axils of leaves. On Smallanthus quichensis (Coult.) H. Rob. Thecaphora smallanthi M. Piepenbr., Paul Hanson & J. Carranza in Piepenbring 2 Sori on stems and leaves. On Bidens pilosa L. Thecaphora pustulata G. P. Clinton in Chardón 1 Sori on leaves, simple spores embedded in host tissue. 3 Sori as pustules protruding from the lamina, spores 13-16 μm. On Galinsoga urticaefolia (H.B.K.) Benth.... Entyloma galinsogae Syd. & P. Syd. 3 Sori as leaf spots. On Acmella oppositifolia (Lam.) R. K. Jansen var. oppositifolia Entyloma spilanthis Speg. (syn. Entyloma ecuadorense Syd.) 1

Meike Piepenbring - 2 - On Bidens pilosa L.... Entyloma bidentis Henn. in Engler On Dahlia imperialis Roezl and/or D. sp. cult.: 4 Spots mostly up to 3 mm diam., sharply delimited, spores 8-12 μm, walls up to 1.5 μm thick. Entyloma doebbeleri M. Piepenbr. 4 Spots ca. 10 mm diam., spores 13-16 μm, walls up to 3.5 μm thick. Entyloma dahliae Syd. & P. Syd. On Delilea biflora (DC.) Kuntze... Entyloma delileae Vánky, Döbbeler & U. Brown in Vánky On Tithonia rotundifolia (Mill.) Blake... Entyloma compositarum Farl. On Viguiera cordata (Hook. & Arn.) D'Arcy, V. sylvatica Klatt. Entyloma compositarum Farl. On Cyperaceae: 1 Spore mass orange; spores smooth, developing inside host tissue. On Scleria melaleuca Reichb. ex Schlechtend. & Cham. Aurantiosporium subnitens (J. Schröter & Henn.) M. Piepenbr., Vánky & Oberw. 1 Spore mass brown or black. 2 Spore masses around filaments of stamina; spores in balls. On Rhynchospora cephalotes (L.) Vahl... Moreaua rhynchosporae-cephalotis (Vánky & T. Vánky) Vánky 2 Spores not in balls. 3 Sori permeated by bundles of sterile, multicellular hyphae (elaters). 4 Sori long cylindrical, with thick, brown, smooth peridia. On Carex chordalis Liebm., C. jamesonii Boott, C. lemanniana Boott... Farysia corniculata Vánky 4 Sori globose, with thin peridia. 2

Meike Piepenbring - 3-5 Spores of irregular shape, ca. 6-8 μm, covered by warts often forming rows. On Carex polystachya Sw. ex Wahlenb.... Farysia chardoniana Zundel 5 Spores mostly spherical, ca. 9-12 μm, covered by warts not forming rows. On Carex polystachya Sw. ex Wahlenb.... Farysia venezuelana Zundel 3 Sori not permeated by elaters with similar celular structure. 6 Sori without peridia; spores around rudimentary host tissue at the tips of rachillae of spikelets; spores flattened. 7 Sori occur in all the spikelets of an infected plant. On Rhynchospora globosa (Kunth) Roem. & Schult.... Ustanciosporium neomontagnei M. Piepenbr. & Begerow in Piepenbring 7 Sori in groups of spikelets forming witches brooms. On Rhynchospora marisculus Lindl. & Nees... Ustanciosporium standleyanum (Zundel) M. Piepenbr. 6 Sori with peridia. 8 Sori replacing flowers, with sack-shaped peridia and unseptate, long sterile cells in the bases of the sori. On Rhynchospora corymbosa (L.) Britton [R. aurea Vahl], R. triflora Vahl Trichocintractia utriculicola (Henn.) M. Piepenbr. 8 Sori without sack-shaped peridia. 9 Spore mass around ovaries, without sterile stroma on the surface of the host tissue. 10 Spores rather light coloured, with lighter coloured and collapsed areas, ca. 15-17 μm. On Carex bonplandii Kunth Anthracoidea uleana (Syd. & P. Syd.) Vánky (syn. Anthracoidea pannucea (Liro) Zambett.) 3

Meike Piepenbring - 4-10 Spores rather dark coloured, without conspicuous lighter coloured areas, ca. 16-18 μm. On Carex chordalis Liebm., C. jamesonii Boott [C. cortesii Liebm.], C. lemanniana Boott Anthracoidea altiphila Vánky & M. Piepenbr. in Vánky 9 Spore mass with sterile stroma formed by sterile fungal hyphae on the surface of the host tissue, around ovaries or other organs. 11 Sori around internodes of stems or inflorescences, with conspicuous shining white peridia when young; spores irregularly roughly warty with ridges. 12 Sori around internodes of stems or inflorescences; spores ca. 15-18 μm. On Rhynchospora holoschoenoides (Rich.) Herter Leucocintractia leucodermoides M. Piepenbr. & Begerow in Piepenbring 12 Sori around all the internodes of inflorescences, not around internodes of the stem; spores ca. 12-16 μm. On Rhynchospora corymbosa (L.) Britton [R. aurea Vahl] Leucocintractia scleriae (DC.) M. Piepenbr., Begerow & Oberw. 11 Sori around the bases of internodes of stems or in inflorescences, with less conspicuous peridia; spores smooth to finely warty. 13 Sori only around ovaries of some spikelets of an infected plant; spores ca. 10-12 μm. On Fimbristylis spadicea (L.) Vahl... Cintractia fimbristylicola Pavgi & Mundk. 13 Sori usually around bases of peduncles, rarely in spikelets. 14 Spores ca. 11-12 μm. On Cyperus odoratus L. [Torulinium odoratum (L.) Hooper], C. rotundus L.... Cintractia limitata G. P. Clinton 14 Spores ca. 14-17 μm. On Fimbristylis dichotoma (L.) Vahl [F. annua (All.) Roem. & Schult.]... Cintractia axicola (Berk.) Cornu (syn. Cintractia peribebuyensis (Speg.) Sawada) 4

Meike Piepenbring - 5 - On Fabaceae: Sori as dark spots of leaves; spores dark, embedded in host tissue. On Phaseolus vulgaris L. Erratomyces patelii (Pavgi & Thirum.) M. Piepenbr. & R. Bauer On Juncaceae: Sori as galls of roots; spores inside hypertrophied host cells. 1 Spores spherical, ca. 11-23, covered by large, blunt warts. On Juncus tenuis Willd. Entorrhiza casparyana (Magnus) Lagerh. 1 Spores ovoid to subglobose, ca. 15-22 μm, covered by warts of medium size. On Juncus bufonius L.... Entorrhiza aschersoniana (Magnus) Lagerh. On Limnocharitaceae: Sori as swollen spots of leaves; spores in balls. On Limnocharis flava (L.) Buchenau Doassansiopsis limnocharidis (Cif.) Vánky On Lobeliaceae: Sori as leaf spots; spores embedded in host tissue. 1 Spores ca. 10-13 μm. On Diastatea micrantha (H.B.K.) McVaugh Entyloma diastateae M. Piepenbr. 1 Spores ca. 12-15 μm. On Lobelia laxiflora H.B.K.... Entyloma lobeliae Farl. On Nymphaeaceae: Sori as leaf spots. 5

Meike Piepenbring - 6-1 Spores single, attached to hyphae. On Nymphaea ampla (Salisb.) DC. Rhamphospora nymphaeae G. Cunn. 1 Spores in complex balls. On Nymphaea blanda G. F. W. Mey. Doassansiopsis ticonis M. Piepenbr. On Poaceae: 1 Sori as stripes in leaves, rarely in inflorescences. 2 Spore mass brown, powdery at maturity; spores ca. 11-12 μm. On Holcus lanatus L. Ustilago striiformis (Westend.) Niessl 2 Spore mass grey, agglutinated; spores ca. 7-8 μm. On Oryza sativa L. Eballistra oryzae (Syd. & P. Syd.) R. Bauer, Begerow, A. Nagler & Oberw. 1 Sori mostly in inflorescences or replacing them, rarely in leaves. 3 Sori as apically whip-shaped stems replacing inflorescences which are not developed; spores ca. 6-7 μm. On Saccharum officinarum L. Sporisorium scitamineum (Syd.) M. Piepenbr., M. Stoll & Oberw. 3 Sori not as apically whip-shaped stems. 4 On species of Sorghum. 5 Sori replacing more or less large parts of the inflorecences, usually not in single spikelets; sori with conspicuous columellae; spores ca. 11-14 μm. On Sorghum bicolor (L.) Moench. Sporisorium reilianum (J. G. Kühn) Langdon & Full. 5 Sori in spikelets. 6

Meike Piepenbring - 7-6 Sori in all the spikelets of an infected inflorescence; one sorus with a thin, fragile peridium and a small columella; spores ca. 7-8 μm. On Sorghum bicolor (L.) Moench. Sporisorium cruentum (J. G. Kühn) Vánky 6 Sori in all the spikelets of an infected inflorescence or in groups of spikelets; sori with thick, persistent peridia; spores ca. 6-7 μm. On Sorghum bicolor (L.) Moench. Sporisorium sorghi Ehrenb. ex Link, in Linné 4 Not on species of Sorghum. 7 Sori destroying the entire inflorescence, all the partical inflorecences or all the spikelets of an infected inflorescence; on an infected plant usually no healthy spikelets are developed. 8 Sori replacing partial inflorecences, with conspicuous columellae; spores ca. 17-19 μm. On Andropogon bicornis L., A. glomeratus (Walter) Britton, Stern. & Poggenb. Sporisorium culmiperdum (J. Schröter) Vánky 8 Sori destroying spikelets, without conspicuous columellae. 9 Spores ca. 15-19 μm. On Paspalum conjugatum Bergius, P. costaricense Mez, P. paniculatum L., P. plicatulum Michx., P. virgatum L.... Ustilago schroeteriana Henn. 9 Spores ca. 6-9 μm. On Stenotaphrum secundatum (Walt.) Kuntze Ustilago affinis Ell. & Everh. in Cockerell 7 At least some spikelets of the infected plant develop normally. 10 Sori in groups of densely situated spikelets (witches brooms), with conspicuous columellae; spores ca. 12-14 μm. On Andropogon bicornis L. Sporisorium holwayi (G. P. Clinton & Zundel) Vánky 10 Sori not in dense groups of spikelets (witches brooms). 7

Meike Piepenbring - 8-11 Sori as large parenchymatous galls (several cm diam.) in different, mostly generative organs; spores ca. 10-12 μm. On Zea mays L. (cult.)... Ustilago maydis (DC.) Corda 11 Sori smaller; not this combination of characteristics. 12 Spores ca. 19-23 μm; sori in ovaries. On Oplismenus burmannii (Retz.) P. Beauv. Tilletia oplismeni-cristati Pat. & Har. in Durán & Fischer 12 Spores smaller. 13 Sori in swollen ovaries; spores ca. 14-15 μm, mixed with balls of y -shaped conidia. On Panicum maximum Jacq., Setaria paniculifera (Steud.) Fourn. Tilletia ayresii Berk. in Massee 13 Spores mostly smaller, not mixed with balls of conidia. 14 Sori in rachillae of spikelets, inconspicuous, with columellae; spores ca. 9-13 μm. On Melinis minutiflora P. Beauv.... Sporisorium melinis (Zundel) Vánky 14 Sori as green galls formed by swollen floral organs, with very short to lacking columellae. 15 Sori in swollen ovaries; spores covered by few very large warts, spores ca. 10-13 μm. On Dichanthelium viscidellum (Scribn.) F. W. Gould Sporisorium veracruzianum (Zundel & Dunlap) M. Piepenbr. 15 Sori in swollen ovaries, stamina, or lodiculae; spores finely warty. 16 Spores ca. 10-11 μm. On Ixophorus unisetus (Presl) Schlechtend. Ustilago ixophori R. Durán 16 Spores ca. 7-10 μm. On Echinochloa colona (L.) Link Ustilago trichophora (Link) Körn. On Polygonaceae: 8

Meike Piepenbring - 9-1 Sori as galls on different organs (leaves, stems, inflorescences); spores in groups embedded in a slimy matrix in host tissue, ca. 11-13 μm. On Polygonum hispidum H.B.K. Melanopsichium pennsylvanicum Hirschh. 1 Sori only in flowers. 2 Sori with peridia and columellae mostly formed by sterile fungal cells; spores, when young, in chains, ca. 11-14 μm, warty. On Polygonum acuminatum H.B.K., P. punctatum Ell. Sphacelotheca cf. koordersiana (Bref.) Zundel 2 Sori without peridia or columellae; spores not in chains, ca. 10-11 μm, reticulate. On Polygonum punctatum Ell.... Microbotryum tenuisporum (Cif.) Vánky On Ranunculaceae: 1 Sori as white to brown, swollen spots in leaves; spores embedded in the host tissue, single, hyaline, ca. 13-17 μm, smooth. On Ranunculus geranioides H.B.K., R. pilosus H.B.K. Entyloma microsporum (Unger) J. Schröter in Rabenhorst 1 Sori as galls in different organs of the host plant; spore mass brown and powdery, when mature; spores in balls formed by central teliospores surrounded by sterile fungal cells. On Ranunculus pilosus H.B.K.... Urocystis ranunculi (Lib.) Moesz On Solanaceae: Sori as spots in leaves. 1 Spores densely packed, ca. 11-14 μm, spore wall two-layered as seen by light microscopy. On Browallia americana L.... Entyloma browalliae Syd. 9

Meike Piepenbring - 10-1 Spores less densely packed, ca. 12-16 μm, spore wall mostly one-layered as seen by light microscopy. On Physalis cordata Mill.... Entyloma australe Speg. On Vitaceae: Sori in large witches brooms formed by hyperamified branches of sterile plants; sori rod-shaped, their central part being filled with a brown, powdery mass of teliospores in pairs. On Cissus rhombifolia Vahl, C. sicyoides L.... Mycosyrinx cissi (DC.) Beck References Piepenbring, M. (1995). Taxonomic studies on Ustilaginales from Costa Rica. Mycol. Res. 99: 783-788. Piepenbring, M. (1995). Trichocintractia, a new genus for Cintractia utriculicola (Ustilaginales). Can. J. Bot. 73: 1089-1096. Piepenbring, M. (1996). Smut fungi (Ustilaginales and Tilletiales) in Costa Rica. Nova Hedwigia Beih. 113: 1-155. Piepenbring, M. (1996). Ecology, seasonal variation, and altitudinal distribution of Costa Rican smut fungi (Ustilaginales and Tilletiales, Basidiomycetes). Rev. Biol. Trop. 44 (Suppl. 4). 115-123. Piepenbring, M. (1996). Two new Entyloma species (Ustilaginales) in Central America. Sydowia 48: 241-249. Piepenbring, M. (1997). The morphology of witches' brooms of Mycosyrinx cissi (Ustilaginales) on Cissus spp. (Vitaceae). Beitr. Biol. Pflanzen. 69: 177-190. Piepenbring, M. (2000). The species of Cintractia s. l. (Ustilaginales, Basidiomycota). Nova Hedwigia 70: 289-372. Piepenbring, M. (2000). New species of smut fungi from the neotropics. Mycol. Res. 105: 757-767. Piepenbring, M., K. Vánky & F. Oberwinkler (1996). Aurantiosporium, a new genus of Ustilaginales. Pl. Syst. Evol. 199: 53-64. 10

Meike Piepenbring - 11 - Piepenbring, M., M. Stoll & F. Oberwinkler (2002). The generic position of Ustilago maydis, Ustilago scitaminea, and Ustilago esculenta (Ustilaginales). Mycol. Progress 1: 71-80. Piepenbring, M. (hopefully 2003). Smut fungi (Ustilaginomycetes p. p. and Microbotryales). Flora Neotropica Monograph (accepted). 11