Terk yt 7 cs , seewtan ges biniaseey siuyeee, AY ; ; ‘ 4 Ate rey eet} bint pela snake Woe i uae . ¢ 3 iy od “hago Satu. BES is aba bathe v pet * ety! es eG one ‘WAR spa Pa aie Aer at Ve, a ree v 3 > ite ie vm ’ ’ i a Bae arte gs” NBs, A 5 * iret Metts MY he re ; a 7 aM gh . g * vine so aed af > ak rel eek _ i me Sa re MANN git at % a 8 3 ei a4 ~ vA ; } Sem RE MONE * ~ 3 \ { y ‘ i 7 ah ‘ Ff oy a ig Sek tS itn) ed. Ma yi DATE DUE EO = 22 = CO Aan BZO = MYCOTAAON AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS WO a WEN ie Gal AE IER AS) ee COM EP hE ey Neel hoe Bee OU A Rel ERG Yeu UE GON ofS TN GanO Ea eVini tet Om A G ES DNC LUD EN Ger ehGUR ES CO-ED! TORS G. Ls HENNEBERT French Language Editor & Book Review Editor RICHARD P. KORF English Language Editor & Managing Editor SUSAN C. GRUFF Assistant Editor Published by MYCOTAXON, LTD., P.O. BOX 264, ITHACA, NY 14850, USA Printed in the United States of America TABLE OF CONTENTS, VOLUME THIRTEEN Nox el April-June 1981 Marcel ewltenGa ln sderemmniscences; RLICHARDEP KORE SD icc ae ck alesse Dascomycetes Exsiccati, fasc. IV, RICHARD P. KORF & SUSAN C. GRUFF.. Alternarta themes and variations, EMORY G. SIMMONS.................. Studies in tropical Corticiaceae (Basidiomycetes) III. Two new species of Laxttextum, KURT HJORTSTAM & LEIF RYVARDEN............. Notes on Zoophthora ocetdentalts (Thaxter) Batko (Entomophthorales: Entomophthoraceae), RYSZARD MIETKIEWSKI, RICHARD S. SOPER & DAN LLANE DA LIA ien eee te rece anes ie Wns, ances Meares Lites tts ak sce sai cette coos New cicada pathogens: Massospora ctcadettae from Australia and Massospora paharitae from Afghanistan, RICHARD S. SOPER............ beotiaceses Lil. Notes onjselected temperate species referred. Co Hel Ocrummandenymenoscy Die se KEN De Pe DUMON | eee ane se teseee. eer ws ene ah ass Lichenes Exsiccati distributed by the University of Colorado Museum, Boulder: Fascicles 1-15, Nos. 1-600, 1961-1979, WILLIAM A. WEBER.. Cultural studies on Porta etnerascens, P. rivulosa, and P. subvermtspora (Aphyllophorales, Basidiomycotina), K. K. NAKASONE.. NSLCWESPCCLOSHOLMAMQL Case DAV FU lime ENKING 3 tre cris ens eee seer eS anaes Candtda paratroptcalts, a new species of Candtda, JOHN G. BAKER, TRAG PO OALKIN, SDAV IDS He PINCUS? G RICHARDOPS D'AMATO. W236 eee so Notes on Corticiaceae (Basidiomycetes). VALIES Twomnewsspecicse cfm 1 crinte ss KURIPHJIORTSTAM.GG .2 oes ts ss IX. Three new combinations in Hypochntctellum, KURT HJORTSTAM..... Cladosporium banttanum and its synonym Cladosportum trtchotdes, MICHAB UA ReEMCGUNNIGSGaDANT Ee BORELL Weems ite ceete canis ett le tm cee. 57s e A preliminary discomycete flora of Macaronesia: Part 2, Hyaloscyphaceae subf. Arachnopezizoideae, RICHARD P. KORF. Part 3, Hyaloscyphaceae subf. Trichoscyphelloideae, LINDA M. KOHN. Restnomycena gen. nov. (Agaricales), an ally of Aydropus, Mycena AROMCGeOSUOY A. mo Pensa REDHLADS GRR ROINGER, © net. te i ae Es kis 6 Stnotermttomyces, a new genus of Amanitaceae from Yunnan, China, MURGANG Semaine to ee eee ect Aree oe retnes tee eterno Tent cr atte ite Se aie vecatare ste Gene a ba Pypewstuaves imathe: !olyporacede U5, species described Dy) Jren. Reve tilenmeLE Hah) VAR ENOem rn ea ener Mitte eee cesta eee hele ee este Sat theaters The publication date,of Arendholz's thesis on leaf-inhabiting Helotiales, WOLF-RUDIGER: ARENDHOLZ G RICHARD P.SKORE. ie... - oe An alternative view of certain taxonomic criteria used in the Entomophthorales (Zygomycetes), RICHARD A. HUMBER........ geeeeeees Les Hymenochaete a éléments hyméniens pinnatifides, J. C. LEGER..... Chlortdtum and some other dematiaceous hyphomycetes growing on decaying wood. Corrections and additions, W. GAMS &@ V. BHOULBOVA = IEGHOVAR See Gate k tate iene cts eRe Mets mW eeoen urea lencrens ot! Trichophyton raubttschekit, sp. nov., JULIUS KANE, IRA F. SALKIN, ERENES WELT ZMANSG UGATHERINERSMLEKAT is sicccte scree coke tere tetetane ote's eons. co) eer tas cate Cyrenella elegans gen. et sp. nov., a dikaryotic anamorph, S. E. GOCHEN AUR tatters steels etek ed. re toners ete Gee tet otal eee etre toes, ret MaNe enetn al, cate comets aie tems fa J.W. CARMICHAEL, W.B. KENDRICK, I.L. CONNERS & Lynne SIGLER; M. SVRCEK & J. KUBICKA; SCRIPTA MYCOLOGICA no. 9; Martyn J. DIBBEN; Leif RYVARDEN & Inger JOHANSEN; Takashi MATSUSHIMA; A. NEUHAUS- LOVA-NOVOTNA & D. GUTHOVA-JARKOVSKA; Helga SCHMITZ; Edmund E. TYLUTKI; W. GAMS, H.A. VAN DER AA, A.J. VANDER PLAATS-NITERINK, R.A. SAMSON, & J.A. STALPERS; C.A.N. VAN OORSCHOT; J. JARVA & E. PARMASTO; Ralph M. SARGENT; G.A. ZENTMYER; Clive JAMES; Le ay, 85 105 112 Lv Michael R. MCGINNIS; J.E. SMITH, D.R. BERRY & B. KRISTIANSEN; Harry J. HUDSON; R.C. COOKE; Adalbert RICKEN; G. BRESADOLA; N.W. SCHAAD Notice to subscribers: replacement of missing issueS.........--+----- NOlez July-September 1981 The genus Pilophorus, H. MARTIN JAHNS..........eeeeec ese ccecrerceeces New dematiaceous hyphomycetes from tropical rain forest litter, S. ONOFRI, D. LUNGHINI, A. RAMBELLI & L. LUSTRATI...............-. Cochliobolus ravenelit sp. nov. and C. tripogonts sp. nov., J emclT es ALCORN 9, eaxGeotas. o\ Upto es 20s Woven lo apler Gatto pogeneie era + raceGs ket Ete ueeo ease tee meneueaeee A psychotropic fungus in Nepal, ROBERT F. SCHROEDER & GASTON GUZMAN. Ascus structure and function in Cochitobolus species, J. L. ALCORN.. A preliminary discomycete flora of Macaronesia: Parte oeeGeoglossaceaes RICHARD] Pe KOR aise te reiete lee ele e sdets cle + i ciena ees The psilopezioid fungi. VII. A new species of Pstlopeata exo France. DONALD H: PFISTER -G. FRANCOISE CANDOUSSAU one. cis caus os) eee Notes on soil fungi isolated from a 15-year-old aspen stand in imnterioryAtlaska wp LOLA Kh. OLIVER SG eKE LTHs VANACLEVE toe oretortesctenees eretets Coprinus sect. Herbtcolae from Canada, notes on extralimital taxa, and the taxonomic position of a low temperature basidiomycete forage crop pathogen from western Canada, S. A. REDHEAD §& Jie em RAQUA TLR 2. citrate «leila. ctehers Be etorerchnegede > see hsea ec ushel seem teat stctetem tel cae Selerotinta bresadolae Rick, a taxonomic synonym of Cibortnta eandoLlecia. (lev. WhetZele (LINDASM ws KOMN tener nis seer tiers ee ore Halystomyces, a new dematiaceous genus from Arizona's Sonoran Desertys EMORY AGS eS IMMONS ic neechetuctoiniak oratetesede ersdolets hore! ot cselets ia chy ctseeret ons Osortomyces, a new genus of the Laboulbeniales from Taiwan, KAT SOUYURIAGUBRADA te tee eieg. 1st riek. eae sr cokerels fubas olen hate Rers Von ope tocc ots tan emai eae The genus Codinaea. Three new species from the Americas, ADRIANNA DREW INGS CGF JregslicgGRANES Sen temgstotzg eh aot. tis Ones sPagins, ve odene teak’ teat ck eee ee APNeWESPECICS OL, Dacryopiyaratromebraai la Bie LOWS een arene ce one Notice-slhirds International MycologicalsConeress. ee ee ee NOLorS October-December 1981 Studies in the lichen genus Psoroma I: Psoroma tenue and Psoroma ernnamomeia, \ Av HENSSEN®&> Bie RENNERS © cw ale cn, eee een ae eee An undescribed pleomorphic species of Codinaea, NORIHIRO TOYAZAKI... The psilopezioid fungi. VIII. Additions to the genus Pachyella, DONALDOH SS PEISTER = § sFRANCOLSESCANDOUSSAU yr. ure cy cee pete nente. 2 ern Pithomyees pavgtt, a new combination for Trichocladium pavgtt and Pithomyces funtculosa, MARY E. PALM & ELWIN L. STEWART............ Validation of the Herpomycetineae and Herpomycetaceae in the Hbaboulbeniales se SABELLE TT SeTAVARES 2) tears oe ot eee Erynta (oy somcetes: Entomophthorales): validations and new species. RICHARD CA SoHUMBER sagen cts ni shrYel val scaled ene eee eet eee ee ee A review of the nonentomogenous Entomophthorales, BRUCE E. TUCKER... Erynta (Zygomycetes: Entomophthorales): emendation, synonymy, and transfers, RICHARD A. HUMBER §& ISRAEL BEN-ZE'EV Revue des med. G. L. HENNEBERT 2) 4G OS 6. 60) 18: OO) 8 as 0) OO 846) 16.8 (C6. 6: “6: 6: 6 i-«. 6: a: BY igh ie eo) whet e Brian SUTTON; C.A.N. VAN OORSHOT; K.T. VAN WARMELO & B.C. SUTTON ; D.M. DRING; J.A. VON ARX; Jan KOHLMEYER & Erika KOHLMEYER; R.C.W. 405 419 A28 431 BERKELEY, G.W. GOODWAY & D.C. ELLWOOD; E.R. WEIBEL; Sultan AHMAD & S.H. IQBAL; R. VANBREUSEGHEM & Ch. DE VROEY; A. GOFFEAU & J.M. WIAME; John WEBSTER; M.L. FARR; T.N. LAKHAMPAL & K.G. MUKERJI; G.M. OLAH, O. REISINGER & G. KILBERTUS; G.M. WATKINS; John ERIKSSON, Kurt HJORTSTAM & Leif RYVARDEN; Erwin F. SMITH & Alfred FISCHER; Peitsa MIKOLA; John PEBERDY; R. HARRISON & G.G. LUNT Notices KVEAGT LLCs OC HCTICGSGON PLES S premier ens. tv arm sie ers ise atiet aleta care Sates eee 525 TMCCEMONBNOUSINEE COSTS AN MAD al. We heer ete Rit west male are ot stecaver eres o2o AU CNOLMLNDE ARE Mercia ea icisnite ois siimte Sek egers coat aeeets starete Mee snet el sas eae stele ate verene : 526 INOUE COMLUNVUSHANG wL UCN MUG Xa vem attests tence erkt te etc Nelc tens oc cusie eel tors: 528 MYCOTAXON@ publication» dates,..12(2)7-sl3 (1b) fand 13(2) 0.0... wees ee 534 REV LOWCIS re ans ere ate cic. e ete tote crete rere at's MPa rae: Mah suet enna etc cela cre witty. 6 555 Errata my O.LUMes.| Ott nese sclera ers lsncy econ sh a et os ge so see tik Ao thn is Baler aruee Patio 535 BEAL AmmeV.O LUMO mE cere tent eteastetee abe terse ome ae eet ore en The! shel onekatal onaiel ohsie ene Hie ee S 555 EVV ata VO lUMe. 1 ON. atte wacle : )O I3 ] LIBRARY ITHACA, N. Y. 14850 @ MAY 1 21981 MYCOT AXON .«N INTERNATIONAL JOURNAL DESIGN 'O°EXPEDITE PUBLICATION OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS Volume XIII April-June 1981 Noel CONTENTS Notcerie "Le Galk va TeMINISCENce..0.0...0 eek bs vie yc bat, s ohare s cies RICHARD P. KORF 1 Diecomycetes* Exsiccati ) fase. IV. ss iene cise see RICHARD P. KORF AND SUSAN C. GRUFF be escriaria themes: and Variations... 27s 4. w'y eke 0% ow eles S dielgieke Bde wee, EMORY G. SIMMONS 16 Studies in tropical Corticiaceae (Basidiomycetes) III. Two new species ICE LORAIN wie x ata sie hioabelta o 24 wes oe siete oh ol K. K. NAKASONE 105 TOM SCE LOS POS MMIC. a Sd cosh Urn Soe. a cig nly dO K viniollals, © uw eho elt gid iwlais DAVID T. JENKINS 112 Candida paratropicalts, a new species of Candida. JOHN G. BAKER, IRA F. SALKIN, DAVID H. PINCUS AND RICHARD F. D'AMATO 115 Notes on Corticiaceae (Basidiomycetes) Witt. tiwornew species: of Tubuliorints. Oe .'s iki + Svc wie oly Vite dances | KURT HJORTSTAM 120 IX. Three new combinations in Hypochnictellum.........eeeeveees KURT HJORTSTAM 124 Cladosportum banttanum and its synonym Cladosportum trichotdes. MICHAEL R. MCGINNIS AND DANTE BORELLI 127 A preliminary discomycete flora of Macaronesia: Part 2, Hyaloscyphaceae subf. Arachnopezizoideae.............. RICHARD P. KORF 137 Part 3, Hyaloscyphaceae subf. Trichoscyphelloideae.............. LINDA M. KOHN 145 Restnomycena gen. nov. (Agaricales), an ally of Hydropus, Mycena and BEPC OTNE wahi's Wp che wie PGE ak Vow Fs sina alas cc ahuo ts typ te wceraletara: dicate S. A. REDHEAD AND R. SINGER 150 Stnotermitomyees, a new genus of Amanitaceae from Yunnan, China......... MU ZANG 171 Type studies in the Polyporaceae 13. Species described by J. H. Léveillé. LEIF RYVARDEN 175 The publication date of Arendholz's thesis on leaf-inhabiting Helotiales. WOLF-RUDIGER ARENDHOLZ AND RICHARD P. KORF 187 An alternative view of certain taxonomic criteria used in the Pucomonnthorales? (Zymomycetes) cic to wc das alse le alk bie wishslelele denims RICHARD A. HUMBER 191 Les Hymenochaete a éléments hyméniens pinnatifides.................. J.C. LEGER 241 Chloriditum and some other dematiaceous hyphomycetes growing on decaying wood, Gorrections. and additions ....5.0 nga + + ite W. GAMS AND V. HOLUBOVA-JECHOVA 257 Trichophyton raubitschekti, sp. nov. : JULIUS KANE, IRA F. SALKIN, IRENE WEITZMAN AND CATHERINE SMITKA 259 Cyrenella elegans gen. et sp. nov., a dikaryotic anamorph....... S. E. GOCHENAUR 267 NESS LA IES ee SON tae hel ee din a A Saag at oe Sopiat Sam Roos, Dante, a's WedelGie pauns G. L. HENNEBERT 278 Notice to subscribers: replacement of miSSing iSSUES.........ceeeeescecrrcereevees 287 [MYCOTAXON for January-March 1981 (12: 313-548) was issued March 17, 1981] ISSN 0093-4666 MYXNAE 13(1) 1-287 (1981) Library of Congress Catalogue Card Number 74-7903 Published quarterly by MYCOTAXON, Ltd., P.O. Box 264, Ithaca NY 14850 For subscription details, availability in microfilm and microfiche, and availability of articles as tear sheets, see back cover gts. eee eee dc Sat Ml MYCOTAXON VOIPmeX [Lie Nowele spp al —4 April-June 1981 MARCELLE LE GAL: A REMINISCENCE RICHARD P. KORF Plant Pathology Herbarium, Cornell University Ithaca, New York 14853 USA Marcelle Le Gal, born Marcelle Choquart on Febru- ary 14th, 1895 at Amiens, France, died in the same Cit yarOleilllicr2) aml 9 ame emeiniluencesOonsmycology, sand particularly on the development of the taxonomy of dis- comycetes, has been immense, even though she only en- Teredeilier iieldmwhens shemwase nearly 140 |years of age. Her early publications dealt primarily with discomycetes and showed jan) exceptional attention to’ detail and clear deliniation of characters such that I, young and wholly inexpernienced,, immediately =accepted her?’ publications as authoritative and as models of procedure. My own work One discomycetesm had» sonly sbegtn sin) 1944, 7and) in 1947 Snemecci tm iniecmnermmoOnumentalmeukecnerches sur ales) orne- mentations sporales des discomycétes operculés," her just-published doctorate thesis (which was to receive the French Academy's coveted "Prix Montagne."). I was Quickiverdrawnmeintop cde lively “correspondence: with “her, auCusCeSpilcmtlcmciilercicem ile OUlnmages susie ioheeriully commented upon and examined problem discomycetes that Deeceh elicrasOVctuaLlcwsVears me nevem Stinting= Ol8 fer time andsadvicer lGwas lO49u before we were? to. meet in) person, on Piemoccaslonmolmdauatripstomburopeans herbaria: when al was searching for type specimens of species of the -Arachno- pezizeae for my own thesis studies. At the Laboratoire de Cryptogamie of the Muséum National d'Histoire Natu- rellcustine Paris = ilmwas ecordially oreeted; bysProfessor Ro- ger Heim and his staff, but above all by Mme Le Gal herself. The memory is one I shall ever cherish, for she devoted time and energy to showing me her collec- tions, and the museum's discomycete holdings, inclu- dingeeivosemo! smerandols (and™ mine), the eminent Emile Boudier. Quiet-spoken, ever helpful, Marcelle Le Gal was from that moment on to be a major influence in my professional life. Her devotion to the study of fungi, her insight and mastery of the fungi she knew, remain an inspiration to all who knew her. An unexpected sur- prise to me as a young graduate student was that she entertained me with a sumptuous meal in her Paris apartment, where I was to meet her greatest joy, her husband, Etienne. They had married in 1922 on her re- turn from the United States where she had earned her M.A. at Columbia University. On several later occasions I was able again to par- take of Mme Le Gal's hospitality and kindness during visits to Paris. The last of these was sad beyond any expectation, for when 1 visited her in 1973 her beloved husband had only recently died. Her grief had caused her to withdraw from duties at the museum to her home in Amiens, and to stop work on the important mono- graph of the genus Scutellinia that had been occupying her for over a .decade. When’ she» learned that 1) was to visit the Paris museum, she made the unprecedented ef- fort of coming to the museum to talk with me about our taxonomic problems, and: offered me the loan of “several of her specimens of a critical genus we discussed s Her sadness over “the loss' Gof cher) husband wacmepeoiouna, ands had clearly devastated! herp emotionally imean cerino both her -personal and, professional mile we siemwacmcoon to turn her back on mycology, perhaps even on humani- ty, as the result of an unfortunate accident she suffer-— ed: in’ Amiens. and) of her grief overmthe Hossmoimherscon— stant companion-husband. Mycology's great loss is that her Scutellinia monograph will apparently remain unpub- lished and incomplete. Marcelle Le gal is one of the very few to have mar- kedly affected my development as a scientist. From her I learned patience and devotion to detailed study of minute structures as keys to relationships. Not always was I to be in her good graces; particularly because of my insistence on following the Code of Nomenclature and the principle’ of ‘prionity, sl ebecameman tpecemem nad boy" in her view. For years I licked the wounds inflic- ted by her vitreolic attack on "le jeune mycologue amé- ricain" in her masterful diatribe against the Code of Nomenclature (Le Gal, 1958). That she eventually for- gave my youthful exuberance was one of my great joys. I shall miss Marcelle Le Gal's cautionary council more than many other mycologists will; our interests co- incided closely, and it is in her footsteps that I have trod most often, secure in the knowledge that her obser- MARCELLE LE GAL (1895-1979) Photographed by the author in the gardens outside the Muséum National d'Histoire Naturelle, Paris, during the 1954 International Botanical Congress vations were dependable and her sense of taxonomic di- rection unerring. A kinder, more loving spirit has never before, perhaps, graced mycology. For those who read French, I commend a touching tribute to her by Patrick Joly (1980), that captures the essence of this most wondrous scientist. Yet another revealing tribute to her contains a complete listing of her mycological papers, the articlel pymener ) long-time associate in the field and laboratory, Henri Romagnesi (1980). REFERENCES CITED JOLY, P. 1980. Marcelle Le Gal (1885-1979). Cryptogamie, Mycol. 1: 93-96. LE GAL, M. 1958. Petite promenade A travers le maquis de la nomenclature. Rev. Mycol. (Paris) 23: Lo pee On ROMAGNESI, H. 1980. Mme Marcelle Le Gal (1895-1979). Bull. Soc. Mycol. France 96: 125-131. MYCOTAXON VOlWexX DL PANO tee pp .seo at April-June 1981 DIS GOMY CETESBEXS NC CAT Ie GRASG® SLV. RICHARD P. KORF AND SUSAN C. GRUFF Plant Pathology Herbarium, Cornell University Ithaca, New York 14853 USA Twenty-five additional numbers comprise this fourth fascicle of Discomycetes Exsiccati [for distribution of sets, see the note on page 15]. We express particular thanks to Dr. Henry Dissing and to Dr. Sigmund Sivertsen for providing us with holotype material of Boudtera denntstt Diss. & Siv. in Dissing to issue here as ISOTYPES (No. 77); a second collection of this species was collected by the senior author of this paper in their company and constitutes AUTHENTIC material from a previously unrepor- Cedmlocali ty, (Now 78), As previously noted (Dissing §& Korf, 1980), No. 84 in this fascicle constitutes ISONEOTYPE material of Ruhlandiella bero- linensts Hennings, the type material of which is presumed to have been destroyed in Berlin during the Second World War. An examination of Rehm's type specimen of Awmarta gregarta Rehm showed that it has warted ascospores, not smooth as is usually assumed. A smooth-spored variant is issued here (No. 90) as Trtchophaea gregarta (Rehm) Boud. f. laevispora Korf §& Gruff, f. nov., the specimens issued being ISOTYPES: Trtchophaeae gregartae f. gregariae similis, sed asco- sports perfecte laevtbus differt. HOLOTYPUS: CUP-MJ 611 (ISOTYPI itn Korf & Gruff, Dtee. Fxe. 90 disperst). PARATYPUS: CUP-MJ 41. On clay bank. Ctnehona Botantcal Garden, elev. 4750 ft., St. Thomas Parish, Jamatca. Lég. R.P.Korf, leader; J.R.Dtxon, K.P. Dumont, R.W.Erb, D.H.Pfister, D.R.Reynolds, A.Y.Rossman & G.J.Samuels. Ciel 71, The typical, warted-spored form is also issued (No. 89) for comparison, as Trtchophaea gregarta f. gregarta. By far the most abundant of Durand's collections of Sarcosoma eyttartotdes Rehm tm Dur., his No. 1305 (= CUP-A 12278), is 6 formally designated here as the LECTOTYPE Ofmthaemspecies, and the specimens issued (No. 79) as Plectanta cyttartotdes (Rehm in Dur.) Korf are thus ISOLECTOTYPES. [All packets of this collection in the Durand (CUP-D) and Atkinson (CUP-A) herbaria in CUP bear the data for collecting site as "Glen Mary,'' while the original description (Durand, 1903) states, 'Most abundant in Glen Burney.'' This latter name does not appear on any of the Sarcosoma packets in these herbaria. | Mr. G. Beaton, of Australia, who has consistently provided us with exciting discomycete finds, has generously given us TOPOTYPE material of the elegant species Underwoodia beatontt Rifai for issue here as No. 91. Dr. Peter Milan Petersen, of the University of Copenhagen, has contributed the material from Greenland of Sarecoleotta globosa (Sommerf. : Fr.) Korf issued as No. 100. Taxonomic and nomenclatural notes again appear on some of the labels: (79) on Plectanita cyttartoitdes as the type species of Plectania sect. Plicosporae; (87) on Karsten's use of the com- bination Seutellinta seutellata, which while the earliest is not validly published since the generic name was not yet va- lidly published; (89) on Karsten's similarly first use of the combination Sepultarta gregarta, likewise not validly pub- lished since its generic name had not yet been validly pro- posed, and on the taxonomic synonymy of Tritchophaea gregarta var. tntermedia with T. gregarta f. gregaria; (95) on an ab- berant collection or unnamed variant of Arachnopeztza cornuta having only 1-celled ascospores; (96) on adoption of Arachnio- peatza leontna instead of A. candtdo-fulva; (97) on discarding the group '"'Anomalae" of Arachnopeztza since the "granulations" on the hairs of A. trabtnellotdes (sole representative of that group) seem to be more like the resinous excretions sometimes evident in the group "Typicae" than like the eranulations on the hairs in Dasyscyphus. ACKNOWLEDGEMENTS Particular thanks are due the US National Science Foundation, which under a series of grants to the senior author has funded the collecting in Jamaica and the Canary Islands, and to the Fulbright Commission that supported the collecting in Japan. The Okinawa collection was by virtue of support from the US Office of Education, the US Civil Administration of the Ryukyu Islands, and the University of the Ryukyus. Collections from Finland and Norway were financed by the University of Copenha- gen and the University of Trondheim. Dr. William Dress, of the Bailey Hortorium, Cornell University, kindly checked the Latin diagnosis. DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 76. Anthracobta macrocystts (Cooke) Boudier, Hist. classif. discomye. Europes sp...0>, 1907. On a burnt Eucalyptus sp. clear-cut. At km mark 14, Bosque de la Esperanza, near Pico de las Process lenenite. canary iisiands. beg: R.P.K., W.C.Denison, L.M.Kohn & M.A.Sherwood 8.1.1976 Dele eRe? ake DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 77. Boudtera denntstt Dissing §& Sivertsen tn Dissing, Kew Bidet eS 5 eel 977 we SOINPE Near a small stream, Nordland, between Berget and Fisksjgmoen, 22 km NW of Mo in Rana, Norway. Leg: H.Dissing, S.Sivertsen §& T.Schumacher GS LX LOS Deten il. Dissing GvS.Sivertsen DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 78. Boudtera dennisii Dissing §& Sivertsen in Dissing, Kew Bridle olen owl 97/24 “AUTHENTIC On mud flat at mouth of brook, among Equtsetum, Calamagrostts neglecta, and Lamprospora ovalispora. Loevvaijakka, near Levajok, along Tana River, Finnmark Fylke, Norway. Doensosoivertscen,eH sDissing & R.P.K. CARS ARLE ees Det? Sisivertsen & H.Dissing DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 79. Pleetanta cyttartotdes (Rehm tm Durand) Korf, Mycologia 49: 110. 1957. [ISOLECTOTYPE of Sarcosoma eyttartotdes Rehm 7m Durand] Oni-deadttwies leaves, etc. Blue Ridge Mountains, Glen Mary, Blowing Rock, North Carolina. NOTES: This is the type species of Plectanita Fuckel sect. Pltcosporae Korf (loc. @tiajee RUE.K. Leer eeUrander( oOo) SUL VEIT ELIOT Deter. TUE DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 80) Pseudoplectanta nigrella (Pers. ': Fr.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 324. 1870. In clay soil on roots of Dtcranopterts linearts and rarely of Pletoblastus ltnearus. University of Ryukyus Recreation Area, Oku, Kumigami-son, Okinawa, Japan. Roce. ome ete DUNO G mmaekO 1 amar kh sKULOSh ma Heya) Wl bed dS Lore: Deemer Ke DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 81. Ptychoverpa bohemtca (Kromb.) Boudier, Hist. classif. agScomyc, (Europes) 1 s54eeel O0n., On ground in mixed woods. Lloyd-Cornell Preserve, McLean, New York. Leg: Mycology Class Zio SS D6Usa hae ah. DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 82. Pulvinula globtfera (Berk. §& Curt. in Berk.) Le Gal, Prodre, ploresMycolssMadacascar 4-794, 1953, NUS Oty otelcOOr Eth elev. in rain) forest, FL NUNGUC we rulert OmR1coOs Leg: G.Abawi 2ipie L96G DEUS R TEs Ke DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 83. Pulvtnula niveo-alba Moravec, Cesk4 Mykol. 23: 231. 1969. On duff. Juuma-Jakalavuoma, Kuusamo, Finland. Leg: S.Sivertsen Or ES AG Dette wows. DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 84. Ruhlandtella berolinensts Hennings, Hedwigia 42: (24). 1903. ISONEOTYPE, designated by Dissing & Korf, Mycotaxon 12: 295. 1980. Oneso bia Burn site along road in Eucalyptus grove at km mark 14, Bosque de la Esperanza, Tenerife, Canary Islands. Legaik.P.K.%, Ro Fogel, GL. Hennebert & LSM /Kohn 299X1T. 1976 pet: HeDussingeg Rey Ke 10 DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 85. Seutellinita erinaceus (Pers. : Fr.) O. Kuntze, Revis. gen. pl. 2: 869. 1891. On mossy wood. Michigan Hollow, Danby, New York. sGamundi ,s Kos. Thind, We. Denison, 9.VIII.1960 Leg: I.J Ree Moone, | Verelewacorgukers Ks Ree Kk. Det: DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 86. Seutellinta pennsylvantca (Seaver) Denison, Mycologia Sila Lobel SO al SOO eae On old wood. "Big Woods," SW of Ann Arbor, Michigan. Leg: L.B.Wehmeyer, R.P.K. G al. 19.VI1I.1948 Det Wc. Denison | DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 87. SCcuLeLlinid, scureliata, (Ll. sa Fre) Lambotte., bh leemucols belpesmoupphs l2299 4 -1837- | On Fagus sp. Lloyd-Cornell Preserve, Slaterville, New York. NOTES: Neither a generic name "Seutellinta (Cooke) Karst." nor the combination "S. scutellata (L.) Cooke" was validly published > i t Fl. Fenn. 11: 145. 1884) [ICBN, Arts. 4h, 43]. R.P , arsten (Medd. Soc. Fauna Leg: M.A.Rosinski Pre1os 2 Det: W.C.Denison 11 DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 88. Scutellinta umbrorum (Fr.) Lambotte, Fl. mycol. belge, Suppl. 1: 300. 1887. One soil: Montezuma Wildlife Preserve, Seneca County, New York. eC cme MUL bed hares 20 GNI LOGZ DOC aR ek’ DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 89. Trtchophaea gregaria (Rehm) Boud. f. gregarta, Hist. Class DiemdiSCONny CmmeEUT Ope map mmo UmLO O74 On soil and rotting plant matter. Peck Foray, Ashokan, New York. NOTES: Neither a generic name "Sepultarta (Cooke) Karst." nor the combination "S. gregarta (Rehm) Karst.'' was validly published by Karsten (Medd. Soc. Fauna FJ. Fenn. 11: 145. 1884) [ICBN, Arts. 41, 43]. Le Gal's T. gregarta var. inter- medta Le Gal [Rev. Mycol. (Paris) 2: 214. 1937) is a synonym, since Rehm's type specimen also has warted ascospores, not smooth as she assumed. R.P.K. Leg: D.Malloch, R.P.K. & al. TORI ao 2 DeGieR. ake DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 90. Trtechophaea gregarta (Rehm) Boud. f. laevtspora Korf §& GUL ieMy CotaxOnel > omLoo le 1 SULYPE Onwclayesoilebank: Trail between Freetown and Wag Water River, near Hardwar Gap, St. Andrew Parish, Jamaica. Becca herwe eR aDEx Ole Kab DUNO Coe he Wi 5 Peps INS ea Dehabrister,. Usk -Reynolds, -A.Y Rossman §& G.J.Samuels DOC SRG Pe KersGio Ga. 4 DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 91. Underwoodia beatontt Rifai, Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect., Su eal Roles. 1968s TOPOTYPE In sandy soil and debris under Melaleuca lanceolata beside cliff walk, Anglesea camping ground, Australia. Leg: G. Beaton ZoeN tile LOGS Det. Geb. DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 92. Urnula cratertum (Schw. : Fr.) Fr., Summa veg. Scand., SeCt wy p0S te, p.) 504.8 O40. On soil and wood. Woods near Westhaven Road, Town of Ithaca, Tompkins County, New York. HEC mn ewe hee OM CN seh el ROLL Gh rake V.1960 Dette RePak. DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 93. Wynnea amertcana Thaxter, Bot. Gaz. 39: 246. 1905. On soil in woods. Z-oumiLes-east of Obi, -at crest, of shuld elev 0G te. Alleghany County, New York. GeMomith sede biuhms ad. MoROni iG Rar ake ZEN LOO Deus, Gi GeMio- ik) DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C., GRUFF, EDITORS 94. Arachnopeaztza aurelta (Pers. : Fr.) Fuckel, Jahrb. NassaudschenmycreinseNaturk. (2 5224' 1305451870. On bark and acorns of Quereus sp. France Brook, Alleghany State Park, New York. Ibe Oeamhy Det Kean Gala | OAV bOGd DEC waht Re ke DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 95. Arachnopeztza cornuta Cli Ss aKonieesl boyd 40158" Tey eae LS) ee (ey ae | On rotted wood. Lloyd-Cornell Preserve, Ringwood, New York. NOTES: Apparently lacking the 2- and 3-septate ascospores by which one usually recognizes this species. R.P.K. Leg. Gel-Hennebert (3096) & RYPLK. WV L962 Det raiRe Pike DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 96. Arachnopeztaza leontna (Schw. : Fr.) Dennis, Kew Bull. Lee OO Leet OOS . On a very rotted log. Bast send of Lake Shikotsu, Ibari Pref. , Hokkaido, Japan. NOTES: This specimen was reported by Korf (Bull. Natl. Sci. Mus. 4: 392, 1959) as A. eandtdo-fulva (Schw.) Korf. Dennis's transfer is correct, since Schweinitz aie CL Leontna earlier than Pezisza candtdo-fulva, and Fries accepted begamoimaigus, Kamei, Y,Otani, R.PvKy 6 al. 20.V.1958 DetewRer Ks 14 DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 97. Arachnopeztza trabitnellotdes (Rehm) Korf, Lloydia 14: POS 9 5 28 ( 8 Goel We On wood. Margaretville, Delaware County, New York. NOTES: The "granulations" on the hairs are more like resinous excretions than the granulations of Daeyscyphus spp. of the ''Typicae" group. This species be- longs close to A. cornuta in Arachnopeaiaa group "'Typicae," and should not be segregated in a group "Anomalae"' as was done by Korf (loc. ctt.). R.P.K. he Ocarina PelNOY eC taegm habe. eh Gals eye! DeGeeRn rake DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS O8MParachnopesiza mintopsts /(EI1s Wakore, aMycotaxonmy. AGS) SESWasy. Ong bark: France Brook, Alleghany State Park, New York. Tee a Rep ko 1G salt Gavi 1o6l DeGeaR yPaks DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 99. Pezoloma larteina (Ell. & Everh.) Korf, Phytologia 21: 2 Ooi Oe. | On duff, mostly Tsuga needles and cones. Peck Foray XIV, Twin Valleys, Lewis, New York. begs Gaapawi, al. PEister 4 Re bak. 15.V1I.1968 Deets RP ak. 15 DISCOMYCETES EXSICCATI DEPARTMENT OF PLANT PATHOLOGY, CORNELL UNIVERSITY RICHARD P. KORF & SUSAN C. GRUFF, EDITORS 100. Sarcoleotia globosa (Sommerf.: Fr.) Korf, Phytologia Ze Orme oe On Sphagnum fuseum. Sydpréven (60°21'N, 45 34'W), Greenland. ever Mialane beter senile OZ) SON LO fel Deter RP. ke DISTRIBUTION OF SETS The first fascicle of this series was deposited in 11 institu- tions (Korf, 1958), with 2 additional ones listed by Korf and Gruff (1978) as receiving fascicles II and III. By error they listed 'BP = Museum of Natural History, Budapest,"' whereas the sete s actually on depositeat BPUN= Institute of Plant Taxono- my and Ecology of Edétv6s L. University. A 14th set is now on deposit at Copenhagen, also beginning with fasc. II. The com- plete list of deposit herbaria is thus: BPI (Beltsville) FH (Cambridge) * NY (New York) BPU (Budapest) * K (Kew) RPG (Paris) C (Copenhagen) * LAH (Lahore) PR (Prague) CUP arthaca) MICH (Ann Arbor) UC (Berkeley) DAOM (Ottawa) UPS (Uppsala) lack IngmrasciGloml . REPERENCEs Ci TED DLSSING gun AND OR cee hORP @LOSOe ePreliminany ssceudres in the genera Ruhlandiella, Sphaerosoma, and Sphaerozone (or- der Pezizales). Mycotaxon 12: 287-306. DURAND, E. J. 1903. The genus Sarcosoma in North America. Jour. Mycol. 9: 102-104. ROLE hoes el OooEuea 1 SCOmyCeLeae exSiccatae, asc. 1.) Myco- LOU a4 Oto oy Been Oona KORE, oR. P. “AND S.C. GRUFF. 1978. Discomycetes exsiccatti, Pascal hq ey COtaxXOn a) tl 3o>Z2U0s. MYCOTAXON VO XL eNOvae lee p Dram O 704 April-June 1981 ALTERNARIA THEMES AND VARIATIONS EMORY G. SIMMONS Department of Botany, University of Massachusetts Amherst, Massachusetts 01003 PROLOGUS Being the initiation of a series of observations on Alternaria species, in this instance on some whose nonbeaked conidia very rarely occur in chains (A. molesta sp. nov., A. mouchaccae nom. nov.) and others whose erostrate conidia prolif- erate in chains by means of interpolated conidi- ophores (A. chlamydospora, A. limaciformis sp. nov., and two commonly misrepresented phytopath- ogens, A. triticina and A. longipes). I. EROSTRATAE, SOLITARIAE Most species of Alternaria produce conidia in chains, some readily and abundantly under conditions of either nature or culture (e. g., A. tenuissima (Kunze in Nees & Nees ex Persoon) Wiltshire), others sparsely and perhaps only tardily after manipulation of culture medium, light quality, and temperature. This latter reluctantly catenu- late group includes, among others, common phytopathogens whose conidia are thin walled and filamentously rostrate (e. g., A. zinniae M. B. Ellis) as well as others whose conidia are relatively thick walled and erostrate (e. g., A. radicina Meier, Drechsler & Eddy). Alternaria species of a third group produce two conid- ia in a chain so rarely, under any condition reported or observed thus far, that any chain can be considered little more than an aberration. Two notable members of this group are A. chrysanthemi Simmons & Crosier (Simmons 1965) and A. helianthi (Hansford) Tubaki & Nishihara (1969). Both of these species are phytopathogens; the nonbeaked conidia of both are relatively large for the genus, with lengths up to 120um and 110um respectively (Fig. 1). 17 An additional member of this group with erostrate, solitary conidia is known in the form of a small-spored species isolated from a skin lesion of a captive harbour porpoise (by S. Andersen, Odense, Denmark; transmitted by G.-A. de Vries, Baarn, The Netherlands, as CBS M 339.77). It is reported here as novel on the basis of examination of axenic cultures, primarily on potato-carrot agar, hay extract agar, and 20% V-8 juice agar (hereafter as PCA, Hay, V-8; see Stevens 1974), in a diurnal fluorescent light/dark eyclenpat 22'C™ 1. Alternaria molesta Simmons, sp. nov. (Fig. 1) [molesta = troublesome (to both Phocaena and Simmons, though in different ways) ] Ex culturis in agaro PCA descripta. Coloniae griseo- brunneae, lanuginosae, postea atrobrunneae, applanatae. Mycelium initio ex hyphis septatis, ramosis, subhyalinis vel pallide stramineis, levibus, 3.5-5.0um crassis composi- tum. Conidiophora simplicia, singulatim ex lateribus hypharum oriunda, recta vel flexuosa, cylindrica, pallide flavo-brunnea, levia, 0-10 septata, plerumque 4-5um crassa, usque ad 100 (plerumque 5-20)um longa, apice rotundato et uniporoso, raro geniculata. Conidia solitaria, subcylin- drica, ovoidea, obclavata vel ellipsoidea, septis trans- versalibus plerumque 3-6, longitudinalibus nullis vel paucis, ad septa definite constricta, pallide straminea, levia, 15-38 x 7-12 (plerumque 27 x 12)um. Origo typi: ex laesione pellis Phocaenae phocaenae, Odense, Daniae. Typus: partes ex Simmons 32-075 (CBS M 339.77) desiccatae et in BPI, CBS, DAOM, IMI, NY conservandae. The type isolate is stable and predictable in its growth and sporulation characteristics under specified culture conditions. It grows well on PCA and V-8 (15mm radially in 4 da), producing colonies that are dark olive- brown with a shallow aerial layer of paler woolly hyphae. Conidium production occurs initially on hyphae near the substrate surface but spreads gradually to the intertwined aerial elements. After the agar surface of an entire culture plate has been overgrown, the colony becomes almost Opaque because of the very dark color of both the submerged and the aerial hyphae. Although the mycelium of colonies on Hay is scarcely visible, it gives rise to conidia readily and abundantly. Conidia of A. molesta are produced one per conidio- phore tip; conidiophore tip proliferation (geniculation) is uncommon. Among many hundreds of conidia viewed in slide 18 preparations, only half-a-dozen had an apical l=cell exten- sion with a terminal scar, and thus presumably had served as the basal unit in a chain of two-conidia. The color of conidium and conidiophore walls is so dilute that septa and disjunction scars appear in bold contrast in transmitted light. A very high percentage of conidia mature as ellips- oid to ovoid units with three transverse septa and at a size of about 27 x 12um. The conidium population is, for an Alternaria species, remarkably restricted in morpholog- ical variation, with very little tendency to produce longi- tudinally asymmetrical or excessively cellular entities. Fig. 1 illustrates conidiophores and a representative range of A. molesta conidium shape, size, and septation. Conid- jum outlines for A. chrysanthemi (Fig. 1, lower left) and A. helianthi (Fig. 1, lower right) are included for pur— poses of comparison. 2. Alternaria mouchaccae (Fig. 2) Arid and semidesert soils harbor remarkable numbers of phaeodictyosporic hyphomycete species (Durrell & Shields 1960; Kuehn 1960; Ranzoni 1968; States 1978). New taxa al- most inevitably have been unearthed and described in Ulo- cladium, Embellisia, and Alternaria. Pertinent to the present discussion is a species published as Ulocladium chlamydosporum Mouchacca (1971), which was described as new on the basis of several isolates from arid and newly culti- vated soil of the New Valley region of Egypt. A second | dematiaceous entity, often associated with U. chlamydospo- rum in nature and morphologically similar to it, later was published as a new species Alternaria chlamydospora Mou- chacca (1973). Study of representative strains.of these two species (IMI 156434 and IMI 156427 respectively; EGS 31-061 and 31-060) reveals that they are congeneric in Alternaria., Their, morphologicalwsimilarity andenature association in the soil tempts one to speculate that they also may be conspecific although culturally in different phases of progressive degeneration (a condition noted by Mouchacca, 1971). However, relying to the greatest extent possible on comparisons of l-conidium reisolates on PCA and Hay agars, of conidiogenesis, and of conidium development before typical cell distortion begins (Figs. 2 & 3), I con- clude that there are two distinguishable species, Alter- naria chlamydospora Mouchacca (discussed below as species no. 3) and Alternaria mouchaccae Simmons, nom. nov. [= Ulo- cladium chlamydosporum Mouchacca, Revue de Mycologie 36(2): 120, 1971; non A. chlamydospora Mouchacca, 1973]. i oa sal sas on PCA; raoh tye Alternaria molesta ex Type (at top), NS chrysanthemi (Vowerd ihe hoe): A. jm @ AL 3k Evioy ie Jo) aL from cGulLtune (lower 20 A. mouchaccae is yet another example of a species that produces nonbeaked solitary conidia when grown on media of low nutrient content. False conidial beaks (secondary conidiophores) are extremely rare under such conditions; only two were found among hundreds of conidia examined. A few young conidium initials might be misinterpreted as ulocladioid on the basis of a narrow isthmian attachment to the conidiophore. However, most young conidia are spher- ical or ovoid from the beginning and have bases in close contact with conidiogenous cells, which themselves exhibit conspicuously pigmented areas around the terminal disjunc- tion scars — characters more nearly typical of Alternaria than of Ulocladium. Conidium basal scars appear always to have been pushed into an eccentric position following the division of any basal cell by a vertical or oblique septum. Enlarging conidia often become conspicuously swollen and distorted, especially when produced in colonies on V-8 agar. Thesstrain studied, Vcultura’typica, s(@Mouchacca 1972). aex— hibits abundant nonsporulating hyphal overgrowth, although conidium production on nonaerial mycelium remains good as long as only conidia are used as inoculum in serial transfers. Considerable experience with moderately unstable iso- lates of this sort suggests that hyphal chlamydospores, stromatic cellular masses, and progressive degeneration originally described for this material, and still present, may be characteristic of the typical culture rather than of the species in toto, and that focusing attention on the | morphology of conidia before they become distorted may permit recognition of A. mouchaccae among future isolates, whether or not accessory structures are present. II. EROSTRATAE, PROLIFICANTES The character "conidium with beak," i.,e., a conidium with an apical portion morphologically distinguishable from the septate sporebody, strongly influences the commonly held generic concept of Alternaria. Neergaard (1945) dis- cussed the nature of the conidial beak, emphasizing its species-diagnosis utility and making a strong point of differentiating between "true beak (eurostrum)" as an inte- gral part of the conidium and "false beak (pseudorostrum), formed like the conidiophore of the species concerned." The nature of the Alternaria false beak remains poorly understood, however, if we judge on the basis of much of the available descriptive literature and posited ranges of conidium length. Evaluation of this literature, a complex DB oO u S) rw) rd = (S) (TRIG) Without locality, Sept. 913 ,on, Cake Dvanehesm iy = inqeon therdround;.C.G.) Llovyan4Gyy ie( TRIG Fruitbody soft, resupinate and widely effused, but loosen- ing at the margin which is striate or fibrillose, about 0.2-0.4 mm thick with the trama distinctly developed, hy- menium smooth, creamish to dull ochraceous, more or less cracked. Hymenial layer somewhat more dense than in the other species of the genus. Hyphal system monomitic with loosely interwoven tramal hyphae, golden yellow, straight and encrusted. The hyphal wall is thickened and the dia- meter may vary from 3,5 to 6 um, often with adventitious septa between each clamped cell. Subhymenial hyphae more irregular, pale coloured to hyaline. Oleiferous hyphae occur more or less abundant, usually with a slight amyloid reaction. All hyphae with clamps. Gloeocystidia arising. from the tramal layer, fusiform, subulate or with obtuse apex, sometimes with schizopapilles, generally 70-80 x 5 um, but the length varies considerably and they may some- times be up to 100 um or more, mostly without amyloid re- action, negative in benzaldehyde. Basidia narrowly cla- vate, 15-25 x 4-5 um, with four sterigmata.; Spores) sub= globose to ellipsoid, echinulate, (4-)4.5(-5) x 3-3.25 um, strongly amyloid. Remarks. The species is somewhat similar to L. lutescens (see below), but is easily distinguished by its slightly larger spores, encrusted hyphae and paler hymenium. Furthermore, all known specimens are resupinate. The hymenial layer is denser and the gloeocystidia are non- amyloid or only very weakly so. Because of its well deve- loped subiculum and negative reaction in benzaldehvde the species seems to belong in Laxitextum and is related to L. bicolor. However, it also has similarities to a group of species in eee ee such as G. propinguum (Jacks.) Parm., G. sibiricum, Parm.,@and Gloeocystidium lacticolor Bres. (type examined). phe Main differences are the distinct subiculum in Laxitextum and that oleiferous hyphae are lacking in these species and the sulfovanilline reaction sis positive, (except in G. lacticolor?). Another oi, Fig. 1. Laxitextum incrustans. a) section through) thes hym= enium b) basidium c) gloeocystidium d) subicular hyphae e) spores. From the holotype. 38 R Fig. 2. Laxitextum lutescens. a) section through the fruit- body b) section through the hymenium c) basidium d) sub- iculur hyphae e) spores. From the holotype. 39 species which also comes close to the group is Gloeocysti- diellum furfuraceum (Bres.) Donk, but it lacks clamps. We have also compared the new species with Scytinostromella humifaciens (Burt) Freeman & Petersen because of its some- what Similar spores and gloeocystidia. However, in that species the spores are smaller and the gloeocystidia react positively in benzaldehyde. Furthermore, itke all species in Scytinostromella, aEsOn pr Humitacvens, Lt 1S) distinctly dimitic with skeletal hyphae. Fig.3. Spores of a) Laxitextum bicolor Colles Ryviee 12092) 5) L. lutescens, holotype c) L. incrustans, holotype d) en- crusted hyphae from L. incrustans, holotype. SEM by L. Ryvarden. 40 LAXITEXTUM LUTESCENS Hjortst. & Ryv. nov. spec. Fuctificatio resupinata vel distincta reflexa, 0.4-0.6 mm crassa, supra infuscata; tramate distincto evoluto, 024- 0.5 mm crasso; hymenium leve, cremeum vel lutescens, leviter rimosum; systema hyphale monomiticum; hyphis cum fibulis, rectis, levibus, laxe intertextis, luteobrunneis, circiter 4 um latis; hyphis subhymenialibus irregularibus, hyalinis; gloeohyphae et gloeocystidia numerosa, amy loides; basidia’ anguste clavata,,.20=-25. x 7 cum, <4 sterigmatibus; sporis subglobosis vel ellipsoidibus, echinulatis, 4 x 3- 3.25 um, valde amyloidibus. HOLOTYPUS: Africa. Ghana. Ashanti Region, Bobiri Forest Reserve, ab. 30 kmsB Of Kumasi... 16.-22. Apri 6097 4a Ryvarden 12875 (0). Fruitbody resupinate to distinctly reflexed, 0.4-0.6 mm thick, pileus deep yellow to pale brown velutinate, azonate, trama well developed, pale yellowish brown, about 0.4-0.5 mm thick, hymenium cream-yellowish to strawcoloured with a light purple-brown tint, slightly cracked. Hyphal sys- tem monomitic, tramal hyphae loosely interwoven, pale yellowish brown, 3-5 um wide, smooth and with clamps. Sub- hymenial hyphae more irregular with the colour pale yellow or hyaline, mostly collapsed. Oleiferous hyphae abundant, 5-6 um wide, distinctly amyloid and with clamps at long intervals. Gloeocystidia numerous, arising from the tramal layer, fusiform, with the apex obtuse, now and then with schizopapilles, 90-120 x-7-9 um, sometimes up to 150-200 um Or more, distinctly amyloid, but negative in benzalde- hyde. << Basidia’ narrowly clavate, 20-257 x) 4 umjawatne oun sterigmata. Spores subglobose to ellipsoid, echinulate, 4X3 S220 UR, eeStrongl ye amyloid. Remarks. Laxitextum lutescens seems to be well located in in the genus and is similar to L. bicolor, but delimited by its deep yellowish colour , the amyloid gloeocystidia and Oleiferous hyphae and the shorter spores. The tramal hyphae of L. bicolor are besides more distinctly brown. References. Boidin, J. 1958. Heterobasidiomycetes saprophytes et Homobasidiomycetes resupinés. V. Essai sur le Genre Stereum Pers. ex S.F. Gray. Rev. Mycol. 23:318-346. Boidin, J. 1960. Le genre Stereum Pers. s.l.au-iCongo Belge. Bull. Jard. Bot. Bruxelles 30:51-74,283-355. Eriksson, J. & Ryvarden, L. 1976. Corticiaceae of North Europe. 4:549-889. Lentz, P.L. 1955. Stereum and allied genera of fungi in the upper Mississippi Valley. U.S. Dept. Agric. Monogr. 24. Talbot, P.H.B. 1951. Studies of some South African re- Supinate Hymenomycetes. Bothalia 6:1-116. MYCOTAXON Vole DLE mNOwEl Ss pple41=49 April-June 1981 NOTES ON ZOOPHTHORA OCCIDENTALIS (THAXTER) BATKO (ENTOMOPHTHORALES : ENTOMOPHTHORACEAE) | RYSZARD MIETKIEWSKI Agricultural and Teachers University 08-110 Siedlce, Poland RICHARD S. SOPER USDA Insect Pathology Research Unit Boyce Thompson Institute Ithaca, New York 14853 STANISLAW BALAZY Wielkopolski National Park 62-040 Puszczykowo, Poland INTRODUCTION The species Entomophthora occtdentalts (Thaxter) 1888 was described by Thaxter (1888) as a species of Empusa. He recorded this pathogen as frequently and commonly infecting aphids on Betula popultfolta Marsh in Maine, U.S.A. Batko (1964b) renamed this species Zooph- thord ocetdentalts (Thaxter) and then placed it in the subgenus Zoophthora Batko (1966b) which includes the group of species referred to as the "Entomophthora sphaerosperma group" (Hutchison 1965; Waterhouse 1975; Remaudiere et al 1976; Zimmermann 1978) and recently defined as Zoophthora Batko sensu stricto (Remaudiére and Hennebert 1980). ik This research supported in part by a grant to the senior author from the International Research Exchange Board (IREX) New York and by a grant from the U.S. Department of Agriculture PL-480 Program PL-ARS-/75 (FG-PO-358). 42 Further notes on the occurence of this species are scarce and data on its morphology constitute only abbreviated , records of the original description. Since the systematics of species within the subgenus Zoophthora has not yet been thoroughly defined and because the biology and morpholog- ical characteristics of Zoophthora phallotdes Batko (1966a) closely resemble those of Z. occtdentalts, it is useful to present the results of our observations of these fungi in recent years. MATERIALS AND METHODS In August and September of 1977 in Orono, Maine, five entomophthoraceous species were discovered on aphids Myzus persicae Sulz. infesting potatoes: Z. occidentalis, Erynta neoaphidis (Remaudiére) Batko, Z. exttialts (Hall and Dunn) Batko, Contdtobolus thrombotdes Drechsler (= Entomophthora virulenta Hall and Dunn), and Entomoph- thora major (Thaxter) Gustafsson. Of these, Z. occtden- talts was predominant. This pathogen was also collected from Aphts fabae Scopoli on pigweed, Chenopodium album L. The strains of this species were examined in the laboratory with application of usual microscopic techniques and suit- able methods of cultivation on egg yolk and Sabouraud's maltose agar (EYSMA) and potato-dextrose agar (PDA) in 16 hours photophase. Laboratory infections were also attempted using several aphid species and caterpillars of Chortstoneura fumtferana Clem. with conidial showers of the fungus. On aJuly 13, 19/8; a morphologically -simitar strain was found in the Wielkopolski National Park, Poland, on an aphid of the tribe Macrosiphina from Deschampsta caespitosa.(L.) P.B. in the forest undergrowth. sthese materials have been preserved as microscopic preparations in lactophenol with cotton blue (LPCB) and the strains from the USA also in living cultures on media. In the course of the studies on the morphology and systematic position of these strains, a comparison was made with the holotype of Zoophthora phallotdes, and data on the para- types of this species were obtained. RESUETS A. Morphology and Diagnostics. The infected aphids produced rhizoids (Fig. 1) from the abdominal part of their bodies as 1-3 compact bundles which firmly attached them to plant leaves. The tips of the hyphae forming these rhizoidal bundles are widened and deeply sinuate (Fig. la). 43 FIG. 1. Rhizoids of Z. occtdentalts from hosts producing (aye conidiarvand™(b) resting spores. Bar’=)'(a) 20), (b) 1Op- During the time of sporulation, the conidiophores covered the bodies with a compact mass of silver-white to white- cream mycelium; on the aphid from D. caespttosa the color of the mycelium was dirty lily. Pseudocystidia tapered with slightly blunt tips are scarce, and not a constant feature in the microstructure. The conidia (Fig. 2) are elongated, with sides bowed slightly outward; conidia may be slightly curved. The conidium narrows slightly towards the papilla, and has a bluntly conical apex. In outline, the papilla is tri-~ angular, and often bears a small central apiculus or arch- wise and obtuse. The single nucleus is oval or elongate, and strains well in LPCB; the plasma appears somewhat hyaline with several granules. Dimensions of the conidia from aphids or from media differed remarkably: From aphids they were found to be 25-26:5u X 8-10yn (av. 30 X 9), L/D 2:3-3:5 (av. 3.0). From EYSMA conidia were 24-42y X 7-10, Cave 318X791) L/D)324—4: 5) (av.2-4.0). Secondary+conidia of the first order are similar to the primary ones buta little shorter. After the transfer of the conidia from EYSMA to PDA, capillispores are formed in great numbers, and a few secondary conidia of further orders usually appear; these 44 PiG. 2 « elongate and slightly curved. Primary conidia may produce secondary capillispores at the end of a fine tube (c); these secondary conidia are more strongly curved than primary conidia and lack a definite papilla (d). Bar=10p. Primary® conidial ofl occidentalis (a. bimare 45 latter conidia are short and considerably different from the primary, with L/D ratio less than 2. Capillispores (Fig. 2c,d) are found relatively seldom on aphids. Capil- lispores from aphids were found to be 20.2-25.1 X 7-9u, and, from media (EYSMA after transfer to PDA), 22-33.3 X iJ De esoe5 ey Cave Os7 ) Resting spores (Fig. 3) were formed in artifically infected aphids or on EYSMA at the temperature 25-27°C. They are smooth, globose, (20.5) 24-30 (32.5)u diameter, Witheaslaehteyel low wall 5-6 thicks 90n EYSMA at 25-27°C, resting spores were larger (20-41.5y) than in the insect. On the basis of many observations of early starges of spore development, it is concluded that resting spores are formed both as azygospores and zygospores, even though the "classical" types of conjuation noted and illustrated by Thaxter (1888, p. 171-172) were not observed. Misidentification of Z. oectdentalts and Z. phallotdes is possible because of the similarities of their published spore dimensions and, more importantly because of imprecise characterizations of the conidial apex in the description of these species. Thaxter (1888, p. 171) described the apex of Z. oectdentalts as "tapering strongly" whereas Batko (1966a, p. 10-11) referred to Z. phallotdes as "sharply pointed". On the basis of the analysis of differ- ences described by Remaudiére et al. (1976), studies which did not consider Z. oectdentalts, the authors intended to provide a diagnosis for Z. phallotdes. However, the com- parison of the conidia of both species shows that the following alternatives would be more suitable: - conidia cylindrical with parallel sides in outline, hemispherically convex papilla and broadly obtuse SC cm i er ee eer ek ee ee emia LL LOtaeS - conidia ellipsoidally spindle- shaped with the sides a little convex, papilla widely conical or slightly convex often with small sharp central apiculus, top part of the spore tapering With somewhat blunt apex... ... . 24.. ocetdentdaite B. Biology and Occurance. JZoophthora occtdentalts was the most frequent pathogen among entomophthoraceous fungi collected from the aphid M. perstcae in Orono, Maine, causing 70-80% among the cases of mycosis; whereas the FIG. 3. Resting spores of Z. occtdentalts are (a) thin walled in the early stages of formation and may be ZY Z0- spores (b); the resting spore contents are granular at first (c) but contain an oil globule at maturity Cae Bar = (a-c.d) e720, (bb) Lone. 47 total mortality caused by entomogenous fungi was very high at the end of the growing season. On the media used in this investigation, the fungus grew and developed well, and produced all forms of spores. Under laboratory conditions, some species of potato aphids - viz., M. persicae, Macrostphum euphorbtae Thom., Aphts nasturtit Kalt., and Acyrtostphon solant Kalt. - were successfully infected with the spores produced on EYSMA. Attempts at similar infec- tion against larvae of spruce budworm, C. fumtferana, failed. The relative absence of Z. occetdentalts from published reports probably derived form its misidentification as Z. radtcans (Brefeld) Batko (= Entomophthora sphaerosperma Fres.). Even the investigations of Shands et al. (1958; 1962; 1963; 1972) on the entomogenous fungi of potato aphicé in Maine, carried out between 1952-1972, did not report the presence of Z. occtdentalts. Likewise, Gustafsson (1965) does not mention it from Sweden. Petch (1939; 1944) on the other hand reported it from two localities in Great Britain but contrary to his usual procedure offered no precise description. Batko (1962) listed it among Polish Entomophthoraceae from an aphid on nettle, Urttca dtotca L., in the Bialowieza National Park, where, despite many repeated thorough searches, this species was found on few aphids only in one locality. Thus it may be interesting to determine the circumstances in which the increase of its frequency in natural conditions occurs, as well as to continue attempts to experimental infections in biological control of aphids. ACKNOWLEDGEMENTS The authors wish to give thanks to Dr. Andrzej Batko (Warsaw University, Department of Plant Systematics and Geography) for rendering the holotype of Z. phallotdes accesible, to Dr. Ru%ena Krejzova (Czechoslovak Academy of Seiences, Laboratory of Insect Pathology, Praha), for furnishing the data on paratypes of this species, and to Dr. Danuta Krzywiec (Polish Academy of Sciences, Institute of Agrobiology Poznan) for identification of the aphids from Deschampsta caespttosa. REBERPNCE SSO LED BATKO, A. 1962. Notes on Entomophthoraceous fungi in Poland. se encomophnaca, 2: 129-139" 48 BATKO, A. 1964a. On the new genera: JZoophthora gen. nov., Triplospertum (Thaxter) gen. nov. and Entomophaga gen. nov. (Phycomycetes: Entomophthoraceae). Bull. Acad. PoleeSci=@ClneLi.. L2C7), Sen Sci ee Lotter azo. BATKO, A. 1964b. Some new combinations in the fungus family Entomophthoraceae (Phycomycetes). Bull. Acad. Pol escis Cle fl.) 12.09), eSer-) Scifi hi0 baaGe OG BATKO, A. 1966a. A new aphidicolous fungus from Poland Zoophthora phallotdes sp. nov. (Entomophthoraceae). Actary Mycol. 2) /—15. BATKO, A. 1966b. On the subgenera of the fungus genus Zoophthora Batko 1964 (Entomophthoraceae). Acta My cole 2: 21521". GUSTAFFSON, M. 1965. On species of the genus Entomoph- vnora Fres. in Sweden. I. Classification and distribution. Lantbrukshoegsk. Ann. 31: 103-212. HUTCHISON, J.A. 1963. The genus Entomophthora in the western hemisphere. Trans. Kansas Acad. Sci. 66: 237-254. PETCH, IT. 1939. Notes 'on‘entomogenous fungi. Trans. Brit. My COM moOCe m2 O02 ita 27 -— Las. PETCH, T. 1944. Notes on entomogenous fungi. Trans. Brit. MyCOl errs OCeag2/Ma Gla 2) Ol—98e REMAUDIERE G., and G.L. HENNEBERT. 1980. Revision systematique de Entomophthora aphtdis Hoffm. in Fres. description de deux nouveaux pathogenes di aphides. Mycotaxon L269 .5 2 1a REMAUDIERE , Geo. UKELLER, 9B. PAR LEROK C= Ee LATGE. 1976. fonmuereciens systematiques et biologiques sur quelques espéces d'Entomophthora du groupe sphaero- sperma pathogenes d'insectes (Phycomycetes: Entomophthoraceae). Entomophaga 21 (2): 163-177. SHANDS, W.A., S.G. THOMPSON, G.W. SIMPSON, and H.E. WAVE. 1958. Preliminary studies of entomogenous fungi for control of potato-infesting aphids in Maine. J. Boon. Ente im Lo4—1 66). SHANDS, W.A., I.M. HALL, and G.W. SIMPSON. 1962. Entomophthoraceous fungi attacking the potato aphid in Northeastern Maine in 1960. J. Econ. Ent. 55: 174-179. SHANDS, W.A., G.W. SIMPSON, and I.M. HALL. 1963. Impor- tance of entomogenous fungi in controlling aphids on potatoes in Northeastern Maine. Bull. Maine Agric. Expewotis Of +eapDe A9 SHANDS, W.A., G.W. SIMPSON, I.M. HALL, and C.C. GORDON. 1972. Further evaluation of entomogenous fungi as a biological control agent of aphid control in Northeastern Maine. Life Sci. Agric. Maine Exp. Stn. Tech ebull woo. eeoceDp. THAXTER, R. 1888. The Entomophthoreae of the United States. Mem. Boston Soc. Nat. Hist. 4: 133-201. WATERHOUSE, G.M. 1975. Key to the species Entomophthora EreS.eDUL em BrLUeMYyCOlmsno0C.1 Oo 4-41. ZIMMERMANN, G. 1978. Zur Biologie, Untersuchungsmethodik und Bestimmung von Entomophthoraceen (Phycomycetes: Entomophthorales) an Blattlausen. Z. angew. Ent. aan | Jad kara ay MYCOTAXON VOU Nee None lem pp.) 00-58 April-June 1981 NEW CICADA PATHOGENS: MASSOSPORA CICADETTAE FROM AUSTRALIA AND MASSOSPORA PAHARIAE FROM AFGHANISTAN* RICHARD S. SOPER USDA, SHAZAR Insect Pathology Research Unit Boyce Thompson Institute at Cornell Tower Road Ithaca, New York 14853 INTRODUCTION The genus Massospora was described by Peck (1879) and emended by Soper (1974). Forbes (1888) and Thaxter (1888) working independently, correctly placed this genus in the Entomophthoraceae. Species of Massospora are pathogenic to cicadas and thus far are found only in adults. These fungi attack the reproductive organs of their host. As the fungus grows, the terminal abdominal segments of the host slough off revealing either conidia or resting spores. Both stages do not normally occur in the same individual. The infected cicadas remain alive and take an active part in transmission of conidia to other adult cicadas or in distribution of the resting spores (Soper et al. 1976a). Only one species, Massos- pora levtspora Soper, has received extensive epizootio- logical study (Soper et al. 1976b). From this investi- gation and the observations of White et al. (1979) on * Partial support for this project was provided by the United States/Australian Cooperative Science Program under travel authorization 2-AS-11 of NSF Contract INT TR a’ YN OT ou Massospora ctcadina Peck, it is apparent that these patho- gens play an important role in the population dynamics of their hosts. Except for one report of Massospora, probably erron- eously identified as M. ctcaditna, from the Japanese cicada Platypleura kaempfert F. (Kobayasi 1951), these fungi were known only in the Western Hemisphere. This paper des- cribes two new Massospora species which indicates a much wider distribution by the addition of Afghanistan and Australian localities. This brings the total of known Massospora species to 13. MASSOSPORA CICADETTAE Soper, sp. nov. CONIDIA ochroleuca in massa, ellipsoideis, papilla gndistincta, parietibus laevibus, 7.3-9.8 x 20.7-28.1- Um jo70er 0.8 x 24.7 + 2.1 Um, & +s) binucleata vel inter— dum trinucleata vel uninucleata, nucleis nunquam bipolari- bus autem ad partem latissimum conidii locatis. SPORAE PERDURES pallida vitellina in massa, reticulis profundis cameras distinctas formantes, cristis angustis laevibus vel raro minute papillatis, 26.8 - 38.7 Um in diam. (OG e ae thaes Tenn oe Gane ee- CONIDIA pale brownish yellow in mass, ellipsoid, Wetheaneindistinct, papilla, smooth-walled, /.3 = 9.9) x Bm moe olen (9.0 +05 8ixr 24,7 492, 1 tim, x ts), bi- nucleate or ocasionally tri- or uninucleate, nuclei located at the broadest part of the conidium, never bi- polar. RESTING SPORES pale egg-yolk yellow in mass, with deep reticulations forming distinct chambers, the narrow ridges smooth or rarely minutely papillate, 26.8 - 38.7 Lm Pmeitamn (5 ).2e oo tim, x tis). (Fig. 1) Holotype: AUSTRALIA: New South Wales: 16 km east of Hay: Growing in the abdomen of an adult cicada, Crcadetta murraytensts Distant, Det. M.S. Moulds. Conidial stage. ieDecembers]979, Coll, M.S) and Bb... Moulds. | CUP: Paratypes: Same data as holotype. Resting spores present. CUP. AUSTRALIA: Queensland: Goomeri: near Kinbombi Falls. Growing in abdomen of Ctcadetta murray- tensts, Det. M.S. Moulds. Resting spore stage present. WOeDecember 1976.8 Coll. M.S. and Bus. Moulds. ~ CUP. Queensland: Aloomba: Growing in abdomen of Cticadetta puer (Walker), Det. K.J. Chandler. Conidial stage ay Figure 1. Spore stages of M. ecteadettae: (A) and (B) scanning electron micrographs of resting Spores; (C) resting spore as seen through light microscope; and (D) conidia. 20 present. 10 February 1973, Coll. K.J. Chandler. CUP. Tas- mania: Davenport. Growing in the abdomen of Cicadetta sp., Conidia and resting spore stages present. No collec- tion data. South Australia Museum. Name: The species name of this Massospora is based on the generic name of its host, Ctcadetta. Host: Cticadetta murraytensts, Cicadetta puer, and Citeadetta species (Cicadidae). This species has the largest conidia yet described (9.0 x 24.7 um) for Massospora. Although Massospora dor- tstanae Soper measure 10.8 x 21.8 Um thus approaching these dimensions, the spore surface is verruculose as opposed to smooth in M. ctcadettae. The nuclei are dis- tinctly bipolar in M. dortstanae, and although M. ctca- dettae is likewise generally binucleate, the nuclei are positioned centrally. The size and ornamentation of resting spores (M. ctcadettae) are similar to the Chilean species Massospora tetttgatts Soper which measures 36.3 Um in diameter. In the absence of conidial characteris- tics these species can be separated by the general lack of papillae on the reticulation of M. ctcadettae resting Spores as compared with numerous minute papillae found on M. tetttgatts resting spores. The location (Australia) and host (Crcadetta sp.) are strong circumstantial sup- port of M. ctcadettae when only resting spores are present. The host, C. puer, is a serious pest on sugar cane in northern Queensland, Australia. Although most cicadas hnaveavery longslite cycles, viz., 4 to 1/7 years, C. puer is thought to complete its cycle in one year (K.J. Chand- ler personal communication). Potentially, M. ctcadettae could have a correspondingly short life cycle, which would be the shortest known for any Massospora species. Intensive epizootiological studies on the periodical cicadas, Magictcada spp. indicate M. ctcadina does not occur in the nymphal stages of the host nor in other genera of cicadas. This indicates M. ctcadina must re- main in the resting spore state for 17 years between the synchronized emergence of the adults (Soper et al. 1976a). A hiatus of 9 years between the occurence of M. levtspora resting spore production in an isolated population of Okanagana rimosa (Say) was given as evidence of a corres- pondingly long survival of M. levtspora (Soper et al. 54 1976b). These observations strongly suggest that the life cycles of Massospora species are controlled in some way by those of their hosts rather than by timing mechanisms intrinsic to the fungi themselves. MASSOSPORA PAHARIAEH Soper, sp. nov. CONIDIA ignotae. SPORAE PERDURES cinnamomeae in massa, reticulis profundis irregularibus indistinctae, cameras indistinctas formantes, cristis late separatis aneustisspapilatais se32.0 — 44 q/epimeiny cL en Oc 2 OM in, Xp S)". CONIDIA unknown. RESTING SPORES dark red-brown in mass, with deep, irregular to indistinct reticulations, ridges widely separated, narrow, bearing numerous minute papillae s2 68 — 44 e/ Um ane diam .a(39.95 ct 422 6 Um, aceon Gi Tomer) Figure 2. Scanning electron micrographs of M. pahartae resting spores, (A) and (B) note the many papillae on the reticulations. 20 Holotype: AFGHANISTAN: Paghman: Growing in the abdomen of an adult cicada, Paharta casyapae (Distant), collected on "shade and fruit trees", Det. R. Froeschner. Resting spores present. 3 July 1963, Coll. E.R. Millet AF 4 68, 23-14756. CUP. Name: The species name of this Massospora is based on the generic name of its host, Faharia. Host: Homoptera: Paharta casyapae (Cicadidae). The resting spores of this species are similar to Massospora levispora Soper in their ornamentation (Soper, 1963, 1974). They can be differentiated by their average larger size 39.5 Um vs. 34.0 Um and fewer reticulations. In mass M. levtspora resting spores are yellowish brown as opposed to the cinnamon brown of M. pahartae. SNOT Ir Care senO BoRECLES The following key is a revised version of the origi- nal (Soper 1974). To identify a species the key can be entered at any point. As additional characters are exam- ined, a unique combination will be found for each species. If only the resting spore is present, it will be necessary to utilize scanning electron microscopy. Complete des-— scriptions of species 1-11 are given by Soper (1974). Species numbers are underlined when the characteristic is unique. 1. Massospora ctcadina 5. Massospora ocypetes Peck Soper 2. Massospora sptnosa 6. Massospora tettigatts Cifera, Machado & Vital Soper’ 3. Massospora levtspora 7. Massospora cartnetae Soper Soper’ 4. Massospora dortstanae 8. Massospora diminuta Soper* Soper * Several orthographic errors were made in the original descriptions with respect to Latin endings. These have since been corrected in the Index of Fungi (1975. 4:312). The corrected spellings are used here. 56 9. Massospora platypedtae 2s Soper 10. Massospora dtceroproctae 13. Soper* 11. Massospora fidtctnae Soper CONIDIAL CH 1-1 Contdtal shape 1-3 aoe UlLipsoidalieZ, 354 339 5 10h, DAs Doc use form aL c. globose 7 d. navicular 4 e. obovate 4, 10 ime VOA- Cle m2 du se se Oy ag Lal g. subglobose 11, 7 1-5 1-2 Number of nuclet ae One 33en) lye 2 b. two is ue oN 4, Oe le Ue LOR Cet hirecw sem 0, el? d., £our or more 6 RESTING SPORE CH 2-1 Ornamentatton of resting spore reticulum absent a. papillae 2) b. papillae minute 6 ee opis ae COUNGEe Cine EEO s dy 16) de Ppapwulaercruncateg2go.s Ut 2-2 Ridges of rettculum a. broad forming small chambers b. irregular forming indistinct ok be Massospora ctcadettae Soper Massospora pahartae Soper ARACTERS Contdtal wall ormamentatton a. -abSen Gyo yu Ossso mes De AVGLLUCOSEGs lyse sag, c. verruculose 4, StL O gael: 1-4 Arrangement of nuclet ae bipolare2 4 man random 1, 3, 6, 7, 12 Contdtal length aa less’ thangs um. be, 8 to Ol mee een Tra oe sLORtowitbe mise eo aera Cen, ee ely 1G ih de U5 se tom2 Or tin eee eee 5a 6 saat @. more, thane2O. lime? sao ae eh, ARACTERS 2-3 Resting spore diameter a. less than 25) Um"3 be eZ Seto eS 0e immo eo st Onmmo 2 Cr 30 ECOR4S) Uimerlt geome mer Byecaiky Ais 1 d. more than 45 Um 1 6 chambers 3, 13 narrow forming distinct chambers 1, 2, 5, 8, 11, 12 Sy HOST 3-1 Genus of eteada attacked Het scarinetdsy. g. Magtetcada 1 De Cicada? <8: h. Okanagana 3 ec. Cteadetta 12 Bh Peak romeem ANS) d. Dtceroprocta 10 ieee OuU ped ial. PeeOresrvanay4. —)) k. Quesada 2 hel) peddemenber’ gos Ie 1. Tetttgates 6 DISTRIBUTION 4-1 Collection localttties Afghanistan 13 Honduras 11 Argentina 5, 7 MeEXTCO e2,pto pL Australia 12 (ees, SHEE) iy Gy Bs Ee spel war dh, Phy AN tL 1G) Venezuela 2 ACKNOWLEDGEMENT S Dr. R. Froeschner, U.S. National Museum, Washington, DC, Mr. M. Moulds, Sidney, Australia, and Mr. K. Chandler, Cairns, Australia, provided the identification of the cicadas. Dr. R. Humber, USDA, SEA-AR Insect Pathology Research Unit, Ithaca, New York, provided the Latin trans- lation of the species descriptions and constructive criti-~ cism of the manuscript. Ms. Myra Libott, Cornell Univer- sity, Ithaca, provided technical support for SEM photo- micrography. Their help is greatly appreciated. LITERATURE CITED Forbes, S.A. 1888. On the present state of our knowledge concerning insect disease. Psyche 5: 3. Kobaysai.e)4-01951'. Notes of fungi 2 (1) 9on the newly found genus Massospora from Japan. J. Jap. Bot. 26: 21-23. Peck, D-H. 1879. Report of the botanist. NY State Mus. Nat. History 3lst Ann. Rept. pp. 19-44. Soper, R.S. 1963. Massospora levispora, a new species of fungus pathogenic to the cicada, Okanagana rimosa. Canepn lev bot wae ee oi =/ O- Soper, R.S. 1974. The genus Massospora entomopathogenic for cicadas. Part I. Taxonomy of the genus 58 Massospora. Mycotaxon 1: 13-40. SOper whe oe eA wae DeLyzer, and) l,f Re) omttlogets ,Oa mec genus Massospora entomopathogenic for cicadas. Part II. Biology of Massospora levispora and its host Okanagana rimosa with notes on Massospora cteadtina on the periodical cicada. Ann. Entomol. Soc. Am. (0958 60-957 SOper. Rh OeR Leh oh momitlinw and Awd. DeLyZer ess La AOD. Epizootiology of Massospora levtspora in an isolated population of Okanagana rimosa. Ann. Entomol. Soc. Am. 69: 275-283. Thaxter, R. 1988. The Entomophthoreae of the United States. Memoirs Boston Soc. Nat. Hist. 4: 133-201. White ad. Mi Lloyd sand. H.62ar.e (19/9 Pacihoveeatoc sion in crowded suburban periodical cicadas: Popu- Patlonussout otecontrol. BEcoltogygoU-mus05— 315. MYCOTAXON VOISE ALIA NOY Lee pp. 59-84 April-June 1981 LEOTIACEAE I11. NOTES ON SELECTED TEMPERATE SPECIES REFERRED TO HELOTIUM AND HYMENOSCYPHUS KENT P. DUMONT The New York Botanical Garden, Bronx, New York 10458 SUMMARY Fourteen temperate species of Helottum and Hymenoscyphus were studied and their taxonomic placement discussed. The accepted species are redescribed and illustrated. .Of the fourteen, six were shown to be good species of Aymenoscyphus, four were demonstrated to be taxonomic synonyms of other spe- cies of Hymenoscyphus, one additional species is a probable synonym, and three were shown to be members of the Sclerotini- aceae. As was recently pointed out by Carpenter (4981), and by Du- montrGl980: 1981) ithe identification of the majority of the mMeOtnopicainspecies Ot inoperculateyDiscomycetesmis ditiivcult Bic matucimes., frustrating dues tosthe lackijof, comprehensive and updated monographs and floristic studies. Dumont (1980) pointed out, in his study of Helottum rufo-corneum, that even Biewmost common species are still datficult to name conmectly because of the lack of adequate literature descriptions and because of existing, confusing synonymies. We have been attempting to work out the names of some of the more commonly encountered species of Inoperculate Discomy- meces, witch we have collected (Carpenter G Dumont, 19738;. Car- penter, 1981; Dumont § Carpenter, 1981; Dumont, 1980; and Haines, 1980) from neotropical regions. Dumont § Carpenter (1981) reported the difficulty they encountered while attempt- ing to identify their own field collections of the Helottum- Hymenoscyphus complex and Helottum-like organisms from Colom- bia and adjacent regions and pointed out the.need sto ireinves-= tigate all of the species of that genus described from the neotropics. Dumont (1981) presented a preliminary summary of all of the species of Helottum-Hymenoscyphus reported from the neotropics. In that study many species were actually excluded from the genus, placed into synonymy, or redistributed to gen- Pramin the, Sclerotiniaceae. While attempting to work out some of the complicated synon- ymies, such as found in Helottium rufo-corneum, it became nec- essary to study the literature and specimens of several spe- 60 cies of temperate members of Helottum-Hymenoscyphus. The pur- posewin the paper aise to present somer or thesintormacionerc. cently uncovered while studying the types of several temperate species described originally as Helotium or placed in Helo- ttum, in hopes that this will add to the growing information on the genus Hymenoscyphus. AS. 1 ebecome more, familian wWithvand study saddi tional speeres of Helottum-Hymenoscyphus, I am beginning to see the emergence Of tWORGTS tinct oY OUPINGSLOF SPeCles MIN tie) Cen sr game Ne mmencts fer to as the Hymenoscyphus caudatus group (which includes the type species H. fructigenus) and the second the Hymenoseyphus epupnyylus vroup. in the first group, the apotheoe tamarems tip itate, and the “sterile tissue (outer ectal excipulum) from the base-=of the stipe to the margin 1s composed ofva weligduevcl- oped and well defined textura porrecta to textura prismatica. Inethe: sf. epvonytlus croup, the apothecia are generaliyvetuyp ius nates to SsuDStinitate, andl the, outermost» trssuc sor ethemetlpe and frequently the lower portion of the receptacle adjoining the stipe as composed of a well to poorly -detined textura elobulosav to texturasanctlaris.) elt assprobebl es thauecthnesemtwe distinct estructura leditsmerencesaarcewor tiy -Oferecognl G10 9c oe ther a separate genus for the #. epiphyliuse group, Or am an- frageneric, rank might .\bevapplicable. Since, the majorityeort the species of Helottum-Hymenoscyphus still remain to be exam- inedyand redescribed, 2t would be premature to, makesaddattrona modi caLvOnsSs auetnis mOOi1n Gt. Thesaccompanyinge taxonomic key 1ncludessnote only thnesspce Giesmstudiedsin tthe present work’ butpencorporaves sa limormerlic species which) have previously studired(DumonteGy Carpenter, 1981; and Dumont, 1980) and accépt-as valid species of kymeno— scypnus. « vnerkey shouldinotebesthoughb or as @etini tue but one to which additional "species should bpemadded atvers they are studied. The methods used are the same as those reported bye Dumon tartans. 1. Key to species of Hymenoscyphus studied 1. Apothecia subsessile, substipitate to turbinate; stipe (or substipe) and base of receptacle of apothecium to the outside composed of tex- tura anoularisvor ¢lobulosas =e 2). (Hymenoseyphus eptphyllus group)2. 2. Ascospores 28-42x3-5um, fusoid, anterior end pointed, posterior ends aErenuiaredy ones nt Moy. seat. eae ee ee eee H. dearnesstt 7p aoc. eee ASCOSPOLeSH lesSe thane 2oningel ON 0 9 wee eae re anes ne ee eee re 3. Ascospores (7-)8-10(-12)x2.5-3.5(-4.5)um, trapezoidal, obo- void, generally equilateral, if inequilateral then not flat- tened on Ones suri ace we. cin .esere eee ener H. immttabults pe 73i 3. Ascospores more than el 2m slong aa cse cect ath oe. te a eee eee Al 4. Apothecia occurring on wood, strongly umbilicate; asco- spores (15~) 15-18 (=20)x(3-) 4—-5im)) oe es H. umbtltecatus (See Dumont, 1980) 4. Apothecia occurring on leaves, turbinate, not umbilicate; ascospores, 15218x5.5=5imae ee eee H. eptphyllus p. 66. (See Dumont, 1980) 1. Apothecia stipitate; stipe (or substipe, if present) composed of tex- tura porrecta to textura prismatica (if any globose cells present in Stipe or receptacle, use first choice) oe eve ee 61 MNES Ta, Oeics tad teat Bp oa (Hymenoscyphus caudatus group)5. 5. Ascospores equilateral or if inequilateral then not flattened, Peleval | Varuale2 O1Gat MCORODOVOLGLOLODDYL LOT... esc taieete ees 6: 6. Apothecia characteristically tany, less than Jmm in diam, dis- coid; ascospores (10-)11-14(-15)x(3-)4(-5)um on leaves; tem- DEVAL Cour teas esha A abbey aBhog dy be ony aye tet dress H. translucens p. 80. 6. Apothecia characteristically large, greater than 2mm in dian, irregular in outline, umbilicate; ascospores (4-)5-7(-9)x1.5-2 Poe PUT RIOR SE OE ry ei eee Re ee ecg ee eal a H. leucopsts (See Dumont, 1980) 5. Ascospores strongly inequilateral, a high proportion flattened..7. 7. Ascospores with a large 'nuclear staining area" visible in phloxine, cotton blue and analine blue dyes, rarely l-septate, (22-) 26-30(-35)x4-5(-6)um, ascus apex papillate; tropical..... Bee eal Shag 5 Fe ent ak he Males Va Ma hres UE Ae PRN Rae ey. shisn ne Cost aie H. sclerogenus (See Dumont & Carpenter, 1981) WN SCOSPOLCSUW1ILNOUD NUCl Cares tAINING Area oon car alaeis occ ele te-0, din 8. 8. Ascospores regularly septate (more than 50% in any amount) Spe Meee Re Meh Sie aay Roce ote sao eet hats 16 ie het chars ¢ Weatens fel gue ‘ape esgueia lore 9. 9. Ascospores l-septate, (17-)18-22(-24)x4-5(-6)um; apo- checidewitnouc nairseat, thesbase, ol thesstape. i.e... ee eee ict ML SOs a) OE ce aoe dab. Cie ayelaces H. mustcola (See Dumont, 1980) 9. Ascospores 3-septate, (20-)24-30(-35)x4-5(-6)um; apo- Eheclomwltlenairsea tetlcaDascnOPetiicsS OLD .9,1 ero. 4 asi SLO Pa Say I Reh. 3 cag tS OOO Pg a A. Peedine (See Dumont & Carpenter, 1981) SrA S COSPOLRESEASOD CALE tg aha cate aie ci erred site aie resis > igvarnate é 102 10. Ascospores (16-)18-22(-26)x2-3um, with a basal cilium Eee PO, Le ate OM ee re Ree Foam ohh wt PE SCULILUS (See Dumont & Carpenter, 1981) LORASCOSPOTCS WItnOUtl a1 Dasa lCi LaUMies os sine ce ee oe 1% Vie enSCOSPOLesenoOked Japl Cally rae cas ties crete: Vze 12. Ascospores (16-)18-23(-30)x3-3.5(-4.5)um, strongly and obviously hooked, tapering evaduad 1 yetOethe, baser, ates H. serottnus (See Dumont’ G Carpenter, 1981} 12. Ascospores (14-)16-23(-26)x4-5(-6)um, if hooked, only slightly and a few per mount, abruptly pointed at basal end...H. caudatus (See Dumont & Carpenter, 1981) TIP ASCOSDOLeS nO tRnOOKeCdn aD LCA LEY si tlre acre el raN 13. Ascospores (11-)12-15(-16)x(2.5-)3-4um, with internal, oily-resinous material...... (See Dumont, 1980) 13. Ascospores without internal resinous, oily MATCTIA IN aimee auch sete ence ate ai ate add. 14. 14. Ascospores frequently curved and then indented, with anterior end pointed; apothecial margin with a gelatinous matrix present; ascospores (10-)12-16 (=18)x3-4(-S)pm-,.f. erraticus ps “66. 14. Ascospores if curved, then not indent- ed, with anterior end rounded; apothe- cial margin lacking a gelatinous ma- trix; ascospores (14-)16-23(-26)x4-5 62 EXsaptin Poe eee Aa Tee My ek a eC Ree Bete: POG ene 6 © H. caudatus (See Dumont §& Carpenter, 1981) 1. Helottum albopunetatum Peck, Annual Rep. New York State MUNG Hat oeLe me 4i/e eee O eos = Hymenoscyphus albopunetatus (Peck) Kuntze, Revis. gen. Pl. 3(3): 40010 91896. NOTES. Helottum albopunetatum was described from New York by Peck (1879) as occurring on leaves, and was fully rede- scribed and illustrated by White (1943), who accépted™=its placement in Helotium. It was transferred to Hymenoscyphus by Kuntze (1898) and accepted in that genus by both Dennis (1964) — and Arendholz (1979). After having studied the type deposited ine NYS, I conclude that the species is the same (as fyumenocsey— phus caudatus, and I place it into synonymy and adopt the name Hymenoseyphus caudatus which has priority. This species fits well within my concept of #. caudatus as presented in Dumont §& Garpente rms Lie | Holotype: U.S.A., New York Adirondack Mts., C. HY Peck (NYS, NY): 2. Helottum caudatum (Karsten) Velenovsky, Monogr. Discom. BOHen etic UO.e 1954" Peztza caudata Karsten, Fungi fenn. exs. 547. 1866. Hymenoscyphus caudatus (Karsten) Dennis, Persoonia 3: 76. 1964. NOTES. Hetlottum caudatum was originally described by Kar- | Stenecromea toliicolous collection made 1n,) Panlandacs peersed | very common temperate and neotropical species. Dennis (1964) jae Arendholz (1979), and Dumont. & Carpenter (1981) accept’ the | species in Fymenoscyphus, a decision with which I concur.” Fou a full description, illustrations, and discussion, see Dumone & Carpenter (1981). 3. Helottum conocarpt Seaver §& Waterston, Mycologia 34: 517. 1942. NOTES. Helottum conocarpt was originally described from Bermuda on leaves. It was accepted in Helottum by White (1943), but neither Dennis (1964) nor Arendholz (1979) treated the species. Dumont (1976) has shown the species to be a mem- ber of the genus Moellerodtseus (Sclerotiniaceae). See Dumont (1976) Bloretulivdéescription.!) pilustrationsmandedaccusstone 4. Helottum dearnesstt (Ellis §& Everhart) White, Mycologia ote AR Te FIGS. 612 am = Phtalea dearnesstt Ellis §& Everhart, Proc. Adad. Nat. Sci. Philadel- phia 1893: 146. 1894. = Hymenoseyphus dearnesstt (Ellis §& Everhart) Kuntze, Revis. gen. Pl. 3(3): 485. 1898. Apothecial morphology — Apothecia minute, scattered or found in small groups, subsessile, to ca lmm in diam, ca 1mm high, when fresh disc drying concave, rehydrating convex. Hymenium when fresh at first subolivaceous, 63 FIG. 1. Hymenoseyphus dearnessit, holotype ex NY, freehand drawings, x 1,000. A. Median longitudinal section of an apothecium through margin. B. An ascus and a paraphysis. C. 6 ascospores drawn after discharge from ascus. becoming yellow, drying flesh-colored to ochraceous, rehydrating lighter and yellowish; receptacle generally concolorous with hymenium when fresh, dry and rehydrated; stipe coloration difficult to detect owing to small size, but appearing dark brown. Apothecial anatomy — Asci 8-spored, 90-110x9-12yum, croziers reported by White to be absent, presence or absence uncertain owing to poor stain- ing of the asci, but possibly present, long-cylindric to clavate, taper- ing towards the base and there generally not becoming expanded to form a small foot, walls ca lym thick, becoming enlarged at the subpapillate to papillate apex and there 3-4ym thick; pore J+, the pore walls staining in- tensely blue in Melzer's Reagent. Ascospores 28-42x3-5ym, details ob- scured in all collections owing to failure of spores to absorb stain ade- 64 FIG. 2. Hymenoscyphus dearnesstt, holotype ex NY, freehand drawings. A. Median longitudinal section of an apothecium at approximately juncture of stipe and receptacle, x 1,000. B. Habit sketch of an apothecium on the@ Substrates xeca 50" quately, biseriate to irregularly uniseriate, hyaline, smooth, aseptate, elongate, fusoid, anterior end pointed, posterior end attenuated and drawn out to a fine point, a cilium reported by White not observed in these studies, in outline equilateral, frequently sigmoid and curved, gut- | tules spherical to irregular in outline, irregularly dispersed in the in- dividual ascospores and generally filling major portion of ascospores and separated by narrow bands of cytoplasm. Paraphyses equal to or slightly exceeding the asci, internally hyaline, occasionally branching towards the base of the asci, septate, filiform, becoming slightly expanded at the apex and there 2-4ym wide, walls thin, smooth and hyaline, Subhymenium absent, without a defined zone or differentiation of tissue beneath the asci. Medullary excipulum poorly developed, in the flanks consisting of narrow, parallel hyphae 2-3yum wide, in the center of the receptacle the 65 parallel hyphae becoming torn apart and interwoven, the individual hyphae hyaline with walls thickened, refractive and smooth. Ectal excipulum: in- ner ectal excipulum absent. Outer ectal excipulum non-gelatinized, re- fractive due to thickened walls of hyphae, to ca 40um wide towards the margin and to ca 60ym towards the stipe, consisting of hyphae originating in the stipe, and at the juncture of stipe and receptacle extending almost perpendicularly to the surface of the apothecium, continuing towards the margin the individual hyphae extending to the surface at gradually lower angles until at the margin the hyphae parallel to each other and to the surface of the apothecium, the individual cells brick-shaped, (6-)8-12 (-20)x4-8um, the walls thickened, hyaline and smooth, at the surface of the apothecium the apically free cells unmodified and remaining appressed to the surface. Outer covering layer absent. Hairs absent. Margin nar- row, consisting of hyphae originating in the receptacle and extending parallel to the surface, as described for the receptacle, but less refrac- tive. Stipe to the outside composed of a pigmented zone of irregularly arranged hyphae, in some areas the individual hyphae losing hyphal orien- tation, and appearing cellular in composition, the individual cells brick-shaped, globose or angular, lightly to intensely pigmented, the walls thickened, non-gelatinized and obviously roughened; inside the pig- mented zone, the hyphae narrow, 2-3um wide, extending parallel to each other and to the surface of the apothecium and continuing into the medul- lary excipulum of the receptacle, the individual cells hyaline or pig- mented light brown, the walls slightly thickened, lightly pigmented or hyaline and generally smooth. Hairs absent. Habitat: On old stems of Stetronema ctltatum. Etymology of the specific epithet: refers to the collector of the ori- ginal and type specimen. Holotype: Canada, London, dead stems of Monarda, May 1980, J. Dearness 1713 (FH). See additional notes below. Tilustrations: White, Farlowia 1: 615, figs. 31-34. 1944.- Dennis, Per- Boonia 5: 41, fig. 16. 1964. NOTES. Phialea déarnessti was described by Ellis | Ever- Marterromea collection made by J. Dearness (1713) in Canada. As White (1944) has pointed out the type collection was made in May 1890; and the host was originally described as Monarda, but Dearness apparently returned to the same locality in June and collected additional material of the new species and con- cluded that the host was not Monarda, but Stetronema ciltatum. To add to the confusion, Dearness gave the new collection the same number as the original material, but fortunately they both do represent the same species. Kuntze (1898) transferred Phtalea dearnessit to Hymenoscy- phus; White (1944) placed it in Helottum. Dennis (1964) ac- cepted its placement in Hymenoscyphus, a decision with which I concur. The species appears to be most clearly related to other substipitate species in the genus, such as H. eptphyl- lus. In these species the stipe is composed of hyphae which do not form a well-structured textura prismatica or porrecta, as in species such as Hymenoscyphus caudatus and its related species, but the hyphae become disoriented with the individual cells losing hyphal orientation and also losing the layered effect. In H. eptphyllus, H. timmutible and Helottum mtdlan- 66 dense the outermost tissue of the stipe toward the receptacle is formed of textura globulosa to textura angularis, Further, | in these species, the ectal excipulum towards the stipe is formed of hyphae originating in the interior of the stipe and extending nearly perpendicularly or at very high angles to the surface of the apothecium. From the stipe to the margin, the hyphae approach the surface at lower angles until at the mar- gin the hyphae are parallel to the surface. In the #. cauda- tus group the ectal excipulum is formed of hyphae originating in the stipe and running parallel (or at a very low angle) to the surface from the stipe to the margin. Hymenoscyphus dear- | nesstt is separated from these other species by the size of LoS HasGOSDOTes.. White (1944), in his description of H. dearnessit, men- tioned that the ascospores produced a delicate cialium 2:-5-3-5 um long. In the portion of the’ type which i -studicd = tiemace cospores only poorly absorbed the various stains utilized; anid, for this reason, I was unable oto observe certatuercacurcs of the anatomy of the species. I did not observe the cilium described by White, but agree with him that the ascospores do become obviously attenuated. 5. Helottum teptphytlum (Persoon ex Persoon)) Fr. > oummanves. Seand Det 500. O49 Peztza eptphylla Persoon, Ann. Bot. (Usteri) 11: 30. 1794. Peztza eptphylla Persoon, Mycol. Europ. 1: 295. 1822. Ht out wi MicheeAcad . S¢i a9 :) Lime 19292 NOTES. Helottum eptphyllum is a very widely distributed species in temperate regions. As mentioned by Dumont (1981), the presumed type of Hymenoscyphus eptphyllus deposited at L contains only sclerotia) of ‘an unidentified fungus 9) UnGileine situation of the type of the species is resolved, I am follow-@ ing the concept of the species as set forth by Dennis (1956) and Arendholz (1979), both of whom illustrate the species. Dumont E1981) shas ftullysdescribedjand) 11 lustratedgautropres. variant of the species. 6. Helotium erraticum White, Farlowia 1: 606. 1944. FIG. 3. Hymenoseyphus eptphyllus (Persoon ex Persoon) Rehm ex Kaufmann, Pap. Apothecial morphology — Apothecia scattered or rarely gregarious, gen- erally produced from leaf blades, small, stipitate, 0.5-1.0mm in diam, to ca 1.0mm high, when fresh cupulate in youth, disc flat with age. Hymenium when fresh white to off-white, drying bright yellow, dull yellow, ochrace- ous, or rarely reddish; rehydrating translucent, pallid to yellow; margin generally concolorous with margin when fresh, dry and rehydrated; recepta- cle when fresh white, drying slightly lighter than hymenium or concolor- ous, yellow to occasionally reddish, rehydrating as hymenium or slightly lighter; stipe generally concolorous with the receptacle when fresh, dry or rehydrated. Apothecial anatomy — Asci 8-spored, 85-110x8-llym, produced from small — replicating croziers, broadly cylindric to long cylindric-clavate, taper- ing to the base and there becoming slightly expanded to form a small foot, wall to ca lum thick, enlarged to 2-3ym at the rounded to truncate apex; ' A ) 1A | ' LAS NAN : \ is \ A N i} 68 pore J+ in Melzer's Reagent. Ascospores (10-)12-16(-18)x3-4(-5)um, bi- seriate above and uniseriate below, biseriate throughout or less commonly obliquely uniseriate, hyaline, smooth, aseptate or rarely l-septate (and then apparently in poorly preserved material), oblong, obovoid, ends rounded to pointed, in outline inequilateral, flattened on one side and generally curved and then slipper-shaped, generally anterior end slightly broader than posterior end, guttules present, more or less equal, irregu- lar in outline, bipolar, filling about half the spore at each end. Para- physes equal to or rarely slightly exceeding the asci, rarely branched near the base of the asci, sparingly septate, internally hyaline, f1ili- form, becoming slightly expanded at the apex and there 2-4ym wide, walls thin, hyaline and smooth. Subhymenium not well differentiated from the medullary excipulum, with a perpendicularly oriented zone to ca 40ym in the center staining more intensely than the remainder of the receptacle and composed of parallel, short, broad hyphae and croziers, the individual cells 2-4(-5)um wide, the walls thin, hyaline and smooth, these hyphae tightly compact and grading into the upper portion of the medullary excip- ulum. Medullary excipulum poorly to well developed, hyaline, obconical, non-refractive, consisting in the center of loosely interwoven, branched, septate hyphae, towards the flank and subhymenium becoming more lightly compact and parallel to subparallel, the individual hyphae 2-5um broad, the walls thin, hyaline and smooth. Ectal excipulum not divided into rec- ognizable layers, to the outside the hyphae more tightly compact and nar- rower than the hyphae in the center of the excipulum; in this region forming a well developed textura porrecta to prismatica, the individual hyphae hyaline, the walls thickened, hyaline and smooth; these hyphae grading toward and into the narrow hyphae of the medullary excipulun. Margin at its base constructed as the upper portionon the flanks out soon becoming characteristically gelatinized, the spaces between the indi- vidual hyphae 1-2(-3)um, the apical cells unmodified. Stipe composed as the flanks, the individual hyphae larger than comparable cells in the flanks. Hairs absent from the margin, flanks, and stipe. Habitat: On leaf blades of unidentified species, leaves of Amelanchter sp., Ulmus sp., Hamamelits sp. and Populus sp., and last year's pods of Ro- binta pseudo-acacta. Etymology of the specific epithet: relevance to the fungus uncertain. Holotype: U.S.A., New York, Coy Glen, near Ithaca, on decaying leaves, 3 Oct 1938, H.sHe Whetzel” G.We bk. White sen. “(White herbenos 25472 leanda 3416, one collection apparently divided into two packets or two specimens. collected on the same day with identical data.) Paratype specimens: U.S.A., New York, Malloryville, W. of North Bog, on leaves of Amelanchter sp., Ulmus sp., and Hamamelis sp., 18 Oct 1941, H. H. Whetzel & Niederhauser s.n. (FH ex CUP 29658); Newfield Gorge, Itha- ca, last year's pods of Robinta pseudo-acacta, 26 Oct 1941, H. H. Whetzel & T. Sproston s.n. (FH ex CUP 29666); Canada, Quebec, Duchesnay, County Portneuf, on leaves of Populus sp., 25 Aug 1938, H. H, Whetzel §& T. Spros- tones-n se s(hH exCGUP: 27850). Illustrations: White, Farlowia 1: 607, figs. 14-18. 1944. Arendholz, Morphologisch-taxonomische Untersuchengen an blattbewohenden Ascomyceten aus der Ordnung der Helotiales, pl. 15, fig. 4, 1979. NOTES. Helottum errattcum was described by White (1944) who indicated that the species was closely related to members of the Helottum epiphyllum group: H. eptphyllum 4H. immuta- 69 bile, Hf. carpinacola; and H. mtdlandense. These latter four share in common one feature which is absent in #. erraticum. ingaliy foun the sectal excapulum:at the: juncture. of the recep- tacle and stipe is composed of globose cells to the outside, with hyphae progressing gradually from high angles to the sur- Pacem Migc hes LOWeCrepOltLOn stosparallelyat ythe margin: I.-have found no globose cells in any of the collections studied here of H. errattcum. I conclude that #. errattcum is closely related to Hymeno- seyphus caudatus. In Helottum errattcum the ascospores are wenerally ‘curved and frequently indented or slipper-shaped, while in Hymenoscyphus caudatus the ascospores are inequilat- Cral and generally not curved ‘and when curved, not slipper- snaped,,) A high proportion of the ascospores in Helotium erra- ttcum have ascospores with pointed apices, while this feature UseabscCitein Humenoscypnus caudatue. »1n #, caudatus the mar- gin 1S composed of thin-walled, brick-shaped cells, while in Helottum errattcum the margin is unusual for a member of dy- menoscyphus. The margin is composed of narrow cells apparent- iMeribeadcodalled Selacinous Matwixn, atestimes, nowever, the hy- Dpidemalcumientiy compact, and it isediiitecult to; anterpret ‘the SaMicticeemoncetiec Wa licemerely=appears thickened. ll Ones cole mectione (CUP sz 9000 lf the hyphae in the ectal excipulum are nar= rower than normal and very thick-walled, and the ascospores momitsceDtatcCam Ine collection appears cto ve Dadly preserved. Waichemaye account 1Orathe abnormal tOrmation of the ‘sterile: Piscticmangumtnemscptatloleinwtne, ascospores a All ouner tea Muresmare Cnaracterislics:0n welotrum errartecum,. The shape a eo ce Se oN Or etic SNCCL1eS. (oS 1S stne Characteristic Suphymenium i conclude sthat this is merely a’ variant collec- cron. I am somewhat concerned about the placement of this species inp hymenoscyphue, since it produces a margin with at least some hyphae in a gelatinous matrix. This feature is more characteristic of the genus Croctcreas than it is of Hymeno- seyphus, However, the ectal excipulum is that of an Hymeno- scyphus. Dennis (1964) did not treat the species, while Arendholz (1979) did place it in Hymenoscyphus as H. erratt- eus. I conclude that the species is perhaps intermediate be- tween the two, but shows more affinities to Hymenoscyphus than Lomcrocicreas. “lowil ly thus; maintain atin Aymenoecyuphust 7. Hetotium fastidtosum Peck, Annual Rep. New York State Mus, ulate gO 7 eee LiSs/) Sys Fl Gey. 4: Apothecial morphology — Apothecia scattered, stipitate, 1-1,5mm in diam, 1-1.5(-3)mm high, when fresh disc flat, drying flat to slightly cu- pulate, rehydrating convex. Hymenium when fresh off-white to pale yellow, drying darker, yellow, ochraceous to flesh-colored, rehydrating lighter, slightly translucent; receptacle when fresh generally slightly darker than hymenium and white to dark yellow, drying and rehydrating darker than hy- menium; stipe narrow, cylindrical, when fresh above concolorous with the lower portion of the receptacle, becoming darker toward the base, drying darker, especially toward the base, or rehydrating lighter and pallid, darker below. 70 Apothecial anatomy — Asci 8-spored, (75-)85-100x9-12um, produced from small croziers, long cylindric-clavate, tapering gradually to the base and there occasionally becoming slightly expanded to form a small foot, wall to ca lum thick, enlarged at the papillate apex and there 2-3ym thick; pore J+, visible as two, dark blue apical dots in Melzer's Reagent. Asco- spores (23-)26-30(-35)x2.5-3.5(-4.5)ym, obliquely uniseriate to irregular- ly arranged, hyaline, smooth, aseptate, subfusoid to clavate, apical end hooked and pointed, basal end pointed, in outline inequilateral, curved and rounded in the upper portion, flattened and straight below, anterior end broader than posterior, with a single row of spherical to irregular guttules filling majority of the spores. Paraphyses equal to or slightly exceeding the asci by 5ym, internally hyaline and devoid of pigmented con- tents, branching occasionally toward the base of the asci, septate, fili- form, not expanded at the apex and there 2-3ym wide, walls thin, smooth and hyaline. Subhymenium not well differentiated from the medullary ex- cipulum, poorly developed, with an indistinct hyaline zone beneath the asci with hyphae more tightly compact than the medullary excipulum below. Medullary excipulum poorly developed in the flanks and in the center of the apothecium, hyaline, non-refractive, consisting of septate, branched, vertically oriented to tightly interwoven hyphae 3-6um wide, the walls thin, hyaline and smooth. Ectal excipulum: inner ectal excipulum well de- fined and well differentiated from the outer ectal excipulum and grading into the medullary excipulum, entire layer hyaline, non-refractive, to ca 10um wide toward the margin and to 25ym toward the stipe, consisting of tightly compact, hyaline hyphae 2-4ym wide, with walls thin, non-refrac- tive, hyaline and smooth. Outer ectal excipulum non-gelatinized, non-re- fractive or rarely appearing slightly refractive due to thickened walls, to ca 20um broad toward the margin and to ca 30m toward the stipe, con- sisting predominantly of a well defined textura prismatica with the indi- vidual hyphae extending parallel to the surface of the apothecium and without apically free hyphal tips; the individual cells toward the margin 8-15x4-8um, 12-25x5-8ym toward the stipe, the walls thin or rarely slight- ~ ly thickened, hyaline and smooth. Outer covering layer present or becom- ing detached and the apothecium then appearing naked, consisting of 1-2 layers 2-5ym broad, the individual hyphae extending parallel to the sur- face, overlapping, terminating before the margin, the apically free cells remaining appressed to the surface of the apothecium and unmodified, the individual cells hyaline or rarely light yellow-brown, the walls non-re- fractive, hyaline to light brown, and smooth. Hairs absent. Margin gen- erally poorly developed, narrow above, broader below, entire zone hyaline or light brown, constructed similarly to the receptacle below, the indi- vidual cells approximately same size as upper receptacle or slightly smaller, apical cells unmodified. Stipe in the upper portion constructed Similarly to the lower portion of the receptacle and apothecial flank, at approximately midpoint and to the outside 1-2 layers of narrow, hyaline or lightly pigmented hyphae (often difficult to detect) with thin, lightly to intensely pigmented and smooth or rarely roughened walls, to the inside a zone of textura prismatica with the individual cells hyaline to light brown, in the central core the hyphae narrow, parallel to slightly inter- woven; hairs absent. FIG, 4. Hymenoseyphus fasttdtosus, holotype ex NYS, freehand drawings, x 1,000. A. Median longitudinal section of an apothecium at approximately midpoint between margin and stipe. B, Median longitudinal section of an apothecium through margin. C. An ascus with 8 ascospores. D. A branching paraphysis. E. 7 ascospores drawn after discharge from ascus. - aN NAN i} ) NUSag Maes Nr Ne . an Ava | Mt iM | \ | AN he Habitat: On leaves of Alnus tneana, Alnus sp., and catkins of Alnus. Etymology of the specific epithet: relevance to the fungus uncertain. Holotype: U.S.A., New York, Forestburgh, on fallen petioles of Alnus leavyes* eocpie, (Ca Hom reckus.D.(NYS), Additional specimens examined: U.S.A., New York, Adirondack Mts., on leaves of Alnus sp., date not given, C. H. Peck s.n. (NYS); Labrador Lake, near Tully, on old overwintered leaves of Alnus ineana, 26 Aug 1935, H. Hy Whetzel & W. L. White s.n. (CUP 24817 ex FH); Lloyd Preserve, McLean; on Alnus tncana, leaves on ground, 6 Sept 1935, H. H. Whetzel, W. L. White & Rogers (White 2042 ex FH). Oregon, Tilly Jane Creek, Mt. Hood, on dead alder leaves and petioles, 26 Aug 1933, J. Kienholz K 137 (BPI ex FH). CANADA, Quebec, Duchesnay, County Portneuf, on petioles and lower midribs of Alnus leaves, 26 Aug 1938, H. H. Whetzel s.n. (FH); locality as previ- ous. collection, 24 Aug 1958.,.H. HL Whetzely sans (fA): Tilustrations: White, Mycologia 34: 1158, fig. 2; 166,, fig. (93 1942" White, Farlowia 1: 153, fig. 7. 1943. Arendholz, Morphologisch-taxono- mische Untersuchungen an blattbewohenden Ascomyceten aus der Ordnung der HelOtlales epi peo eld On eee LOO. NOTES. Helotium fasttditosum was described by Peck from the Uport.as OCGUTTING Jon, beavesvon Alnus Pe lheuspeciesmuce san om produce a stroma and is ‘surely referrable to Hymenoscyphus as reported recently by Arendholz (1979) who also reported it foy Cheetirsiatimestrom Europe, “Structurally stnessterie surssuc of the apothecium is similar to the common species Hymenoscy- phus caudatus and Hymenoscyphus serottnus, while the asco- spore shape 1S like {that of 4. serozinus = Dumont GeGarnenger (LOSI) spointed out -thateuntiie recently #.Mecrotiiucewosmecie erally regarded as /a species Occurring .onehey bac eousme tens However, they have demonstrated from their studies of neotrop- ical collections that the species jis extremely variable wand occurs on leaves as well as on stems. They further reported the ascospores to be (16-)18-23(-30)x3-3.5(€-5)um and indicatedm that the base of the stipe generally stains light pink’ in Melg zer's Reagent,. a reaction also observed in the material stud- ied here. Dennis (1964) reported the ascospore measurements for H. serottnus to be 18-28x3-4um, while Seaver reported them to be 4x22-24um & Rehm (1893 in 1887-1896) found them to be 30-36x4-6uym. The ascospores studied in the present investiga- tion of Helottum fastidtosum were (23-)26-30x2.5-3.5(-4.5)um and would thus appear to fall within the reported range for hymenoscyphus serotinus, and is a probable synonym of #. sero- tinus., I have been unable to find a-type, of #. Sserottinuc) ang am thus following the generally accepted concept as presented by Dennis (1956, 1964). A final decision on the, tentative Synonymy can be made only after type material is located or a neotype designated for H. serotinus. 8. Helottum fraternum Peck, Annual Rep. New York State Mus. Deg) AF 18:79 = Hymenoscyphus fraternus (Peck) Dennis, Persoonia 3: 76, 1964, NOTES. Helottum fraternum was originally described from New York State as occurring on leaves, and was accepted by ETS TEs acetone al ee Ses | | | ae White (1942). Dennis (1964) transferred the species to Hymen- oseyphus; Arendholz (1979) did not treat the species. Dumont (1981) redescribed and illustrated the species. He found that the species formed a well-defined substratal stroma, produced an ectal excipulum with hyphae embedded in a gelatinous matrix and transferred the species to Poeculum (Sclerotiniaceae). For peeuliedescraption, allustrations and discussion, see Dumont (1981). 9. Helottum tmmutabtle Fuckel, Jahrb. Nassauischen Vereins Na CUT Kees 52 Oe OO Om Gils FIGS.5. = Hymenoscyphus tmmutabtlts (Fuckel) Dennis, Persoonia 3: 76, 1964. Apothecial morphology — Apothecia scattered, arising from leaf blades, short stipitate, ca 1.0-1.5mm in diam, to ca 0,5-0.75mm high, when fresh flat to slightly convex, drying same, rehydrating flat to slightly cupu- late, Hymenium when fresh white, drying pale yellow, yellow-brown, or reddish orange, rehydrating slightly lighter and pallid; margin concolor- ous with hymenium; receptacle when fresh white, drying pale yellow-orange, rehydrating lighter and concolorous in the lower portion with the upper portion of the stipe; stipe in the upper portion similar to receptacle, coloration difficult to observe owing to small size, Apothecial anatomy -—- Asci (70-)80-100(-105)x(6-)8-9um, produced from small croziers, clavate to clavate-cylindric, gradually tapering toward the base and there becoming expanded to form a small foot; wall ca lum wide; pore J+. Ascospores (7-)8-10(-12)x2.5-3.5(-4.5)um, generally bise- riate, but occasionally uniseriate, hyaline, smooth, obovoid to trapezoid- al, anterior end round or rarely pointed, posterior generally slightly pointed, in outline more or less equilateral, if inequilateral only slightly so and not flattened on one side, walls slightly thickened, egut- tulate or less commonly with polar guttulate areas (which are generally obscured and difficult to detect) or in youth biguttulate with guttules disappearing with age. Paraphyses equal to or slightly exceeding the asci, internally hyaline, branching toward the base of the asci, septate, filiform, becoming slightly expanded at the apex and there 2-3ym wide, walls thin, smooth and hyaline. Subhymenium not well differentiated from the medullary excipulum, consisting of parallel, vertically oriented hy- phae grading into medullary excipulum. Medullary excipulum well devel- oped, non-refractive, hyaline to pigmented light brown, consisting of sep- tate, branched, parallel or loosely to tightly interwoven hyphae 2-4ym wide, the walls thin, non-refractive, hyaline or rarely pigmented light brown, and smooth. Ectal excipulum: inner ectal excipulum poorly defined and differentiated from medullary excipulum, entire layer non-refractive, hyaline to pigmented light brown, consisting of tightly compact, more or less parallel, hyaline hyphae 2-3um wide, the walls thin, non-refractive, hyaline and smooth. Outer ectal excipulum non-refractive, entire layer hyaline to subhyaline, variable to ca 35um broad toward the margin and the same toward the stipe, at the intersection of the stipe and the receptacle to the outside composed of thin-walled, globose cells comprising a well developed textura globulosa, to the inside the hyphae parallel, above the juncture the isodiametric cells giving way to hyphae originating in the stipe and extending almost perpendicularly or at very high angles to the surface, progressing toward the margin the hyphae extending at lower and lower angles to the surface until at the margin the hyphae parallel to the surface of the apothecium, the individual cells with walls thin to slight- 74 ly thickened, hyaline or rarely pigmented light brown and smooth. Outer covering layer as a distinct layer absent, but with an area to the outside formed of the apical cells of the hyphae extending at low to high angles to the surface and continuing to the surface, and then becoming somewhat expanded and frequently large and clavate, these cells hyaline to slight- ly yellow-brown with walls thin to slightly thickened and frequently roughened. Margin constructed similarly to the upper portion of the flank, with the individual cells smaller, the apical cells generally un- modified. Hairs absent on the receptacle and margin. Stipe in the upper portion constructed similarly to the lower portion of the receptacle, be- low to the outside the cells globose with parallel narrow hyphae to the inside; hairs absent. Habitat: Leaves of Populus tremula, P. ntgra, Quercus pedunculata, Quercus sp., Ulmus sp., Robinia pseudo-acacta. Etymology of the specific epithet: relevance to the fungus uncertain. Holotype: Germany, Boss Pr. Ebenbach, ad Popult tremulae folia putrida, Fuckel (Fuckel, Fungi rhenani no. 2388, NY probably distributed in Herbier Barbey-Boissier no. 1215.). Tilustrations: Dennis, Mycol. Pap.62:°93, fig.) 85.) 1950 me Nn ecum rans lowia 1: 143, fig. 3. 1943. Arendholz, Morphologisch-taxonomische Unter- suchengen an blattbewohenden Ascomyceten aus der Ordnung der Helotiales, Die bin etl OS me 2 ost 7 OR nopres. SAccordins to: White *(1943) “andwArendholza(lo7o 1m Helottum tmmutabtle is a widely distributed species in Europe Dennis (1964)* transferred the species to Hymenoscyphus, a de- cision with which I agree, and it appears most» closely relatea tO Helottum epiphytlum, since? both havevsimilares truccuiesor the sterile tissue of the apothecium; the outermost layers of the stipe and base of the receptacle are composed of globose cells, and the receptacle is composed of hyphae originating at the stipe and at the base extending almost perpendicularly to the surface. Further, from the stipe to the margin the hyphae extend at progressively lower angles until toward the margin the hyphae are parallel to the surface. I have studied, few collections of #.; tmmutabctleswbuts based on the shape of jthe ascospores, it appears distinets comer. eptphyllum. There is overlap in the measurements, dH. epitphyl- Zum 12-24x3-5ym (combined measurements according to Dennis, 1956, White, 1943 and Arendholz, 1979) and in #. tmmutabile they are said to be 10-13x4-5um.. It%would) appear that asco- sporen size is of little value in separating these two spe- cies. I have attempted to locate the type specimen of Peziza eptphylla in the Persoon herbarium at L, but no apothecia re- main on the leaf, and all that is present in the type are sev- eral apparently unrelated sclerotia, probably non-Sclerotinia- ceae. Thus, a final decision on the correct placement of #. FIG. 5. Hymenoseyphus immtabtilis, Fuckel, Fungi rhenani 2388 ex NY, free- hand drawings, A, Median longitudinal section through an apothecium show- ing lower portion of receptacle and margin, x 1,000, B, An ascus with 8 ascospores, C. 6 ascospores drawn after discharge from ascus, x 1,000, D. Habit sketch of an apothecium on the substrate, x ca 50, 76 eptphyllum and its relationship with H. tmmutabile can be made | only after a neotype specimen is designated for Peztza ept- Divl layed clr es eam eno Canow prepared to do. 10. Helottum lindert White, Farlowia 1: 154. 1943. vores. White described Helotium lindert from collections from. the U.S.A. Dennis. (1964) ‘didenot *tream™ themspecies, while Arendholz (1979) placed H. linderi into synonymy with Hymenoscyphus caudatus, a decision with which I concur. Holotype: U.S.A., Tenn., Chimney Trail, Great Smokey Mts. Nat. Park, 3200, ft. on fallen leaves, #18 Aug 1959.5. D. 7H slinders (En). 11. Helottum midlandense White, Farlowia 1: 605. 1944. FIG...6% Stroma — Substratal, visible on the fruits only as blackened areas of the substrate; in section the blackened areas composed of an irregularly formed rind composed of irregular to epidermoid cells in face view, also visible in cross section of the base of the stipe of the apothecium; not known in culture, Apothecial morphology — Apothecia solitary, gregarious or occasionally arising in clusters of 3-7, stipitate, unknown in fresh condition, when . dry ca 0.5mm in diam and ca 0.5mm high, disc drying flat to slightly cupu- late, rehydrating flat and expanded, Hymenium drying shades of yellow to flesh-colored, rehydrating glassy-translucent; margin drying lighter than hymenium, rehydrating concolorous with hymenium; receptacle drying concol- orous with the margin, lighter than hymenium, rehydrating glassy-translu- cent and concolorous with hymenium; stipe concolorous with receptacle in dry and rehydrated conditions, cylindrical, short, generally less than 0.5) mm high and wide, occasionally subpapillate in appearance. Apothecial anatomy — Asci 8-spored, (40-)52-58(-65)x(5-)6-7um, pro- duced from small, replicating croziers, cylindric to slightly clavate, gradually tapering toward the base and there becoming slightly expanded to form a small foot, wall to ca lum thick, slightly enlarged at the apex and there to ca 2um thick; pore apparently J- in Melzer's Reagent. Ascospores (S-)6-8(-9)x2-3um, obliquely uniseriate, irregularly biseriate or biseri- — ate throughout, hyaline, smooth (a few devoid of pigment possibly lightly punctate), aseptate, obovoid to obpyriform, ends rounded or rarely slight- ~ ly pointed, in outline generally equilateral, anterior end obviously broader than posterior end, occasionally slightly indented below median point of spores; guttules generally absent or rarely with one tiny (ca lum or less), transversely positioned at the narrowest portion of the spore and appearing as a septum. Paraphyses equal to or slightly exceeding the asci, internally hyaline, branching toward the base of the asci, septate, cylindrical, generally not becoming expanded at the apex and there 2-3ym wide, walls thin, smooth, and hyaline. Subhymenium well developed, well BIG, 6. Helottum mtdlandense, Stevens 59 ex NY, freehand drawings, 1,000, A. Median longitudinal section of an apothecium through margin, B, Median ~ longitudinal section of an apothecium through the juncture of receptacle and stipe. C. An ascus with 8 ascospores and a paraphysis. D. 9 asco- spores drawn after discharge from ascus. 78 differentiated from the medullary excipulum below, hyaline, to ca 50um 4 broad in the center, narrower toward the margin, consisting of parallel to, slightly interwoven, vertically interwoven hyphae, the individual hyphae hyaline, 1.5-3um wide, the walls thin, hyaline and smooth. Medullary ex- cipulum poorly developed, obconical, non-refractive, hyaline, consisting of septate, branched, loosely to tightly interwoven (to parallel in the flank) hyphae 2-3(-4)um wide, the walls thin, hyaline and smooth. Ectal excipulum: inner ectal excipulum well to poorly defined, well-differenti- | ated from the outer ectal excipulum and grading into the medullary excipu- | lum, entire layer non-refractive, hyaline, 15-20um broad toward the margin | and approximately the same toward the stipe, consisting of parallel to slightly interwoven, hyaline hyphae 2-3um wide, the walls thin, non-re- fractive, hyaline and smooth. Outer ectal excipulum non-gelatinized, non-| refractive, entire layer hyaline, toward the stipe ca 50yum broad, consist-| ing of large, globose, angular to irregular cells forming a well defined | textura globulosa to angularis, the individual cells (5-)8-12(-20) um wide, | the walls thin to slightly thickened, hyaline and smooth. Outer covering layer and hairs absent; progressing towards the margin the outer ectal ex- | cipulum becoming narrower, and grading from a textura angularis and globu- | losa to textura prismatica, the individual cells smaller and forming more | and more regular rectangularly shaped cells until at the margin the layer composed of a well defined textura prismatica, lacking an outer covering layer, Margin simple, the brick-shaped cells if continuing into margin only a short distance and the remainder simple and composed of the narrow | cells of the inner ectal excipulum and the adjoining paraphyses, hairs | also absent in the margin. Stipe constructed similarly to the lower por- | tion of the receptacle, to the outside a zone to ca 70Oym broad composed of | globose to angular cells and grading into narrow, parallel hyphae in the interior of the stipe; the individual ‘cellsiwith wallssthingtossligngly thickened, hyaline and smooth or rarely roughened; at the base rind cells visible; hairs absent. Habitat: Old pods of Gledttsta trtacanthos L. and reported by White (1944) to occur also on petioles and midveins of Quercus sp. Etymology of the specific epithet: refers to the part of the country where the fungus was originally collected. Holotype (not examined): Iowa, Homestead, pods of Gledttsta trtacan- thos, 26 Sept 1931, G. W. Martin 5186 (FH). Illustrations: White, Farlowia 1: 607, "figs. 10-13. 1944" Specimen examined: Kansas, Lawrence, on pods (? of Gledttsta sp.), 1890, W. C. Stevens 59 (paratype ex NY). NOTES. When White (1944) described Helotium midlandense, he noted that the stipe to the outside was composed of globose | to angular cells, and he illustrated the ectal excipulum as formed of brick-shaped cells in the lower portion approaching the stipe. I have examined a paratype collection and concludem that the majority of the ectal excipulum is composed of a well defined textura globulosa to angularis, but that toward the margin the tissue becomes more organized into a well defined textura prismatica, and not toward the stipe as illustrated by White. Further, I have found a well developed rind on the substrate in association with the apothecia and in section have observed rind cells at the base of the stipe. I have also noted that the spores are relatively small for the Scle- 1s) otiniaceae and are oboyoid to obpyriform, Since the species Peoduces arsubstrata le stroma. 1t should be referred to the pelenoviniaceae, —iBecause (i)) the ectal; excipulum is composed of a textura globulosa, (ii) small pyriform ascospores are present, and (111) it occurs on fruits, the species is prob- Bplybest referred Co Ciboria. At present, J] am- uncertain of species concepts in Ctborta and of the separation between Ci- borta and Moellerodtseus; I will not make a new combination in Ctborta, but will defer action until the species are better known. I have observed in several apothecia a rather high propor- tion of misshapen and malformed ascospores, some of which pro- duced an apparent septum. I cannot satisfactorily explain the presence of the irregularly formed ascospores, nor the occar sional production of a single septum. My observations are then different from White's, who did not» report the presence of a stroma and referred it to the Leotiaceae (Helottum). Neither Dennis (1964) nor Arendholz (1979) treat the species. ice Herortum pnytlogenon Rehm, Hedwigia 24: 14. 13885. = Hymenoscyphus phyllogenon (Rehm) Kuntze, Revis. gen, Pl, 3(3): 485, TSOse NOTES. Helottum phyllogenon was originally described from Hungary on leaves of "poplar." Kuntze, (1898) transferred the species to Hymenosecyphus, and the placement there was accepted Dy Dennis (1964) and by Arendholz (1979). I find the type collection to be indistinguishable from #. caudatus, following Poewconcept Of Dumont Gg. Carpenter (1981). They gave the asco- PoOore Measurements O12. caudatus to be (14-)16-23(-26)x4-5 (-6)ym, whereas in the original description of H, phyllogenon the :ascospore measurements were 12-15x3.5um. White (1943) gave them as 11-14x3.8-5um, Dennis (1956) 14-16x4-6um, Arend- Zc, oye oxo. OAM. tL tind =the apothecial “structure of the type of H. phyllogenon indistinguishable from H. cauda- tus, and conclude that they represent the same species, with the ascospore measurements of the type of H. phyllogenon fall- ing at the lower limits of #. caudatus and perhaps extending the limits somewhat lower than previously reported by Dumont § Barpenter (19381 )*, Holotype: Hungary: bei Ungerisch-Altenburg (Ungarn), an faulen Pallel- Blattern, Oct 1883, Linhart (ex FH, Rehm, Ascomyceten 768). 13. Helotium phyllophilum (Desmaziéres) Fries, Summa veg. Scandi. 2500. 1849. Peziza phyllophtla Desmaziéres, Ann., Sci. Nat. Bot., Sér. 2, 17: 98. 1842; PZ. Crypt (du Nord) France éd. 1. no. 1159. 1842; éd. eiser- leno. 659. 91842. = Hymenoscyphus phyllophtlus (Desmaziéres) Kuntze, Revis. gen. Pl. 3(3): 485. 1898. Wl NOTES. ‘Peziza phyllophila was originally described as oc- curring on leaves of Acer and Fagus from France and was trans- 80 ferred to Helottum by Fries and to Hymenoscyphus by Kuntze (1898). Its placement in Hymenoscyphus was accepted by both Dennis (1964) and Arendholz (1979). I find the sterile tissue# of the apothecium of the lectotype specimen of the species to] be indistinguishable from that of H. caudatus. The apothecia of H. phyllophilus were originally thought to be smaller than those of H. caudatus, but as Dumont §& Carpenter have pointed out, the apothecia of tropical collections are frequently less@ than lmm in diameter and those of the type of H. phyllophilus | would then fall within the range of H. caudatus. In the original description the measurements of the asco- spores were not given, but they were reported to be 1-septate.} Dennis (1956) reported the ascospores to be 14-16x3.5-4um, White (1943) gave them as 11.5-15x3.2-4ym, and Rehm (1893, in 1887-1896) had them 10-15x3-3.5ym. Rehm further placed Helo- tium phyllophtlum and H. phyllogenon into synonymy. The only difference which I consider noteworthy between Hymenoscyphus caudatus and H..phyllophilus is.that in the latter aliotache ascospores in the type collection are l-septate, whereas, as pointed out by Dumont §& Carpenter (1981), in H. caudatus they are.only rarely l1-septate. However, I have noted in other clearly marked species, such as H. rufocorneus (see Dumont, 1981) where for some unexplained reason all of the ascospores in a single collection become 1l-septate. I do not consider | this then to represent the single criterion upon which to sep# arate two species, H. caudatustand) A. phyblophilus. s hatnet as prefer to consider them as probable synonyms. There 1S some question as to where, the name fe2z7zaquony lo phittla" was actually first published wand as tos whagetnescype | specimen actually is. Desmazi@éres (1842) presented a formal description of the species and a description also appeared on the labels of two specimens cited by Desmaziéres; "Pl. Crypt. @ éd. J, no. 1159; éd. 2, no. 659.'"" As nearly as (can, be (detcr am mined (Sayre, 1969), fascicles containing these two specimens were issued simultaneously. I have been unable to determine if the article appeared before the exsiccati specimens. In this regard, White (1943) selected no. 1159 as the type, but indicated that he examined two specimens, one in the bound) set of the exsiccata at FH and the other from the Curtis col-@ lections. I select the portion deposited in the bound exsic- @ cata as the lectotype specimen for Péztiza phytlopnitla. Lectotype specimen: Plantes Cryptogames de France, éd. 1, no. 1159 (ex FH, bound set). 14. Helottum translucens White, Farlowia 1; 149. 1943. FIG. 7. Hymenoscyphus transtlucens, holotype ex FH, freehand drawings, x 1,000. A. Median longitudinal section of an apothecium through margin. B. Median longitudinal section of an apothecium through margin. C. An ascus with 8 ascospores. D. A branching paraphysis. EE. 8 ascospores drawn after discharge from ascus. 82 = Hymenoscyphus translucens (White) Arendholz, Morph.-tax. Untersuch. blattbe. Ascomyceten Helotiales p. 78. 1979. Apothecial morphology — Apothecia tiny, solitary to occasionally gre- garious, arising from leaf blades, especially veins of various sizes, with a minute stipe, to ca 0.5mm in diam and to ca 0.4mm high, when fresh disc flat, drying and rehydrating same. Hymenium pallid to hyaline when fresh, | drying off-white to pale ochraceous, rehydrating lighter, pale flesh-col- ored to off-white; margin concolorous with hymenium when fresh, dry and rehydrated; receptacle generally concolorous with the hymenium when fresh, | dry and rehydrated; stipe, difficult to observe coloration owing to minute | Size. | Apothecial anatomy — Asci 8-spored, 80-95x9-llum, probably produced from tiny croziers, long cylindric-clavate, gradually tapering toward the base and there not expanded to form a small foot, wall ca lum wide, slightly enlarged at the rounded to subtruncate apex and there 2(-3)um thick; pore J+. Ascospores (10-)11-14(-15)x(3-)4(-5)um, uniseriate to bi-| seriate throughout, hyaline, smooth, aseptate or very rarely l-septate, trapezoidal, obovoid, ends rounded or less commonly slightly pointed, in | outline generally equilateral, if inequilateral only slightly so and then not flattened on one side, anterior end broader than posterior end, fre- quently appearing eguttulate or with two polar guttulate areas generally | obscure but granular and oily in composition, walls slightly thickened and occasionally to ca lym wide. Paraphyses equal to or slightly exceeding | the asci, internally hyaline, branching at the base of the asci and toward™ the middle of the asci, filiform, becoming slightly expanded at the apex | and there (2-)3-4ym wide, walls thin, smooth and hyaline. Subhymenium not” well differentiated from the medullary excipulum, but with a tendency of || the hyphae at the base of the asci to be slightly more compact and narrow-_ er than the hyphae in the center of the receptacle. Medullary excipulum poorly developed, obconical, non-refractive, hyaline, consisting of .sep- tate, branched, more or less parallel to slightly interwoven hyphae 2-4 (-5)um wide, the walls thin, non-refractive, hyaline and smooth. Ectal excipulum: inner ectal excipulum poorly defined and differentiated from medullary excipulum. Outer ectal excipulum non-refractive, non-gelati- nized, entire layer hyaline to subhyaline, ca 15-20ym wide toward the mar-_ gin and to ca 20-25ym toward the stipe, consisting predominantly of tex- | tura prismatica with the individual hyphae extending parallel to or at low to high angles to the surface of the apothecium and without apically free hyphal tips; the individual cells frequently collapsing, becoming disori- ented owing to drying process and tissue then appearing to be composed of narrow hyphae; individual cells toward the margin 5-12(-15)x3-5ym and 12- 22x3-7ym toward the stipe, the walls thin to slightly thickened and ca 1 um broad, hyaline to light brown and smooth. Outer covering layer pres- ent, but difficult to detect, 1-2 hyphal layers and 1-3um broad, the in- dividual hyphae extending parallel to the surface of the apothecium, over- | lapping, terminating before the margin, the individual cells subhyaline, the walls non-refractive, thin, pigmented light brown to intense brown, smooth to frequently slightly roughened. Hairs absent. Margin poorly de- veloped, narrow above, broader below, entire zone hyaline to light brown, constructed similarly to the flank below, the individual cells smaller. Stipe in the upper portion constructed similarly to the lower portion of the flanks, not becoming noticeably interwoven in the center, toward the base the individual hyphae becoming disrupted, and frequently losing hy- phal orientation, but generally oriented at a high angle to the surface. Hairs absent. 83 Habitat: Leaves of Acer sp., Quercus sp. and Fagus stlvattca (fide Arendholz, 1979). Etymology of the specific epithet: refers to the color of the apotheci- um in its fresh condition. Holotype: U.S.A., Mass., swamp at east side of Great Blue Hill, near Milton, on decaying leaves of Acer sp., 11 Nov 1941, D. H. Linder, E. V. seeler G W. L. White s.n. (FH). Paratype: U.S.A., Mass., Stony Brook Reservation, West Tuxbury, fallen leaves of Quercus sp., 11 Nov 1941, W. L. White s.n. (FH). Pilustracions: White, Farlowia 1:31595,, £19. 6..1943- Arendholzs Mor- phologisch-taxonomische Untersuchungen an blattbewohenden Ascomyceten aus MoreHelotrales plomi0, L19S: 5, eo.unl 9/9. NOCD OMe NentS deSCEINDCION Ole HeLOL TUM Lrane ucenss White (1943) concluded that this new species was closely related to Helottum caudatum, and I am in agreement with this. dHelottum Pauaarum 1S) dadistinguashnedy trom Hi translucens in®that the apo- mneCiavOt A. evrransilucens, are smaller than those of f° *caudatum and the ascospores in H#:-caudatum are predominantly ‘inéequilat- erateancetrequenctly Nnooked apically sn) contrase the spores of Pomcrane pucenceare trapezoldalwin shape: andsgenerally equilat- eral. s lhe shape of the ascospores in”. translucens 1S sug= PoctiViewOt ChOSCmOLa-m7Tmnicap1 cow mine Structure: OL the apo- thecia in these two species is fundamentally different. See discussion under H. tmmutabtle for a comparison of apothecial structure. PhemspecresmWwasenO tac reatedebysDenn1s. (1956,.1904).,, butt accept the placement of #. translucens in Hymenoscyphus by mrendholz (¢1979).. Acknowledgements anesauthorawisnesto express is gratitude, to the directors of those institutions who have provided the loan of material for the present study, injparticular, Dr. D. H. Pfister, of FH has been most helpful in the loan of specimens and assistance in working out bibliographic citations. Fi- nancial support for the present study has been provided by the National Science Foundation grant, GB. 28593. The line drawings have been prepared by Bobbi Angell. LITERATURE CITED Arendholz, W.-R. 1979. Morphologisch-taxonomische Untersuchungen an blattbewohenden Ascomyceten aus der Ordnung der Helotiales. A disser- tation presented to the faculty of Hamburg University. 1-115 pp., pl. 1-16. Carpenter, S. E. 1981. Monograph of Croctecreas (Ascomycetes, Helotiales, Leotiaceae). Mem. New York Bot. Gard. 32: 1- in=press. & K. P. Dumont. 1978. Los Hongos de Colombia-IV. Bisporella trisep- tata and its allies in Colombia. Caldasia 12(58): 3359-348, Dennis, R. W. G. 1956. A revision of the British Helotiaceae in the her- barium of the Royal Botanic Gardens, Kew, with notes on related Europe- anpspectes, e Mycol. ap., 02251-2116, apl 1. 1964. Remarks on the genus AHymenoscyphus S. F. Gray, with observa- tions on sundry species referred by Saccardo and others to the general 84 Helotium, Peztzella or Phtalea. Persoonia 3: 29-80. Desmaziéres, J. B. H. J. 1842. Cryptogames nouvelles. (Ann. Sci 7sNat. Botewsoen 2 salir la Liss Dumont, K. P. .1971. Sclerotiniaceae II. Lambertella. Mem. New York Bot. Gard’: 22(4) te-1=7 3" 1976. Sclerotiniaceae XI. On Moellerodiscus. Mycologia 68: 233- PLONE 1980. Sclerotiniaceae XVI. On Helotium rufo-corneum and Helottum fraternum. Mycotaxon 12(1): 255-277. 1981. Leotiaceae II. A preliminary survey of the neotropical species referred to Helottum and Hymenoscyphus. Mycotaxon 12(2): 313- SAG & S. E. Carpenter. 1981. Leotiaceae IV. Los Hongos de Colombia-VII. _ Hymenoscyphus caudatus and related speci¢s from Colombia and adjacent regions. Caldasia (in press). Haines, J. H. 1980. Studies in the Hyaloscyphaceae 1: Some species of Dasyscyphus on. tropical ferns. Mycotaxong ll 31389-21673 Kuntze; 0; 18984) sReviss gen. Pl 445(3)? Gl-576- Peck; C.,H#) 1879. Reportof the botanist.) Annual) Rep. New, Yorkeotate Musii5517 19-60. Rehm, H. 1887-1897. Die Pilze. Ascomyceten: Hysteriaceen und Discomy- ceten. In Rabenhorst's Kryptogamen-Flora von Deutchland, Oesterreich Undederecchwre zee cenUllawico) ela 75. Sayre, G. 1969. Cryptogamae Exsiccatae - An annotated Bibliography of Published Exsiccatae of Algae, Lichens, Hepaticae, and Musci. Mem. New LOTKBOtUs Gard 1619 (0) cme Lae White, W. L. 1942. Studies in the genus Helottum, I. A review of the species described by Peck: Mycologia 34: 154-1797 1943. Studies in the genus Helottum, III. History and diagnosis of certain European and North American foliicolous species. Farlowia 1: 135-170. 1944. Studies in the genus Helottum, IV. Some miscellaneous spe- cies meebariowiarl: 8599-6172 MYCOTAXON Vode ALIS No eal epp. S024 04 April-June 1981 LICHENES EXSICCATI DISTRIBUTED BY THE UNIVERSITY OF COLORADO MUSEUM, BOULDER FASCICLES 1-15, NOS. 1-600, 1961-1979 WILLIAM A. WEBER Univ. of Colorado Museum, Campus Box 218, Boulder, 0 80309 SUMMARY Revision of identifications and nomenclature for the first 600 numbers of LICH. EXSICC. OOLO is provided, and the following new combinations are proposed: Cladina galapagosensis, C. polia, Hetero- dermia barbifera, H. circinalis, H. stellata and H. verrucifera. Validation is provided for the publi- cation of three new species distributed in the ex- siccati: Lecanora pseudopinguis, L. texana and Psora cerebriformis. Lichenes Exsiccati, distributed by the University of Colorado Museum (standard abbreviation LICH. EXS. COLO) began in 1961 with the distribu- tion of Fascicle 1 (1-40). The latest fascicle was Fascicle 15 (56]- 600), distributed in 1979. Some of the early fascicles were distributed in 100 sets, but the current ones are reduced to 60 sets. The exsiccati are offered in exchange to most of the active lichen herbaria of the world. A brief notice was published in Taxon 13:31. 1964, wherein the institutions receiving the complete series were listed. The present pa- per provides an alphabetical arrangement of the numbers thus far distri- buted, with updated nomenclature and corrections of identifications up to the present moment. Additional corrections will be welcomed. In the following list, the first name given is that under which the number was distributed. When that name is enclosed by brackets, this in- dicates that the name has been changed, amd the current name is preceded by an (=) sign. If the identification has been changed, no (=) sign pre- cedes the second name. 86 TAXON Acarospora badiofusca (Nyl.) Th. Fr. [A. flava (Bell.) Trev.] =A. chlorophana (Wg.) Ach. A. nitida H. Magn. A. schleicheri (Ach. ) Mass. [Alectoria chalybeiformis (L.) S. Gray] =Bryoria chalybeiformis (L.) Brodo & Hawksw. [A. fremontii Tuck.] Bryoria tortuosa (Merr.) Brodo & Hawksw. [A. glabra Mot.] Bryoria lanestris (L.) Brodo & Hawksw. A. lata Tayl. A. nigricans (Ach.) Nyl. A. ochroleuca (Hoffm.) Mass. A. poeltii Bystrek [A. pubescens (L.) R. H. Howe] =Pseudephebe pubes- cens (L.) Choisy A. sarmentosa (Ach.) Ach. A. smithii DuRietz [A. tenuis E. Dahl] Bryoria chalybeiformis (L.) Brodo & Hawksw. [Anaptychia barbifera (Nyl.) Trevis. ] =Heterodermia barbifera comb. nov. [A. circinalis (A. Zahlbr.) W. A. Web.] =Heterodermia circinalis comb. nov. [A. diademata (Tayl.) Kurok.] =Heterodermia diadem- ata (Tayl.) Awasthi A. erinacea (Ach.) Trevis. [A. leucomela (L.) Mass. f. verrucifera Kurok. ] =Heterodermia verrucifera comb. nov. [A. multiciliata Kurok.] =Heterodermia multiciliata Follm. & Redon [Anaptychia obscurata (Nyl.) Vain.] =Heterodermia obscurata (Nyl.) Trevis. [A. stellata (Vain.) Kurok.] =Heterodermia stellata (Vain.) W. A. Web., comb. nov. Anthracothecium ochraceoflavum (Nyl.) Muell.-Arg. Anzia angustata (Pers.) Muell.—Arg. A. gregoriana Muell.—Arg. A. hypoleuca Muell.-Arg. A. wilsonii Raes. Arthonia glebosa Tuck. Norway California Wyoming California Tasmania Canada N. Guinea Canada Norway California N. Guinea Colorado N. Guinea Galapagos Arizona Mexico Mexico Galapagos Chile Australia Galapagos Galapagos Australia N. Guinea N. Guinea Australia Colorado p35 508 145 281 377 378 311 53 A. impolita (Ehrh.) Borr. ex Hook. & Sowerby A. platyspilea Nyl. A. rubella (Fee) Nyl. Arthopyrenia halodytes (Nyl.) Arn. [Arthothelium spilomatoides (Nyl.) A. Zahlbr.] A. galapagoense Huneck & Follmann TYPE OOLL. Aspicilia alpina (Sommerf.) Arn. A. desertorum (Kremp.) Mereschk. [A. hispida Mereschk.] A. desertorum (Kremp.) Mereschk. [A. mutabilis (Ach.) Koerb.] =Pachyospora mutabilis (Ach. ) Mass. A. quartzitica W. A. Web. TYPE COLL. A. transbaicalica Oxner A. sp. indet. (fruticose modification) Bacidia albescens (Arn.) Zwackh. B. herrei A. Zahlbr. B. millegrana (Tayl.) A. Zahlbr. Baeomyces absolutus Hepp in Zollinger B. trachypus Nyl. B. weberi J. W. Thomson TYPE OOLL. Belonia americana Fink ex Hedrick Bryoria tortuosa (Merr.) Brodo & D. Hawksw. Buellia curtisii (Tuck.) W. A. Web. B. flavoareolata (Nyl.) Muell.-Arg. B. galapagona W. A. Web. TYPE COLL. B. glaziouana (Kremp.) Muell.-Arg. B. novomexicana B. de Lesd. B. oidalea (Tuck.) Tuck. B. punctata (Hoffm.) Mass. [B. spuria (Schaer.) Anzi] undescribed taxon. . cf. taltalensis Dodge . triphragmioides Anzi sp. indet. . Sp. indet. . zahlbruckneri Steiner Calicium abietinum Pers. [C. hemisphaericum Howard] =C. adequatulum Nyl. Caloplaca amabilis A. Zahlbr. C. aurantiaca (Lightf.) Th. Fr. Incorrect, possibly an | Gd) Oo) | [C. bracteata (Hoffm.) Mass.] Fulgensia desertorum (Tomin) Poelt, cited specimen! [C. chrysophthalma Degel.] Determination doubt ful. C. cinnanomea (Th. Fr.) Oliv. California California USSR California California Oregon Galapagos N. Guinea N. Guinea N. Guinea N. Guinea Texas Canada Louisiana Chile Galapagos Galapagos Arizona California Colorado Mexico Chile Colorado Peru Peru Arizona Idaho Montana Arizona Arizona Colorado N. Mexico Colorado 88 C. cladodes (Tuck.) A. Zahlbr. in Engler C. epithallina Lynge C. fernandeziana (A. Zahlbr.) Follm. & Redon C. lamprocheila (DC.) Flagey C. microphyllina (Tuck. ) Hasse C. modesta (A. Zahlbr.) Fink C. pollinii (Mass.) Jatta C. stanfordensis H. Magn. C. subnitida (Malme) A. Zahlbr. C. trachyphylla (Tuck.) A. Zahlbr. [Caloplacopsis submexicana (B. de Lesd.) B. de Lesd. ] =Candelina submexicana (B. de Lesd.) Poelt Candelariella deflexa (Nyl.) A. Zahlbr. C. rosulans (Muell.—Arg.) A. Zahlbr. C. spraguei (Tuck.) A. Zahlbr. Catillaria griffithii (Sm.) Malme C. sp. indet. Cavernularia lophyrea (Ach.) Degel. Cetraria canadensis (Raes.) Raes. C. cucullata (Bell.) Ach. C. delisei (Bory ex Schaer.) Th. Fr. C. ericetorum Opiz C. fendleri (Nyl.) Tuck. [C. fendleri f. coralligera W. A. Web. TYPE OOLL.] =C. coralligera (W. A. Web.) Hale [c. glauca (L.) Ach.] =Platismatia glauca (L.) W. Culb. & C. Culb. C. idahoensis Essl. C. islandica (L.) Ach. C. pinastri (Scop.) S. Gray [C. richardsonii Hook. in Richards.] =Masonhalea rich- ardsonii (Hook. ip Richards.) Kaernefelt C. scutata (Wulf.) Poetsch C. tilesii Ach. C. sp. indet. Cetrelia chicitae (W. Culb.) W. Culb. & C. Culb. Chaenotheca brunneola (Ach.) Muell.~Arg. Chiodecton effusum Fee C. farinaceum Fee C. sp. indet. C. sp. indet. Chondropsis semiviridis (F, Muell. ex Nyl.) Nyl. ex Cromb. i ae Colorado Colorado Chile Colorado Arizona Arizona Minnesota California Texas Colorado Arizona Colorado Colorado Colorado Colorado California Chile Canada Idaho Alaska Norway Colorado Arizona Arizona Colorado Idaho Colorado Australia Colorado Alaska Tasmania Alaska N. Guinea N. Guinea Idaho Galapagos Galapagos Chile Chile Australia Australia Cladia aggregata (Sw.) Nyl. C. ferdinandii (Muell.-Arg.) R. Filson C. retipora (Labill.) Nyl. [C. retipora (Labill.) Nyl.] C. corallaizon R. Filson C. schizopora (Nyl.) Nyl. C. sullivanii (Muell.-Arg.) W. Martin [Cladonia alpestris (L.) Rabenh.] =Cladina stellaris (Opiz) Pouzar & Vezda [Cladonia cariosa (Ach.) Spreng. ] C._ceratophylla (Sw.) Spreng. C. chlorophaea (Flk.) Spreng. C. coccifera (L.) Willd. C, corymbescens Nyl. C.e a Leighton C. foliacea (Huds.) Willd. [C. galapagosensis Ahti] =Cladina galapagosensis comb. nov. C. leporina Fr. C. multiformis Merrill C. robbinsii Evans C. pertricosa Kremp. C. cf. pityrea (Flk.) Fr. [C. polia R. Sant.] =Cladina polia (R. Sant.) comb. pOve’ C. scabriuscula (Del. in Duby) Nyl. c. solitaria solitaria H. Magn. TC. squamosa squamosa (Scop.) Hoffm.] C. dilleniana Flk. fide J. W. Thomson ; [C. subtenuis (des Abb.) Evans] =Cladina subtenuis (des Abb.) Hale & W. Culb. C.. sulphurina (Michx.) Fr. C. verticillata (Hoffm.) Schaer. C. vulcanica Zoll. in Hasskarl [C. xanthoclada Muell.-Arg.] C. capitellata (Tayl.) e Raby ee. C. zopfii Vain. Coelocaulon aculeatum (Schreb.) Gyel. [C. australiense nom. nud. | Kaernefelt TYPE COLL. Coenogonium implexum Nyl. Collema cristatum (L.) G. H. Web. in Wiggers Cetraria australiense Nepal Australia Galapagos Australia Australia Australia Australia Australia Tasmania Alaska Kansas Galapagos Galapagos N. Guinea N. Guinea Colorado Denmark Galapagos Alabama Colorado — N. Mexico Australia Australia Galapagos Australia Chile N. Guinea N. Guinea Kansas Montana Australia N. Guinea Australia Denmark Colorado Canada Australia Tasmania Colorado 90 C. durietzii Degel. C. leucocarpum Hook. f. & Tayl. C. polycarpum Hoffm. Conotrema urceolatum (Ach.) Tuck. [Cora pavonia (Sw.) Fr.] =Dictyonema montanum (Sw. ) Marm. ex Follm. [Cornicularia aculeata (Schreb.) Ach.] =Coelocaulon aculeatum (Schreb.) Gyel. [C. californica (Tuck.) DuRietz] =Coelocaulon cali- fornicum (Tuck. ) Howe f. C. nommerica (Gum. ) DuRietz [Cypheliopsis bolanderi (Tuck.) Vain.] =Thelomma mammosum (Hepp in Hartung) Mass. The collection is mixed, containing also T. santessonii L, Tibell Cyphelium inquinans (Sm.) Trev. stocoleus ebeneus (Dillwyn) Thwaites Dactylina madreporiformis (Ach. ) Tuck. Darbishirella gracillima (Kremp.) A. Zahlbr. Dendrographa leucophaea (Tuck. ) Darb. D. minor Darb. Dermatoc leptophyllum (Ach.) Vain. D. miniatum Ta) Mann D. plumbeum B. de Lesd. D. reticulatum H. Magn. D. rivulorun (Arn.) Dalla Torre & Sarnth. D. tuckermanii (Rav. ex Mont.) A. Zahlbr. D. vellereum Zschacke _ Dictyonema irpicinum Mont. D. sericeum (Sw.) Berk. Diploicea canescens (Dicks.) Mass. Diploschistes ocellatus (Vill.) Norm. [D. scruposus (Schreb.) Nomm.] D. cf. canadensis Raes. Dirina catalinariae Hasse D. herrei A. Zahlbr. D. limitata Nyl. Dolichocarpus chilensis R. Sant. Endocarpon pulvinatum Th. Fr. E. pusillum Hedw. Enterographa atacamensis Dodge [Enterographa atacamensis Dodge] Roccellina olivacea Follm. Ephebe lanata (L.) Vain. Evernia prunastri (L.) Ach. EI ROBIE RICA EERE OE RR Australia Australia Colorado W. Virginia Galapagos Colorado Oregon Oregon Mexico Colorado Colorado Colorado Colorado Chile Mexico Mexico Calif. USSR Arizona Colorado Oregon Colorado Arizona USSR N. Guinea N. Guinea California Mexico Colorado Mexico Galapagos Chile Chile Colorado Colorado Colorado Peru Chile Colorado California 438 Everniastrum neocirrhatum (Hale & V. Wirth) Hale E. vexans Hale Glypholecia scabra (Pers.) Muell.-Arg. Graphis caesiella Vain. G. striatula (Ach.) Spreng. Gyrostomum scyphuliferum (Ach. ) Nyl. Haematonma babingtonii Mass. H. subpuniceum (Muell.-Arg.) B. de Lesd. Heppia lutosa (Ach.) Nyl. Heterodea melleri (Hampe) Nyl. Heterodermia leucomelos (L.) Poelt [Hubbsia lumbricoides W. A. Web. TYPE COLL.] =Reinkella californica Raes. Hydrothyria venosa Russell Hypogymnia billardieri (Mont.) R. Filson H. krogii Ohlsson H. lugubris (Pers.) Krog H. mundata (Nyl.) Rassadina H. oroarctica Krog H. physodes (L.) Nyl. Icmadophila ericetorum (L.) A. Zahlbr. Ingaderia pulcherrima Darb. Koerberia biformis Mass. Lasallia pensylvanica (Hoffm.) Llano L. pustulata (L.) Mereschk. ssp. papulosa (Ach. ) W. A. Web. Lecanactis californica Tuck. L. cf. myriadea (Fee) A. Zahlbr. L. (subg. Bactrospora) sp. indet. [Lecanora arizonica (Tuck.) W. A. Web.] =Omphalo- dium arizonicum (Tuck. ) Tuck. L. badia (Hoffm.) Ach. [L. bolanderi Tuck.] lL. phryganitis Tuck. [L. cancriformis (Hoffm.) Vain.] L. caesiorubella Ach. ssp. merrillii Imsh. & Brodo L. carpinea (L.) Vain. L. chlarotera Nyl. L. christoi W. A. Web. TYPE OOLL. L. confusa Almborn [L. conizaea (Ach.) Nyl.] Incorrect, but no alter- native identification available. [L. conizaeoides Nyl. ex Cromb.] L. confusa Almb. L. coquimbensis A, Zahlbr. L. frustulosa (Dicks. ) Ach. L. garovaglii (Koerb.) A. Zahlbr. ments (Disregard com- Mexico Galapagos USSR Galapagos Galapagos Louisiana New Zealand Texas Iowa Australia Mexico Mexico California Australia N. Carolina Australia Australia Colorado Colorado Colorado California Chile California N. Mexico Montana California California California Colorado Colorado Oregon Galapagos California Chile Colorado Colorado 92 [L. cf. glabrata (Ach.) Malme] Probably incorrect but no alternative name available. [L. lentigera (G. Web.) Ach.] =Squamarina lentigera (G. Web.) Poelt ([L. marginalis Hasse] =Rhizoplaca marginalis (Hasse) W. A. Web. [1.. melanophthalma Ram.] =Rhizoplaca melanophthalma (Ran. ) Leuckert & Poelt L. mellea W. A. Web. TYPE COLL. [L. peltata (Ram.) Steud.] =Rhizoplaca peltata (Ram. ) Leuckert & Poelt L. phryganitis Tuck. L. pinguis Tuck. L. polytropa (Ehrh.) Rabenh. L. pringlei M. Lamb L. pseudopinguis W. A. Web. sp. nov. TYPE COLL. [L. rubina (Vill.) Ach.] =Rhizoplaca chrysoleuca (Sm.) Choisy L. saligna (Schrad.) A. Zahlbr. L. texana W. A. Web., sp. nov. TYPE COLL. [L. thomsonii H. ent =L. novomexicana B. de Lesd. ex H. Magn. L., stirps varia (Ehrh.) Ach. Lecidea aspidula Kremp. [L. atrobrunnea (Ram. ): Schaer. ] Arn. L. berengeriana (Mass.) Th. Fr. L. leucothallina [L. cinnabarina Somerf. | (Ach.) Raes. [Lecidea decipiens Ach.] =Psora decipiens Hoffm. (Ignore the incorrect synonymy on the label) L. dolodes Nyl. ex Hasse L. elabens Fr. L. elata Schaer. Protoblastenia russula [L. friesii Ach. in Liljebl.] =Hypocenomyce friesii (Ach. in Liljebl.) G. Schneider [L. icterica (Mont.) Tayl.] =Psora icterica (Mont.) Muell.—Arg. L. insularis Nyl. L. leucothallina Arn. L. limosa Ach. [L. luridella Tuck. ] L. luridella =Psora luridella (Tuck.) Fink L. lyngei Degel. Texas Colorado California Colorado Wyoming California Colorado Colorado California Mexico Colorado Colorado Galapagos Colorado Colorado ‘Texas Colorado Colorado Australia Colorado ‘Colorado Oregon Australia Colorado California Australia Colorado Australia N. Mexico Montana Montana N. Guinea Colorado Colorado 436 18 451 273 155 428 272 435 424 [L. novomexicana (B. de Lesd.) W. A. Web. ex R. Ander— son] =Psora nipponica (A. Zahlbr.) G. Schneider L. nylanderi (Anzi) Th. Fr. [L. quadricolor (Dicks.) Borr.] =L. granulosa (Hoffm. ) Ach. [L. rubiformis (Wahlenb. ex Ach.) Ach.] Psora tuck ermanii R. Anderson ex G. Schneider, invalid. no Latin description. __ [L. rubiformis (Wahlenb. ex Ach.) Ach.] Psora cere- briformis W. A. Web., sp. nov. [L. rufonigra (Tuck.) Nyl.] =Psorula rufonigra (Tuck. ) G. Schneider [L. symmicta (Ach.) Ach.] Lecanora cadubriae (Mass.) Hedlund [L. texana W. A. Web.] =Xanthopsora texana (W. A. Web.) G. Schneider & W. A. Web. TYPE OOLL. [L. wallrothii Flk. ex Spreng.] =Trapeliopsis wall- rothii (Flk. ex Spreng.) H. Hertel & G. Schneider Lecidella elaeochroma (Ach. ) Haszl. [Leprocaulon albicans (Th. Fr.) Nyl. ex Hue] rocaulon gracilescens M. Lamb & Ward L. arbuscula (Nyl.) Nyl. ex Hue [L. microscopicum (Vill.) Gams ex D. Hawksw. ] L. tenellum (Tuck.) Nyl. Leprocaulon tenellum (Tuck.) Nyl. Leptogium brebissonii Mont. in Webb L. burnetiae Dodge var. hirsutum (Sierk) P. M. Joerg. L. cyanescens (Ach. ) Koerb. L. denticulatum Nyl. [L. foveolatum Nyl.] L. punctulatum Nyl. [L. furfuraceum (Harm. ) Sierk] Lesd. ) Dodge L. hypotrachynum Muell-Arg. L. javanicum (Mont. & v. d. Bosch) Mont. L. lichenoides (L.) A. Zahlbr. L. ml legranum Sierk L. moluccanum (Pers. in Gaud.) Vain. L. phyllocarpum (Pers.) Mont. L. rugosum Sierk SEES vulpina (L.) Hue L. wulpina (L.) Hue f. californica (Lev.) W. A. Web. Richina Lichina confinis (0. F. Muell.) C. A. Agardh L. papillosum (B. de | Idaho Minnesota Arizona Colorado Colorado Colorado Texas Colorado Texas California Colorado Arizona Qolorado N. Guinea Galapagos Peru Australia Colorado Galapagos Colorado Mexico Galapagos Arizona Australia N. Guinea Colorado Oregon Arizona Arizona N. Guinea Mexico Arizona California Nevada Tasmania 93 224 536 103 124 94 L. tasmanica Henssen Lobaria hallii Tuck. L. retigera (Bory) Trevis. [L. stictaeformis (Schaer.) Trevis.] L. discolor (Bory in Del.) Hue Lopadium pezizoideum (Ach. ) Koerb. Maronella laricina Steiner ISOTYPES Mastodia tessellata auct. (See Brodo, The Bryologist 79: 396-398. 1977, for nomenclatural discussion) Melanaria melanospora (Nyl.) Erichsen [Menegazzia aeneofusca (Muell.-Arg.) R. Sant.] M. nothofagi P. James ined. M. pertusa (Schrank) Stein Micarea denigrata Fr.) Hedlund Microthelia micula Koerb. Mycoglaena myricae (Nyl.) R. C. Harris Neophyllis melacarpa (F. Wils.) F. Wils. G. Schneider maintains this over Yoshimura's placement in Gymnoderma Nephroma arcticum (L.) Torss. N. australe A, Richard N. cellulosum (Sm. ex Ach.) Ach. N. helveticum Ach. N. resupinatum (L.) Ach. [Nephromopsis californica Gyel.] =Cetraria orbata (Nyl.) Fink Neuropogon acromelanus (Stirt.) M. Lamb N. sulphureus (Koenig) Hellb. Normandina pulchella (Borr.) Nyl. Ocellularia alba (Fee) Muell.-Arg. Ochrolechia grimmiae Lynge O.upsaliensis (L.) Mass. Omphalaria kansana Tuck. Determination doubtful. Opegrapha saxicola Ach. Pannaria leucophaea (Vahl) P. M. Joerg. [P. nigrata Muell.-Arg.] Possibly P. tavaresii P. M. Joerg., cf. Opera Bot. 45:70. 1978. P. pezizoides (G. Web.) Trevis. Parathelium sp. indet. [Parmelia bostrychodes A, Zahlbr.] =Hypotrachyna bostrychodes (A. Zahlbr.) Hale [P. caperata (L.) Ach.] =Pseudoparmelia caperata (L.) Hale [P. chiricahuensis R. Anderson & W. A. Web. ] =Neofuscelia chiricahuensis (R. Anderson & W. A. Web.) Essl. Tasmania Oregon India N. Guinea Idaho Austria Alaska Chile Australia N. Guinea Colorado Arizona Colorado Australia Alaska Tasmania Australia Colorado Alaska California Tasmania Spitzbergen Australia Australia Spitzbergen Colorado Colorado California USSR Australia Colorado Chile N. Guinea Australia Arizona Texas [P. conspersa (Ach.) Ach. ] chroa (Tuck. ) Hale [P. dichotoma Muell.-Arg. ] toma (Muell.-Arg.) Hale [P. dominicana Vain. ] Hale [P. elegantula (A. Zahlbr.) Szatala] elegantula (A, Zahlbr.) Essl. [P. endosulphurea (Hillm.) Hale] sulphureun (Hillm. ) Hale [P. exasperatula Nyl.] =Melanelia exasperatula (Nyl.) Ess1. [P. furfuracea (L.) Ach.] Pseudevernia intensa (Nyl.) Hale & W. Culb. [P. galbina Ach.] =Parmelina galbina (Ach.) Hale [P. isidiotyla Nyl.] Neofuscelia loxodes (Nyl.) Essl. [P. novomexicana Gyel. ] (Gyel.) Hale [P. perlata (Huds.) Ach. ] (Huds. ) Hale [P. physodes (L.) Ach. ] Nyl. P. praesignis Nyl. P. pseudotenuirima Gyel, [P. pulla (Schreb.) Ach.] =Neofuscelia pulla (Ach.) Essl. [P. pulvinata Fee] Hale [P. revoluta Flk.] =Hypotrachyna revoluta (Flk.) Hale [P. rutidota Hook. f. & Tayl.] =Pseudoparmelia rutidota (Hook. f. & Tayl.) Hale P. saxatilis (L.) Ach. [P. saximontana R. Anderson & W. A. Web.] TYPE COLL. =Melanelia substygia (R. Anderson & W. A. Web.) Essl. P. signifera Nyl. P. subalbicans Stirt. [P. subcrinita Nyl.] =Parmotrema subtinctorium (Nyl.) Hale P. subrudecta Nyl. Xanthoparmelia chloro- =Xanthoparmelia dicho- =Melanelia =Parmotrema endo- =Parmotrema perlatun =Hypogymnia physodes (L.) psy OL uae ayia pL nse S [P. substrigosa Hale in W. A. Web.] =Xanthoparmelia substrigosa (Hale in W. A. Web.) Hale [P. subtinctoria A. Zahlbr.] =Parmotrema subtinc- torium (A. Zahlbr.) Hale =Parmotrema dominicana (Vain. ) =Xanthoparmelia novomexicana Wyoming Australia Galapagos Colorado Australia Colorado Arizona Australia Colorado N. Mexico Australia Australia Arizona Qolorado Australia Colorado Colorado Australia Australia Dominica Australia Australia Australia Arizona 95 29 96 [P. taractica Kremp.] =Xanthoparmelia taractica (Kremp.) Hale [P. tinctorum Nyl.] =Parmotrema tinctorum (Nyl.) Hale [P. ulcerosa A. Zahlbr.] =Xanthoparmelia ulcerosa (A. Zahlbr.) Hale [P. ulophyllodes (Vain.) Sav.] =P. soredica Nyl. [P. weberi Hale] =Xanthoparmelia weberi (Hale) Hale [P. wyomingica (Gyel.) Hale] =Xanthoparmelia wyo- mingica (Gyel.) Hale Parmeliopsis aleurites (Ach. ) Nyl. P. placorodia (Ach.) Nyl. Peccania sp. indet. Peltigera aphthosa (L.) Willd. Peltigera dolichorhiza (Nyl.) Nyl. P. horizontalis (Huds.) Baumg. P. horizontalis (Huds.) Baumg. f. zopfii (Gyel.) J. W. Thomson P. polydactyla (Neck. ) Hoffm. [P. scabrosa Th. Fr.] P. sp. nov. Vitikainen ined. P. venosa (L.) Willd. Peltula cylindrica Wetmore P. obscurans (Nyl.) Gyel. var. deserticola (A. Zahlbr.) Wetmore Pertusaria californica Dibben P. saximontana Wetmore P, xanthodes Muell.-Arg. Pertusaria (Subg. Lecanorastrum) sp. indet. [Phaeographina tridacna W. A. Web. sp. nov. ined.] P. isidiosa (Vain.) A. Zahlbr., det. Nakanishi. Phaeographis exaltata (Mont. & v. d. Bosch) Muell.- Arg. P. sp. indet. Phylliscum demangeonii (Moug. ) Nyl. Physcia adscendens (Th. Fr.) Oliv. em. Bitt. [P. aegialita (Ach.) Nyl.] Dirinaria subconfluens Awasthi, cited specimen. © P. caesia (Hoffm.) Hampe Identification incorrect, but no alternative available. P. callosa Nyl. P. duplicorticata W. A. Web. & J. W. Thomson TYPE COLL. Arizona Galapagos Venezuela Colorado Arizona Wyoming Arizona Colorado Arizona Mexico Colorado Australia N. Guinea Colorado N. Guinea Colorado Alaska Oregon Colorado Colorado Georgia Arizona California Colorado Louisiana Tasmania N. Guinea Mexico Galapagos Colorado Colorado Australia Australia Colorado Calif. [P. pulverulenta (Schreb.) Hampe f. coralloidea Suza in Nadv.| Physconia sp. indet. [P. setosa (Ach.) Nyl.] P. hispidula (Ach.) Frey [P. setosa (Ach.) Nyl.] Identification incorrect. P. stellaris (L.) Nyl. P. tribacoides Nyl. P. vitii Nadv. Physconia detersa (Nyl.) Poelt P. muscigena (Ach.) Poelt Physma byrsinum (Ach. ) Muell.—Arg. Pilophoron robustum Th. Fr. Placynthium nigrum (Huds.) S. Gray Platismatia stenophylla (Tuck.) W. Qulb. & C. Qulb. Polychidium muscicola (Sw.) S. Gray Porina epiphylla Fee P. rubentior (Stirt.) Muell.—Arg. P. sp. indet. Pseudocyphellaria argyracea (Del. ) Vain. P. australiensis H. Magn. [P. durvillei (el.) Vain.] P. hirta (G. Forst.) D. Gall. & P. James [P. freycinetii (Del.) Malme] Identification dubious. P. gilva (Ach.) Malme P. glabra (Hook. f. & Tayl.) Dodge P. impressa (Hook. f. & Tayl.) Vain. P. neglecta (Muell.-Arg.) H. Magn. [P. orygmaea (Ach.) Malme] P. hirta (G. Forst.) D. Gall. & P. James P. sp. indet. P. rainierensis Imshaug Psoroma soccatum R. Br. in Cromb. P. sphinctrinum (Mont.) Nyl. Psorotichia minuta H. Magn. P. sp. indet. Pyrenotrichum splitgerberi Mont. Pyrenula cerina Eschw. P. nitidella (Flk. ex Schaer.) Muell.-Arg. var. maculata R. C. Harris TYPE OOLL. Pyxine pringlei Imshaug [Ramalina ceruchis (Ach.) DeNot.] =Desmazieria ceruchis (Ach.) Trevis. [R. combeoides Nyl.] =Desmazieria combeoides (Nyl.) Follm. & Huneck R. complanata (Sw. ex Ach.) Ach. N. Mexico Arizona Australia Colorado Louisiana Germany Colorado Colorado Australia Australia Tasmania Australia Tasmania N. Guinea Tasmania Tasmania Australia Tasmania Australia Washington Australia Tasmania Colorado Texas Louisiana Galapagos Texas Galapagos Mexico Mexico Galapagos a7 98 R. farinacea (L.) Ach. R. fraxinea (L.) Ach. Identification doubtful. R. geniculata Hook. f. & Tayl. var. compacta Muell.— Arg. [R. homalea Ach.] =Desmazieria homalea (Ach. ) Mont. R. javanica Nyl. R. linearis (Sw.) Ach. R. menziesii Tayl., non Tuck. R. pacifica Asahina R. peruviana Ach. R. sinensis Jatta R. sorediantha Nyl. R. subleptocarpha Rundel & Bowler R. usnea (1. ) Howe f. [Ramalodium succulentum (R. Br.) Nyl.] Arctomia fruticosa Henssen & Weber. sp. nov. ined. Reinkella lirellina Darb. R. parishii Hasse Rhizocarpon macrosporum Raes. Rinodina applanata H, Magn. [R. archaea (Ach.) Vain. em. Malme] Sheard questions the identification. R. calculiformis W. A. Web. TYPE COLL. R. coloradiana H. Magn. R. svillensis H. Magn. [R. oreina (Ach.) Mass.] =Dimelaena oreina (Ach.) Norm. [R. radiata (Tuck.) Tuck.] =Dimelaena radiata (Tuck.) Hale & W. Culb. R. turfacea (Wahlenb.) Koerb. Roccella babingtonii Mont. R. fimbriata Darb. R. fucoides (Dicks.) Vain. R. portentosa Darb. [R. portentosa Darb.] R. galapagoensis Follm. TYPE COLL. Roccellaria mollis (Hampe) A. Zahlbr. Roccellina condensata Darb. Galapagos California Australia Tasmania Mexico California Philippines Galapagos N. Mexico Galapagos Galapagos Mexico Panama Galapagos Tasmania Peru California Colorado Louisiana California - California Mexico Colorado California Colorado California Colorado Mexico Galapagos Mexico Mexico Italy Chile Galapagos Galapagos Chile Peru Chile [R. lutosa Follm.] Roccellina luteola Follm. Sarcogyne clavus (Ram. ) Kremp. S. regularis Koerb. Schismatomma cupressum Herre Schistophoron tenue Stirt. Schizopelte californica Th. Fr. Siphula coriacea Tayl. ex Nyl. Siphula fragilis (Hook. f. & Tayl.) J. Murray, nom. nud, x ria S. sp. indet.? Speerschneidera euploca (Tuck. ) Trevis. Sphaerophorus melanocarpus (Sw. ) DC. S. tener Laur. Sporastatia testudinea (Ach.) Mass. Staurothele clopima (Wahlenb. ex Ach.) Th. Fr. Stereocaulon glareosum (Sav.) H. Magn. S. leprocauloides M. Lamb TYPE OOLL. S. massartianum Hue [S. microcarpum Muell.-Arg. ] TYPE OOLL. S. myriocarpum Th. Fr. [Stereocaulon piliferum Th. Fr.] misspelling for S. piluliferun th. Fr. S. pseudomassartianum M. Lamb ex Frey S. weberi M. Lamb, S. ramilosum (Sw.) Raeusch var. nudatum (Muel1.-Arg.) Muell.—Arg. S. rivulorum H. Magn. S. staufferi M. Lamb ex Frey S. virgatum Ach. ex Spreng. Sticta boschiana Mont. & v. d. Bosch ex Jungh. S. filix (Sw.) Nyl. ait S. [Pseudocyphellaria] cf. fragillima Bab. ex Hook., sensu A. Zahlbr. S. fuliginosa (Dicks. ) Ach. S. rubella Hook. f. & Tayl. -S. weigelii Isert. ex Ach. Strigula elegans (Fee) Muell.-Arg. Teloschistes exilis (Michx.) Vain. T. fasciculatus Hillm. T. flavicans (Sw.) Norm. T. velifer F. Wils. Chile Texas Texas California Galapagos Califormia Australia Australia Tasmania California Texas Tasmania Tasmania Colorado California Colorado Montana N. Guinea N. Guinea Philippines Galapagos N. Guinea India N. Guinea N. Guinea N. Guinea Australia Colorado N. Guinea Dominica N. Guinea Australia N. Guinea Australia Tasmania Arizona Louisiana Australia Texas Australia Galapagos Australia 100 T. villosus (Ach. ) Nor. Thamolia vermicularis (Sw.) Schaer. T. vermicularis (Sw.) Ach. ssp. solida (Sato) W. A. Web. [Thelidea corrugata Hue] =Knightiella splachnirima (Hook. f. & Tayl.) Gyel. Thelotrema diminitum Hale, Cited specimen. Thrombium epigaeum (Pers.) Wallr. Thysanothecium hookeri Berk. & Mont. T. hyalinum (Tayl.) Nyl. Toninia bullata (Mey. & Flot.) A. Zahlbr. [T. ruginosa (Tuck.) Herre] T. sp. nov. ined. Teetristiseinesrr, Trypethelium grossum Muel1.-Arg. T. mastoideum (Ach.) Ach. T. tropicum (Ach.) Muell.-Arg. Umbilicaria arctica (Ach.) Nyl. U. coriacea Imshaug U. cylindrica (L.) Del. ex Duby U. decussata (Vill.) A. Zahlbr. U. deusta (Huds.) Baume. . havaasii Llano . hirsuta! (Sw. ) Ach. . hyperborea (Ach.) Hof fm. . kraschenimnikovii (Sav.) A. Zahlbr. . phaea Tuck. . proboscidea (L.) Schrad. subglabra (Nyl.) Harm. U. vellea (L.) Ach. qaqa, a qj, ca = [U. vellea (L.) Ach.] U. cinereorufescens (Schaer.) Frey sensu Llano Usnea angulata Ach. U. angulosa (Muell.—Arg.) Mot. U. arbusculiformis Mot. U. capillacea Mot. U. cavernosa Tuck. in Agassiz U. cf. cladocarpa Fee U. comosa (Ach.) Roehl. [U. diplotypus Vain.] U. herrei Hale. nom. nud. U. aff. elongata Mot. U. flexilis Stirt. Mexico Colorado Alaska Australia N. Guinea Tasmania Australia Colorado Australia Australia Australia Colorado Colorado N. Guinea Louisiana Louisiana Norway Norway Australia Australia Colorado Norway Colorado Colorado Colorado California Australia Australia Colorado Colorado Mexico Australia Mexico Tasmania Tasmania Arizona Mexico Galapagos Arizona Kansas Texas Galapagos Tasmania N. Guinea . herrei Hale, nom. mud. . 1gniaria Mot. . inermis Mot. . cf. merrillii Mot. . microcarpoides (Muell.-Arg.) Mot. . mutabilis Stirt. . paradoxa (A. Zahlbr.) Mot. . poliotrix Kremp. aIaI1al acl aiciacjia [U. rubiginea (Michx.) Mass.] U. rubicunda Stirt. U. scabrata Nyl. ssp. nylanderiana Mot. U. scabrida Tayl. [U. sorediifera auctt.] =U. fulvoreagens (Raes.) Raes. cf. Brodo in The Bryologist 79:406. 1976 [1977]. U. spilota Stirt. U. cf. strigosa (Ach.) Eaton U. torquescens Stirt. U. torulosa (Muell.-Arg.) A. Zahlbr. [U. tristis Mot.] U. florida (L.) Wigg., fide Asahina, J. Jap. Bot. 43:65. 1968. eae U. xanthopoga Nyl. Verrucaria laevata Ach. ‘Xanthoparmelia wyomingica (Gyel.) Hale Xanthoria candelaria (L.) Th. fr. X. fallax (Hepp ex Arn.) Arn. sens. latiss. X. polycarpa (Ehrh.) Rieb. X. polycarpa (Ehrh.) Rieb. Identification doubtful. [Xylographa spilomatica (Anzi) Th. Fr.] =X. vitiligo Ach. ) Laundon X. vitiligo (Ach. ) Laundon NEW COMBINATIONS Cladina galapagosensis (Ahti) W. A. Web. Ahti, Ann. Bot. Soc. 'Vanamo' 32(1):46. 1961. Cladina polia (R. Sant.) W. A. Web. orth. error) R. Sant., Ark. Bot. 30A(10): 15. 1952. Heterodemmia barbifera (Nyl.) W. A. Web. syn, Lich. 1:416. 1860. Heterodemmia circinalis (A. Zahlbr.) W. A. Web. Arizona Chile Tasmania Mexico Australia Texas Galapagos Australia Australia Australia Australia USSR Tasmania Tasmania Colorado Colorado Wyoming Australia Mexico Australia Australia Tasmania N. Mexico Tasmania S. Dakota Colorado Mexico Texas Colorado California Colorado Montana Cladonia galapagosensis Cladonia polia (as "pohlia”, Physcia barbifera Nyl., Anaptychia leuco- melaena var. mltifida f. circinalis A. Zahlbr., Beih. Bot. Centralbl. 19(2):84. 1905. 102 Heterodermia stellata (Vain.) W. A. Web. Anaptychia podocarpa var. stellata Vain., Acta Soc. Fl. Faun. Fenn. 7:131. 1890. Heterodemnia verrucifera (Kurokawa) W. A. Web. Anaptychia leucome- laena f. verrucifera Kurokawa, Nova Hedwigia, Beih. 6:72. 1962. VALIDATIONS OF NEW SPECTES LECANORA PSEUDOPINGUIS W. A. Web., sp. nov. Thallus saxicolous crus- taceus effusus indeterminatus rimoso-areolatus, areolis minus quam 1 pm diam ca. 0.5 mm crassis laevibus pallide citrinis vel albescentibus so- redia et isidia desunt. Cortex 30m crassus, granulis lutescentibus in- spersis, medulla fulvescens, C+ aurantiaca, P-, K-, IKI-, UV+ rubro-viol- ascens. Apothecia sessilia usque ad 1.5 mm diam margine crasso laevi vel undulato-crenulato disco plano livido pulverulento, epihymenio flavo-gra- nuloso C+ aurantiaco hymenio 50-60 jam crasso sporis 8/nae 7-9 x 3-5 um. Pycnidia cylindrica 200Mm alta x 100 sm lata margine prominenti elevato, conidiis arcuatis acicularis 15-20 ym longis. Ad saxa wulcanica praeci- pue litoralis. Thallus saxicolous, crustaceous, effuse, indeterminate, irregularly ri- mose-areolate with areoles less than | mm diam, ca. 0.5 mm thick, smooth, pale citrine to albescent, lacking soredia or isidia. Cortex 30 pm thick, inspersed with yellowish granules; medulla discolored, C+ orange, P-, K-, IKI-, UV+ red-violet. Apothecia sessile, up to 1.5 mm diam, with thick smooth or crenulate-wavy margin, the disk plane, livid, pulveru- lent; epihymenium with yellow granules, C+ orange; hymenium 50-60 Mm high, spores 8/ascus, 7-9 x 3-5 44m; pycnidia cylindric, 200 jim deep x 10 yim wide, with prominent raised rim resembling the apothecial margin; pyc- noconidia arcuate, acicular, 15-20141m long. On volcanic rocks, primarily littoral. HOLOTYPE. Ecuador. Galapagos Islands. Isla Santa Cruz, vicinity of Academy Bay, on exposed point along shore just east of Darwin Research Station, abundant on rocks just above high tide mark, 10 April 1976, Weber & Lanier, Lich. Exsicc. OOLO No. 500 (COLO). Further distribu- tion. Floreana I., Jervis I., Wenman I., S. Plazas Islet, Champion I., and probably on every island of the archipelago. One population has been found inland from the seacoast but on a seaward exposure: Santiago I., on rocks at rim of easternmost crater in the highlands, 400 msm, 10 May 1971, L. H. Pike 2112 (COLO L-55204). This is a most unusual occurrence and may be correlated with atmospheric conditions (upwelling of fog) pe- culiar to the site. L. pseudopinguis is one of the dominant species co- loring the rocks of the shoreline yellow on many of the islands. _ Except for its effuse thallus the new species resembles the lobate littoral Californian Lecanora pinguis Tuck. Dr. Chicita Culberson, to whom I am indebted for analyses of the chemistry of L. pseudopinguis and 103 the next species, reported: "'L. pseudopinguis contained a compound (Re classes A4-5, B5, C4-5; spot color with 10% H»S0, and heat is orange brown) in high proportion that is probably a xanthone pigment....We have seen this same pigment before in a sample from Ahmadjian that was also from the Galapagos Islands but that had been identified as Buellia stra- minea. The proportions of the minor products of Ahmadjian's sample were different from the Lecanora, and the Lecanora also contained a trace of thiophaninic acid and another unknow pigment (probably not a xanthone) that is common in lichens and that we have previously called "pigment SV-1". LECANORA (Eulecanora) TEXANA W. A. Web., sp. nov. Prothallus destitutus. Thallus effusus indeterminatus laevis vel tartareus non- vel indistinctissime lobatus, usque ad 2 mm crassus, irregulariter rimoso- areolatus vel continuus, pallide flavus. Pseudocortex prosenchymaticus 30-40 41m crassus ex hyphis erectis, strato algarum 30-50 yam alto, con- tinuo. Medulla densissima, alba, subtus fulvescens, K-, KC+ luteus. Apothecium immersum vel adnatum, usque ad 2 mm diam, margine vestigiali, excipulo proprio ex hyphis erectis parallelis, disco carneo, plano vel convexiusculo, epihymenio hyalino granuloso, granulis in K dissolvens, hymenio 50-60 jAm alto, subhymenio ex hyphis verticalibus, hypothecio hya- lino subtus algifero, ascis IKI+ coeruleis, paraphysibus simplicibus, 2 m diam, gracilibus, non capitatis, sporis 8-nae anguste-ellipsoideis rectis vel plus minusve curvatis, 10-14 x 4-5 wm. Pycnidium ostiolo aeruginoso loculo 200 pm diam, pycnoconidiis Eulecanoroideis acicularis, arcte curvatis vel sigmideis, 10-15 x 0.5 jam. Ad saxa granitica, areas amplas rupium superficieram verticalibus tectus. Prothallus lacking. Thallus effuse, indeterminate, smooth to tar- tareous, very distinctly if at all marginally lobate, up to 2 mm thick, irregularly rimose-areolate or continuous, pale yellow; pseudocortex pro- senchymatous, 30-40 ym thick, of erect hyphae; algal layer 30-50 m high, continuous; medulla very dense, white, becoming discolored brownish be- low, K-. KC+ yellow wash. Apothecia immersed to adnate, up to 2 m dian, the margin vestigial or represented by a few small bead-like remnants; proper exciple of erect parallel hyphae; disk flesh-colored, plane to slightly convex; epihymenium hyaline, granulose, the granules dissolving in K; hymenium 50-60 pm high, difficult to differentiate fram subhymenium of vertical hyphae; hypothecium hyaline, with continuous or broken algal layer below; asci IKI+ blue; paraphyses simple, 2 jAm diam, slender, not capitate; spores 8/ascus, narrowly ellipsoid, straight or somewhat curved, 10-14 x 4-5qm. Pycnidia appearing as black dots on thallus sur- face, the ostiolar margin aeruginose, the loculus 200 Mm diam; pycno- conidia of Fulecanora type, acicular, curved in a semicircle, 10-15 x 0.5 fam. On granite rocks, covering large areas of the vertical faces, rarely fruiting, but when fertile with clusters of apothecia in small plaques of the thallus. 104 HOLOTYPE. Texas, U. S. A. Gillespie Co. [not Llano as given on the label]: trail to 'Balanced Rock'"', a granite dome rising above the sur- rounding plain, 4 mi n of Fredericksburg in the Texas "Hill Country"; abundant and dominant on vertical N-facing massive granite blocks, 29 April 1974, Weber, Lich. Exsicc. COLO No. 451 (COLO). Lecanora texana is apparently closely related to L. sulphurea (Hoffm.) Ach., but the apothecium is pale from the beginning, never blackening. Dr. C. Culberson reported: "L. texana contained usnic acid, zeorin and a trace of atranorin. In addition there were two or three un- identified triterpenoids that are probably related to zeorin but which do not correspond to leucotylin or leucotylic acid (the only known compounds of this type in our collection). There is also a curious unidentified phenolic compound (R¢ classes A3, B2-3, C5; spot color with 10% H»SO, and heat is yellow). This may be a new lichen product." PSORA CEREBRIFORMIS W. A. Web., sp. nov. Thallus terricola creta- ceus tumulos hygroscopicos usque ad 5 cm diam formantes, squamilis dense aggregatis sed non imbricatis primo planis mox alte convexis rimosis mar— ginibus revolutis. Cortex argillaceus pseudoparenchymaticus 50-60 m crassus, strato epinecroso crasso albo tectus. Stratum gonidialium con- tinuum. Medulla alba solida crassa, coacto rhizo-hyphoso substrato af- fixo. Cortex et medulla K-, C-, KC-, P-, IKI-. Apothecia 1-2 m diam atra marginalia convexa numerosa plerumque aggregata, excipulo ex hyphis parallelis pallide ferrugineo, ascis clavatis 30-40 m longis, sporis el- lipsoideis (8)10-12(17) x 5-6 m. Epihymenium ferrugineum K+ vinosum C-, Pp. IKI. Thallus terricolous, chalk-white, forming hygroscopic convex mounds up to 5 cm or more diam; squamles densely aggregated but not overlapp- ing, up to 5 mm diam, at first plane but very soon highly convex, deeply cracked, with the margins rolled under. Cortex pale clay-colored, of pseudoparenchyma type, 50-60 m thick, covered by a thick white epinecral pruinose layer. Algal layer continuous. Medulla white, solid, thick, attached to the substrate by a rhizo-hyphal felt. Reactions K-, C-. KC-. P-. IKI-. Apothecia 1-2 mm diam, black, marginal, highly convex, abundant and often clustered; exciple of parallel hyphae, pale reddish- brown; asci clavate,30-40 m long; spores ellipsoid, (8)10-12(17) x 5-6 m; epihymenium reddish-brown, K+ vinose, C-, P-, IKI-. HOLOTYPE. U. S. A. Colorado. Montrose Co.: on gypsum knolls, floor of Paradox Valley 4 mi E of Bedrock, 1,500 msm, 30 May 1960, Shushan, Anderson & Weber, Lich. Exsicc. COLO No. 24 (COLO). MYCOTAXON WO lee AIL ORmeL ee Oye meio LY April-June 1981 CULTURAL STUDIES ON PORIA CINERASCENS,, P. RIVULOSA, AND P. SUBVERMISPORA (Aphyllophorales, Basidiomycotina) K. K. NAKASONE Center for Forest Mycology Research 1/ Forest Products Laboratory— USDA, Forest Service Madison, oWLs 5 05/05 ABSTRACT The culture complex involving Poria cinerascens, P. rivulosa, and P. subvermispora is examined. Cultural descriptions are provided for each species. Although nearly identical, cultures of these three species can be separated by temperature studies, growth on cabbage extract agar, and microscopic characteristics. INTRODUCTION Sporophores of Poria cinerascens (Bres.) Sacc. et Syd., Poria rivulosa (Berk. et Curt.) Cke., and Poria subvermispora Pilat are dis- tinct and easily identified by observing the hyphal systems and basid- iospores (Lowe 1966). However, when grown in pure culture they are nearly indistinguishable. Nobles (1948) treated P. rivulosa (as P. albipellucida Baxter) and P. cinerascens together under one culture description. Later, Nobles (1965) and Stalpers (1978) distinguished these fungi on minor or variable characters. Because of the difficulty in identifying these cultures, the im- portance of individual species of the complex in the decay of products, preservative-treated wood, and other wood forms cannot be assessed accurately. Duncan and Lombard (1965), in their summary report on Hymenomycetes associated with decay of wood products, report these fungi collectively as the Poria cinerascens complex. Thus, this study was undertaken to resolve the P. cinerascens culture complex. MATERIALS AND METHODS Cultures were grown at 25° C in the dark on 1.5% malt extract agar (MEA) in glass petri plates and were examined at weekly intervals. 1/ Maintained at Madison, Wis., in cooperation with the University of Wisconsin. 106 Cultures were grown also on 0.5% gallic acid agar (GAA), on 0.5% tan- nic acid agar (TAA) (Davidson et al., 1938), and on cabbage extract agar (CAB) at 25° C and measured at 7 and 14 days. Cabbage extract agar was made from fresh cabbage because growth | of these fungi on commercial media was not satisfactory. CAB is pre- pared as follows: Place 500 gm chopped, fresh cabbage in 1000 ml dis- tilled water, cover, and bring to a boil. Simmer about 15 hours (over several days if necessary) and add about 1,000 ml distilled water during this time. Filter through cheese cloth and bring final volume of cabbage extract to 1000 ml. Add 20 gm agar, 2.5 gm diabasic sodium phosphate, 5 gm sodium chloride, and 10 gm Difco Bacto-peptone to the 1000 ml of cabbage extract and autoclave. Unless otherwise indicated, all cultures studied are of polyspo- rous origin. Monosporous isolates were obtained from fruiting in cul- ture. Key patterns of 2-week-old cultures follow the format of Davidson et al. (1942). Species codes of 6-week-old cultures follow the format of Nobles (1965). Cultures are on deposit at the Center for Forest Mycology Research. Sporophores associated with the cultures were previously identi- fied by Drs] J: L. Lowe, Ri L. Gilbertson, or) M.J.,3Llarsen. 7 sAL though the sporophores were not reexamined by the author, dikaryotic- monokaryotic matings were done with all cultures studied. Cultures listed under each species had successful matings with one or more hap- loids of that species. CULTURE DESCRIPTIONS Poria Cinerascens (Bres.) Sacc. et Syd., Syil. Fung. 1671615 81902" Growth characters: Growth on MEA rapid, plates covered in 1 wk; mats white, appressed, thin, subfelty, with a network of thin, radial strands, sometimes slightly raised and downy or felty at 2 wk, may | develop small white balls of mycelium at 6 wk; margin even, appressed; odor none; agar discoloration none; not fruiting by 6 wk. Oxidase reactions at 1 wk on GAA moderately strong, mat 37-51 mm diam; on TAA strong, mat 12-26 mm diam. Optimum temperature 36° C (Fig. 1). Growth on CAB O-trace at 1 wk. Microscopic characters: Hyphae of advancing zone 6-8 wm diam, thin-walled, simple septate, branched, becoming thick-walled and scat- tered by 2 wk. Hyphae of submerged mycelium 2-4 wm diam, thin-walled, nodose septate, branched. Hyphae of aerial mat (a) similar to sub- merged hyphae except occasionally encrusted with hyaline crystals; (b) "binding hyphae" very slender, 0.5 um diam, aseptate, richly branched, hyaline, nonstaining in phloxine, rare to abundant at 2 wk; (c) fiber hyphae (Fig. 2) 4-6 um diam, thick-walled, walls thinning toward apex, aseptate, hyaline nonstaining in phloxine, apparently lacking or numer- ous at 2 wk. Chlamydospores globose to limoniform, 8-13.5 x 8-13.5 um, thin-walled, walls thickening in age, hyaline, terminal or intercalary, few to abundant in aerial and submerged mycelium. Bulbils sometimes present in submerged mycelium. Key patterns: A-P-F-1-2-10; A-P-F-1-2-10-16; A-P-F-1-2-11; A-P-F- 1=2-11-16; A-P-F-1-2-10-14. Species code?) 2740072 (8 )) (22 494.6238, 42-55 250. Monosporous cultures: Eight monosporous cultures of HHB 76 (4 of each mating type) examined were similar to the polysporous cultures except that they did not have clamp connections. 60 PORIA SUBVERM/ISPORA 50 40 RADIUS (mm) S 20 PORIA RIVULOSA PORIA CINERASCENS /6 20 24 28 32 36 40 44 TEMPERATURE (°C) Fig. 1. Average radial growth of Poria species on MEA after 4 days at 8 temperatures. (M 149 135) 10 um M 149 140 Fig. 2. Fiber hyphae from a culture of Poria cinerascens (FP 100506) at 2 wk. Fig. 3. "Binding hyphae" from cultures of Poria rivulosa and P. subvermispora at 2 wk. (M 149 140) 107 108 Incompatibility system: Poria cinerascens has been reported by Nobles et al. (1957) to be heterothallic and bipolar. Pairing of | 17 monosporous cultures of HHB 76 confirm their finding: Ay =01,45,0., Sells. Osos A, Se Oe Onis Pe Ole slates Cultural description: Nobles (1948); Singh (1966). Cultures studied: U.S.A.: MARYLAND--FP 100506 on pine, Prince Georges County; SOUTH DAKOTA--FP 105939 on Pinus ponderosa Dougl. ex Laws: (ponderosa pine), Black Hills National Forest, TEXAS-- HHB 76 cn Pinus sp., San Jacinto County; WISCONSIN--FP 105349 on Pinus banksiana Lamb. (jack pine), Wood County. Remarks: The development of fiber hyphae and optimum temperature of 36° C are diagnostic for P. cinerascens. Poria cinerascems can quickly be separated from P. rivulosa and P. subvermispora by growing the cultures on MEA at 40° C (Fig. 1). At 4 days P. cinerascens grows significantly more than the other species. Cultures of P. cinerascens will often deteriorate after growing several years on artificial media. Deteriorated cultures are slow growing, sodden, and do not develop fiber hyphae. Poria cinerascens is widely distributed in the U.S.A. and is associated with a white rot of conifers. Poria rivulosa (Berk. et Curt.) Cke., Grevillea 14:109. 1886. Growth characters: Growth on MEA rapid, plates covered in 1 wk; mats white, azonate, at 1 wk appressed, thin, subfelty, at 2 wk slightly raised, thicker, subfelty to felty; margin even, appressed; odor none; agar discoloration none; not fruiting by 6 wk. Oxidase reactions at 1 wk on GAA moderate, mat 64-90 mm diam; on TAA strong, mat 19-29 mm diam. Optimum temperature about 32° C (Fig. 1). Growth on CAB 20-48 mm diam at 1 wk. Microscopic characters: Hyphae of advancing zone 6-7.5 mm diam thin-walled, simple septate, branched, by 2 wk often becoming thick- walled, rare to abundant, absent in older cultures. Hyphae of sub- merged mycelium 3-5 mm diam, thin-walled, nodose septate, branched. Hyphae of aerial mat (a) 1.5-5 mm diam, similar to submerged hyphae, sometimes encrusted with hyaline crystals; (b) "binding hyphae" (Fig. 3) very slender, 0.5 mm diam, aseptate, richly branched, hyaline, nonstaining in phloxine, usually present by 2 wk in clusters, difficult to separate. Chlamydospores globose to limoniform, 8.5-11.5 x 8.5-11.5 mm, thin-walled, becoming thick-walled in age, hyaline, termi- nal or intercalary, abundant in aerial mat. Key patterns: -A-P-F-1-2-10-16; A-P-F-1-2-10; A-P-F-1-2-10-14-16; A>P-F=1]=2-L0=14% Species code: 2.4.(7).(8).34.36.38.41.42.55.59. Monosporous cultures: Eight monosporous cultures of FP 133035 (4 of each mating type) were similar to the polysporous cultures except that thick-walled advancing zone hyphae were present in 6-wk-old cul- tures, and they lacked clamp connections and "binding hyphae." Incompatibility system: Nobles et al. (1957) reported that P. rivulosa is heterothallic and bipolar. Pairings of 17 monosporous Cute tures of FP 133035 confirm their result: A) =01,35455, 9.12 15 LS, i A, 2 05 7.,6,.0. bilan Cultural descriptions: Buckland (1946), Baxter (1937), and Nobles (1948) as Poria albipellucida Baxter; Stalpers (1978) as P. lindbladii (Berk. et Br. ex Berk.) Cooke. 109 Cultures studied: U.S.A.: CALIFORNIA--Piirto 26 (rot) in Sequoiadendron giganteum (Lindl.) Bucholz (giant sequoia); JLL 10602 on Sequoia sempervirens (D. Don) Endl. (redwood), Del Norte County; FP 104207 (rot) in redwood, Wilson Creek; IDAHO--JLL 6987 on conifer, Kootenai County; OREGON--FP 133035 (rot) in Tsuga sp., Benton County, FP 133406 and FP 133696 on conifer, Lincoln County. CANADA: BRITISH COLUMBIA--DAOM 9757 on Pseudotsuga menziesii (Mirb.) Franco (Douglas-fir). Remarks: Poria rivulosa is most similar culturally to P. subvermispora. They can be separated, however, by growing the cul- tures on CAB. Poria rivulosa will grow 20-40 mm in diameter at 1 wk but P. subvermispora only O-trace. Poria rivulosa is common in the northwest U.S.A. and British Columbia on conifers. It is associated with a white ring rot of living western redcedar, Thuja plicata Donn ex D. Don (Buckland 1946) and living redwood (Kimmey and Lightle 1955). Poria subvermispora Pilacesstud iam Bote Cech 5:26 L94Gs Growth characters: Growth on MEA rapid, plates covered in 1 wk; mats white, azonate, at 1 wk appressed and subfelty or raised and downy, at 2 wk appressed, subfelty to felty, or raised and woolly, sometimes tufted; margin even, appressed; odor none; agar slightly bleached in age; not fruiting by 6 wk. Oxidase reactions at 1 wk on GAA moderate, mat 69-90 mm diam; on TAA strong, mat 22-30 mm diam. Optimum temperature 32° C (Fig. 1). Growth on CAB O-trace at 1 wk. Microscopic characters: Hyphae of advancing zone 6-7 wm diam, thin-walled, simple septate, infrequently branched, by 2 wk becoming thick-walled, rare. Hyphae of submerged mycelium 2-4 wm diam, thin- walled or with slight wall thickening, nodose septate, branched. Hyphae of aerial mat (a) similar to submerged hyphae, occasionally encrusted with hyaline crystals; (b) "binding hyphae" very slender (Fig. 3), 0.5 wm diam, aseptate, richly branched, hyaline nonstaining in phloxine, usually present in clusters by 2 wk, difficult to sepa- rate; (c) fiber hyphae rare, 2 wm diam, thick-walled, aseptate, hya- line nonstaining in phloxine, infrequently branched, seen only in 6-wk-old cultures of FP 90031. Chlamydospores globose to limoniform, 1125-13.5 (-16) x 7.5-13.5 (-16) um, thin-walled, walls thickening in age, hyaline, terminal or intercalary, numerous in submerged and aerial mycelium. Key patterns: A-P-F-1-2-10-16; A-P-F-1-2-10; A-P-F-1-2-10-14-16; eesti 2—10=14. Species codes.) 2-4, (7)-(8).34.36.38.41.(42).54.55.59. Monosporous cultures: An examination of 8 monosporous cultures obtained from JLL 14807 (4 isolates of each mating type) reveals that they are similar to the polysporous isolates except that their mats tend to be appressed, develop delicate radial to reticulate patterns, and lack clamp connections. Incompatibility system: Nobles et al. (1957), as P. notata Overh., and Domanski (1969), as Fibuloporia subvermispora (Pilat) Doman., reported that P. subvermispora is bipolar. Pairings of 17 monosporous cultures of JLL 14807 confirm their results: AL 1 ae Bros ol2, 13,15, 16,18 519; A, Seow Ln lis Lee La Cultural descriptions: Baxter (1947) (as P. quercuum Baxter); Nobles (1948); Domanski (1969); Stalpers (1978). 110 Cultures studied: U.S.A.: ARIZONA--JLL 8904 on ponderosa pine, Cochise County; COLORADO--JLL 6133 on conifer, Garfield County; JLL 6332 on lodgepole pine, Larimer County; FLORIDA--FP 104027 (sporo- phore tissue) on Quercus laurifolia Michx. (laurel oak), Duval County; MARYLAND--FP 90031 on oak and FP 105752 on hardwood board, Prince Georges County; NEW YORK--JLL 3292 on conifer, Essex County; JLL 15225 on hardwood, Ulster County; OREGON--ME 485 (rot) in Douglas- fir wood chips, Clatsop County. CANADA: Saskatchewan--JLL 14807 on Populus tremuloides Michx. (aspen). Remarks: Poria subvermispora is widely distributed throughout continental U.S.A. It is associated with a white rot of conifers and hardwoods and has been isolated several times from wood chip piles in Washington and Oregon. DISCUSSION Cultures of P. cinerascens, P. rivulosa, and P. subvermispora can be identified by noting several critical characters. These are sum- marized in Table 1. Dikaryotic-monokaryotic pairings can be used also to identify cultures once monosporous cultures are obtained. Cultures of the three species will fruit on MEA and discharge spores if kept long enough. The shape and size of the basidiospores can also be used to differentiate between the species (Table 1). Development of the slender "binding hyphae" found in the aerial mat of all three species has not been recorded previously. Their development in culture suggests their presence in the sporophore; how- ever, this is not the case. Poria rivulosa and P. subvermispora have monomitic hyphal systems (Lowe 1966). However, Poria cinerascens spo- rophores have a trimitic hyphal system (Lowe 1966), and this is sup- ported in culture. Table 1.--Characteristics that differentiate cultures of Poria cinerascens, P. rivulosa, and P. subvermispora Optimum Growth Growth BE taal growth at 1/ ony Host ate temperature 40° C-— CAB-— size— Ree oi ae REE ed thin ae my mm; ?s ero as ym BOEEG 36 To ela Gp Oo Hane rec eee - -trace Conifers aaa GUAR ee Eee 1. Ser 220 5S 0e="6.0 P. rivulosa SZ La 20 - 48 Conifers me 4, Oo=e 500 Conifers 3250 ORO P. subvermispora OZ Tae BO-L race and x hardwoods 1.0 peed ico 1/ Average radial growth after 4 days on malt extract agar. 2/ Average mat diameters on cabbage extract agar after 7 days ate 5o Se 3/ Spore measurements taken from Lowe (1966). yal Culturally, these three species belong to a unique group of fungi. The rapid growth, production of extracellular oxidases, simple septate hyphae of the advancing zone that later give rise to nodose septate hyphae, and the bipolar mating system are characters shared by all spe- cies of Group 54 (Nobles 1958). Phlebia subserialis (Bourd. et Galz.) Donk, Phlebia subochracea (Bres.) Erikss. et Ryv., and Hyphodontia setulosa (Berk. et Curt.) Maas G. are the most culturally similar to the Poria species discussed. ACKNOWLEDGMENTS Ms. Ruth G. Payne is responsible for the successful development and application of the cabbage extract medium. Critical reviews of this manuscript, by Drs.) Row. Gilbertson, H.-H. Burdsall, Jr.=, and Mrs. F. F. Lombard are greatly appreciated. LITERATURE CITED BAXTER, D. V. 1937. Some resupinate polypores from the region of the Great Lakes. IX. Pap. Michigan Acad. Sci. 23:285-305, (published in 1938). BAXTER, D. V. 1947. Some resupinate polypores from the region of the Great Lakes. XIX. Pap. Michigan Acad. Sci. 33:9-30, (published in 1949). BUCKLAND, D. C. 1946. Investigations of decay in western red cedar in British Columbia. Can. J. Res., C., 24:158-181. DAVIDSON, R. W., W. A. CAMPBELL, and D. J. BLAISDELL. 1938. Differ- ention of wood-decaying fungi by their reactions on gallic or tannic acid medium. J. Agric. Res. 57:683-695. DAVIDSON, R. W., W. A. CAMPBELL, and D. B. VAUGHAN. 1942. Fungi causing decay of living oaks in the eastern United States and their cultural identifi- cation. USDA Tech. Bull. 785, 65 p. DOMANSKI, S. 1969. Wood- inhabiting fungi of Bialowieza virgin forest in Poland. X. Fibuloporia subvermispora (Pilat) Doman. comb. nov. and its diagnose. Acta Soc. Bot. Pol. 39:531-538. DUNCAN, C. G., and F. F. LOMBARD. 1965. Fungi associated with principal decays in wood products in the United States. eos POr.e serv, Res. Pap. WO-4, 31) p.> ~KIMMEY, J. W.,' and P. C. LIGHTLE. 1955. Fungi associated with cull in red-wood. For. Sci. 1:104-110. LOWE, J. L. 1966. Polyporaceae of North America. The genus Poria. State Univ. Coll. of Forestry, Syracuse Univ. Tech. Bull’ 90, 183 p. NOBLES, M. K. 1948. Studies in forest pathology. VI. Identification of cultures of wood-rotting fungi. Can. J. Res., Co, 26:281-431. NOBLES, M. K. 1958. Cultural characters as a guide to the taxonomy and phylogeny of the Polyporaceae. Can. J. Bot. 36:883-926. NOBLES, M. K. 1965. Identification of cultures of wood- inhabiting Hymenomycetes. Can. J. Bot. 43:1097-1139. NOBLES, M. K., R. MACRAE, and B. P. TOMLIN. 1957. Results of interfertility tests on some species of Hymenomycetes. Can. J. Bot. 35:377-387. SINGH, B. 1966. Studies on Indian Poria. II. Diagnosis of five species as new records. Indian Forester 92:680-683. STALPERS, J. A. 1978. Identi- fication of wood-inhabiting Aphyllophorales in pure culture. Centraal- bureau voor Schimmelcultures. Studies in Mycology: 16, 248 p. MYCOTAXON VGieecl Lint NO.e eee Dt eee or April-June 1981 A NEW SPECIES OF AMANITA DAVID T. JENKINS Department of Biology, University of Akabama in Biuningham, BuUuntngham, AL 35294 ABSTRACT Amanita martginata (fig. 1) is described as new from Tennessee. It is assigned to Section Lepidella, Subsection Solttariae. Amanita martginata Jenkins, sp. nov. Holotype: Tennessee, Loop Road, Cades Cove, Great Smoky Mountains National Park, 17. viii. 72, David T. Jenkins 580(DTJ).* Pileus 110 mm mense diametro, convexus, margo leviter appendicu- latus, non striatus; labrum distinctum sterileque subter, album conver- tens ad gilvum colorem; reliquae volvae ita verrucae adnatae factaeque in inaequalem formam, gilvae cum cacumenibus in vero colore, potius in circlos qui easdem medias partes habent. Lamellae stipatissimae, sol- utae. Stipes 90 x 20 mm, solidus, fibrillosus squameusque adversus basem; tegumen ad apicem partim, pendens, album, delicatum; bulbus ad basem in ovi formam factus, gilvus; maculans flammeus fuscusque adversus fundum; aliquando rimae longitudine; reliquae volvae ita circli squam- arum adnatarum reflexarumque leviter, qui easdem medias partes habent. Sporae 5.46-6.25 x 7.0-10.15 um. Fruit body medium to large, solitary. PILEUS: 110 mm diam, con- vex, margin non-striate, very slightly appendiculate, but with a dis- tinct, sterile margin, up to 3 mm wide, pileipellis fairly easily sep- arable, fibrillose between volval remnants, white to creamy-white, flesh white, up to 15 mm thick at center, tapering toward margin; universal veil remnants as fibrous, adnate, irregular-shaped warts, up to 7 mm wide at base, cream with tips cream to pale tannish-cream, more or less concentric ring arrangement, becoming decidedly smaller toward margin. LAMELLAE: very crowded, free, fairly broad, cream, edges smooth; lam- ellulae numerous, attenuate. STIPE: 90 x 20 mm, solid, white, fibril- lose-scaly beneath annulus, becoming lacerate-scaly toward base; partial veil apical, creamy-white, pendant, large and thick, striate above, floccose below, double-edged, very delicate, soon falling off, ring of universal veil material approximately 10 mm below partial veil. Basal bulb ovoid, cream at apex, staining orangish-brown toward bottom, 65 x 45 mm, occasionally with longitudinal splits; universal veil rem- nants as concentric rings of adnate, slightly recurved scales, quite thick, on upper part of bulb. Faint chloride of lime smell. /PILEIPELLIS: filamentous hyphae densely interwoven, slightly gelatinized, hyaline, 2-7 um diam. PILEUS TRAMA: filamentous hyphae ar nn EES IEEEnEEeeed * DTJ = The author's herbarium ies undifferentiated, moderately branched, clamps rare, 3-8 um diam; inflated cells abundant, mostly terminal, variform, mostly elongate, up to 162 x 31 um. LAMELLA TRAMA: bilateral; filamentous hyphae undifferentiated, moderately branched, no clamps, 3-8 um diam; inflated cells terminal, elongate. SUBHYMENIUM: hyphae ramose, clamps rare. BASIDIA: up to 48 x 3.9-8.6 um, 4-sterigmate, thin walled, without clamps. UNIVERSAL VEIL: filamentous hyphae on pileus moderately abun- dant, sparsely to moderately branched, without clamps, 3-6 um diam, irregularly disposed; inflated cells abundant, globose, subglobose, broadly elliptic, elliptic, clavate, cylindric, most being quite small, not exceeding 62 x 46 um, usually short, terminal chains, irregularly disposed: tissue on basal bulb similar, but with a slightly higher percentage of filamentous hyphae. STIPE TRAMA: filamentous hyphae undifferentiated and abundant, sparsely branched, without clamps, 3-6 um diam; the presence of gloeoplerous hyphae conspicuous; inflated cells terminal, clavate, longitudinally oriented, up to 312 x 25 um. PARTIAL VEIL: upper surface mostly inflated cells, elliptic, ventri- cose, clavate, up to 125 x 63 um, mostly terminal; filamentous hyphae Sparse, sparsely branched, without clamps, 2-7 um diam: lower surface Similar to that above, but with many more smaller inflated cells, vari- form; filamentous hyphae also sparse, similar to above. Fig. 1 Amanita marginata Jenkins 580(DTJ) 114 SPORES: 6 50460-6725. 7-0-10.15eum, (Ei lecorlmoc, eye Ba51) elliptic to elongate, adaxially flattened, thin walled, hyaline, weakly amyloid, spore print white; contents guttulate; apiculus sublateral, short cylindric. Habitat and distribution: terrestrial, mixed coniferous and deciduous forest, Tennessee. Collections examined: road cut, Loop Road, Cades Cove, Great Smoky Mountains National Park, Tennessee, 17. viii. 72, David T. Jenkins 580(DTJ). Discussion: Amanita marginata is assigned to Section Lepidella, subsection Salitariae. This is due to the presence of such characters as amyloid spores, an appendiculate pileus margin, a white to cream pileus color, and variform volval elements. As attempt has also been made to assign this organism to a part- icular stirps. Following the organization of Bas (1969) this organism would seem to belong to Stirps Strobiliformis, based upon the absence of basidial clamps, white volval remnants with moderately abundant, interwoven elements, and the production of irregularly shaped volval warts on the pileus. Although this organism is apparently related to the members of Stirps Strobihiformts, only slight changes in a particular character would allow it to fit into other stirps. For example, if the basidia had clamps this organism could be placed into Stirps Vingineoides due to its small to medium, detersile volval warts and the irregularly disposed universal veil remnants, consisting of moderately abundant hyphae and inflated cells. If the elements of the volval remnants on the pileus had a more erect-parallel disposition, it would be more | closely related to Stirps Polypynramis, due primarily to its color and small to medium, detersile warts. As the Amantta flora of the United States, and in particular the southeastern region, is further studied the variation and diversity becomes increasingly apparent. As can be seen with Amanita marginata it is frequently quite difficult to determine the specific relationship with other members of this section based upon the current classification system. Therefore, the establishment of new stirps within this section may be necessary in the future. ETNERATURESCITED BaSsmce 1969. Morphology and Subdivision of Amanita and a Monograph on its Section Lepidella. Persoonia 5(4): 285-579. MYCOTAXON VO) eX ule NOM os EDD cg aLio nla Api June wLooL CANDIDA PARATROPICALIS, A NEW SPECIES OF CANDIDA JOHN G. BAKER* Analytab Products, Inc. Division of Ayernst Laboratonrtes Plainview, New York 11803 TRAS ESS OALKIN Division of Labonatonies and Research New York State Department of Health ALbany, New York 12201 DAVID H.) PINCUS Analytab Products, Inc. Diviston of Ayenst Labornatonrtes Plainview, New York 11803 RICHARD F. D'AMATO Diviston of Microbtology Department of Pathology Catholic Medicak Center of Braookkyn and Queens Jamaica, New York 11432 An unusual Candida isolated from clinical specimens has been found to be sufficiently different from existing taxa within this genus to warrant its establishment as a new species, C. paratnropicakrss. MATERIALS AND METHODS Twenty-nine isolates of the new Candida were recov- ered from specimens of human blood, sputa, urine, bronchial washings, throat swabs, a lung biopsy, and a decubitus ulcer. Their morphologic and physiologic characters were studied by procedures previously described (Baker, Salkin, Pincus #eand DraAmato, Lo Silay 9D) TAXONOMIC PART Division: Fungi imperfecti. Form class: Blastomycetes. MOrMeOrdeL sm Crypetococcales. RorvmEconihy, scl tOCOCCaceae. Candida panatrzopreatis Baker, Salkin, Pincus et D'Amato, St enOVem he LouUrem 1). *Present address: Microbiology Department, Lahey Clintc mpunl ngtonsMall, Burlington, Massachusetts, 01803: 16 Fig. 1. Candida paratropicakits on cornmeal + 1% Tween 80 agar vatters edaVseuncubati Onsate omc. Coloniae in extracti malti agaro cultae 2 mm diam usque sunt, ochroleucae farinulentae, lucidae vel subluci- dae, planae subconvexae vel umbonatae, marginibus fila- mentosis patulis. Per dies 14 in glucoso-fermenti extracto cum aqua peptonica immersis sedimentum pelliculumque formantur, cellulis fertilibus ovoideis vel elongato- ovoideis (2.5--7.5 x 3.0--12.5 um) cum quibus blasticonidia gemmantia numerosa collo angusto conjunguntur; pseudo- hyphae hyphaeque genuinae adsunt. In farinae Zeae agaro per dies 7 cultae fermenti cellulae ovoideae vel elongato- ovoideae (2.5--6.5 x 3.5--14.0 um) cum pseudohyphis hyphisque genuinis formantur. Status sexualis ignotus. Notulis physiologicis (tab4 1)\ cum. (Ci 2r0opceals (Gast. ) Berkhout congruit, sed sucrosum melezitosum fermentare necnon L-arabinosum assimilare nequit simulque methyl-D- glucosidum sucrosum melezitosumque modo variabili assimi- at. Habitat: Man. Holotype: 79MR8, isolated from human blood at autopsy, New York, New York. The holotype slide preparations, as well as living cultures, have been deposited in the American Type Culture Collection, Rockville, Maryland (accession number ATCC 42678), and in the New York Botanical Garden Culture Collection, Bronx, New York. The epithet C. parxatnopicakis was chosen to emphasize the resemblance of this species to C. tropicalis. Ley Colonies on malt-extract agar at 14 days are 2 mm in diameter; yellowish-white; pasty; glossy to dull glossy; and umbonate, flat, or slightly convex, with spreading filamentous margins. At 14 days in glucose -- yeast- extract -- peptone water a sediment and pellicle have formed; reproductive cells are ovoid to long-ovoid (2.5- Pe Oe es OK. Omi) wi tCoemuLte ple budding, blastoconidia, each attached by a narrow neck; and pseudohyphae and true hyphae are present. When grown on cornmeal agar for 7 daysymeene celisvare ovoid! to Long-ovoid (2.5-6.5 x 93.5- 14.0 um) with well-ramified pseudohyphae and true hyphae. The teleomorph is unknown. Physiologic characters (Table 1) are similar to those of C. tnopicalss (Cast.) Berkhout excepLelom(1) the inability of C. paratrzop.calrcs to fer- ment sucrose and melezitose and assimilate L-arabinose and (ii) its variable assimilation of methyl-D-glucoside, Sucrose, and melezitose. DISCUSSION The morphology of C. panratnopicakis (shape and size of blastoconidia, development of blastoconidia in sparse clusters at hyphal nodes, absence of chlamydospores and germ tubes) suggests a close similarity to C. troptcalts and its sSucrose-negative variant (Ahearn, Meyer, Mitchell, NICHOLSON wednasloreanim~, 097 /-) baker etsal., §19:31a)) sHow- ever, the new species' inability to ferment sucrose and melezitose clearly differentiates it from C. troptcalrs (van Uden and Buckley, 1970). The utilization of methyl- D-glucoside, melezitose, and sorbose by many isolates of C. panatnoprcaliss (24%, 52%, and 90% respectively; Baker et al., 198la) and its inability to assimilate inulin distinguish this taxon from the sucrose-negative form of CEeLLODL CAC CA Anearn etials, 1977). Since isolates of C. panatroprcakts frequently (41%) do not assimilate sucrose, cursory examination of such isolates could contribute to their misidentification as C. 4tellatoradea. The lack of recognition of this new species despite its relative frequent recovery from diverse clinical specimens in several laboratories suggests that C. panra- tnopteakss has been and probably still is being misidenti- fied. ACKNOWLEDGMENTS The authors express their appreciation to Dr. Rupert Barneby for preparation of the Latin translation of the HtaGnNOSIGNatOnDrSs Gi AvuandsandsGa eb. Roberts atom sup] plying some of the cultures used in this study, and especially to Dr. Michael R. McGinnis for providing taxo- nomic assistance. 118 TABLE 1. FERMENTATION AND ASSIMILATION PATTERNS OF 29 C. paratropicalis ISOLATES? Tsolates positive (3%) Substrate Fermentation Assimilation Cellobiose 0 69 Dextrose 100 100 Galactose 100 100 inwian 0 0 Lactose 0 0 Maltose 100 100 Melezitose 0 52 Melibiose 0 0 Raffinose ) 0 Sucrose 0 59 Trehalose 100 100 D-Arabinose 0 L-Arabinose 3) Cierrceacia 100 Dulce 0 Brevcnienco | 0 Ethanol 100 Ethylamine hydrochloride 100 Glucitol 100 Glycerol 14 THOS oll 0 2-Keto-gluconate 100 DL-Lactic acid 86 Mannitol 100 Methyl-D-glucoside 24 Potassium nitrate 0 Ribitol 100 Ribose 10 Rhamnose 0 Salicin 24 Sorbose 90 Succinic acid 100 Xylose 100 SS Results obtained with Wickerham broth technique at 3 weeks. LITBERATURESECLTED Ahearn, D. G., S. A. Meyer, G. Mitchell, M. A. Nicholson, and A. I. Ibrahim. 1977. Sucrose-negative. variants of Candadaltuopi. calc so. Clin Microbiol. 5: 494-496. 119 Baker MU lmG Wales! Soa ki nD tage LnCuUS and Reg ks: D'Amato. 198la. Diagnostic characters of an atypi- CamCanalid eC line Mi CroObLol. (in press). Dakar Gomer moa Lk in De ieee 1 NGUS jj eand Re be D'Amato. 198lb. Use of rapid auxanographic proced- ures for recognition of an atypical Candida. J. GloneeMLeroo.O bm ULE GOSS Jr. van Uden, N., and H. Buckley. 1970. Genus 2. Candida Berkhout, peo9 3206 /ee inn) seLoddersed.)7,) The yeasts, a taxonomic study, 2nd ed. North-Holland Publishing Co., Amsterdam. MYCOTAXON Wioiliee Xe NOt alee DOeel 20 bce5 April-June =voSL ca. Eee , NOTES ON CORTICIACEAES CBASIDIOMYCE Geapeav tke Two new species of Tubulicrinis. Kurt Hjortstam Malaregatan 12 S-441 35 Alingsas, Sweden SUMMARY Tubulicrinis cinctoides and |l-povalisporus are described from North Europe and Africa respectively. Jhe first species is similar Ome CUnCLUG eG sh CUM anu trae Ll ihinbe Cmy its more globose spores and cystidia of di- vergent appearance. The African species is related to the I. glebulosus-—qroup in its ecystidial morphology, but well defined py having oval spores. The type of T. cinctus was studied. LUBULITCRINT SS CINCTOLIDES Hjortst. nO Vass eC usu Commer or. Species Tubulicrini cincto affinis sed sporis oblique sub- grobosis @4.75—)5 x0 (8.75-)4-4 5ovums el yocysGuci tom nds — stinete capitatis, 60-/70(-90) x 5-6 um. Holotypus: Sweden. Ostergdtland. V. Tollstad par., Omberg, Storpissan state forest reserve, in herb-rich Picea forest, on ‘decayed trunk of Picea abies. 1979-11-09. K Hjgortstami& ie ad en gbac kt.weH jen clea ben Ge). Paratypus: ‘do. Hjm 11393 (priv.herb.). Rruitbody resupinateswetfuse, atetirstyreti culate mtLnen more or less continuous, (thin, whitish «Wi che thescysttaia projecting, each with a globule of excreted matter easily seen under a lens (50-100 X). Subiculum very sparse, composed of thin-walled, more or less uniform hyphae, about 2-2.5 um wide, with a clamp at each septum. Lyocystidia faintly amyloid, mostly greyish in Melzer’s reagent, cylindricai, up to 60-70(-90) um long and 5-6 um wide near the bi-, seldom tri-furcate base; the capillary lumen ending more or less abruptly; neck-width (3.5-)4-5 um, the apices of the cystidia slightly expanded but indistinct- ly capitate, usually not more than 6-7 Um*in diam. Basidia small, 12(-15) x (4.5-) 5 (-6) um, with four sterig- mata and a clamp at the base, not amyloid. Spores obliquely subglobose, thin-walled, with neither amy- loid non eyanophilous geaction, (4.75-) 5, x) (oe7>— ea 4 Atal | 5 um e. ee | 5 um -— 5 um oO 09) Fig.1. Tubulicrinis cinctoides a) cystidia b) basidium ¢) spores. = Coll. Hjm 11348 (holotypus). T. cinctus d) hy- pha,e) ecystidium f) spotes.’— Coli. Cunn. 17428 (holotypus). um, with a distinct apiculus. Remarks. The new species T. cinctoides is undoubtedly very Sina late Lomlomc nc tuseG.H.CUnn a bDubesSeemsmro beswel igide li — mited by its more globose and slightly larger spores. In the type of the latter (as far as known the only material gathered) the spores are more or less ellipsoid and measure WA Ani a1) Xe2 1 9-5 62503) UM (Cine thesoriginel. descrip= tion 4-425 x 3-5.5-um), while most spores of 1. cinctoides measure 4.75-5 x 3.75-4 um. The lyocystidia of the two spe- cies show a slight but clear difference. In T. cinctus the apical bulb is fairly distinct, starting somewhat ‘abruptly from the neck (fig.1,e), and thick-walled in the basal part, while the cystidial-bulb of the new species is less marked and usually consists only of the thin-walled part of the apex (fig.1,a). 122 7 — Sym sae Ne b ite oN j—-__| C) 5 um o Cc) C a Fig.2. Tubulicrinis ovalisporus a) cystidia b) basidium c) spores. - Coll. Ryv. 11308 (holotypus). TUBUPTCRENTLOSOVAE FSRBURUSEHYOrtGte n0V .mSDe.c. mn 1 mecr. specs ese TuUbULICrInIe Gglebulosomaltinis esedmiructincatLtone subtiliter odontioide et sporis ovalibus 4.5 (-5) x 2.5- 2.75 (-3) ums; lyocystidiis cylindricis, pallide amyloidibus (fere griseis), circiter 100 x 5-6 um, versus apicem levi- ter angustum. Holotypus: Africa. Malawi. Southern Prov., Mulanje distr., Mulanje Mts., Lichenmya Plateau, alt. 1800-2000 m.a.s.l., on coniferous wood, probably Widdingtonia. 1973-03-09/10. L. Ryvarden 11308 (0). Isotypus: in GB. Panatvypusi do wcll. Any Varden. o0 2mula. 123 Fruitbody resupinate, effuse, thin, composed of very small aculei, about 15-20/mm, whitish or becoming yellowish brown, with numerous cystidia projecting above the hymenium and easily observed under a lens (50-100 X), each encrusted at the apex with brownish, excreted material. Subiculum thin, mostly inconspicuous, composed of thin- walled, uniform hyphae with clamps, usually 2-3 um wide. Lyocystidia numerous, cylindrical, faintly amyloid, about 100 um long and 5-6 um wide near the bi-furcate base, ta- pering slight towards the apex and with a neck-width of 4-5 um, the capillary lumen ending gradually at the slight- ByeLAape lung DULeOLULUSe, aplCal part. Basidia usually 13-15 x 4.5-5 um, with four sterigmata and with a basal clamp, not amyloid. Spores ovate or ellipsoid, thin-walled, 4.5 (-5) x 2.5-2.75 (-3) um, with neither amyloid nor cyanophilous reaction. Remarks. Owing to the cystidial morphology this new species seems to be closely related to T. angustus (Rog.& Weres.) Donk and T. glebulosus (Bres.) Donk but is distinguished by MvuomOovatemuomellipsoi1dmespores. shes tinely sodontroid 7 ructi-— fication is somewhat unusual in other species of the genus, but recently Jilich described one species (T. corneri) WO iCnem according tomthesdescription.aismquitesclosemto I: hamatus (Jacks.) Donk but distinctly toothed, with fairly Largeeaculei ma nmnuresnamaverta le mastmilaregrowth suructure can be demonstrated for example in T. inornatus (Jacks. & Rog.) Donk,where it is less pronounced than in T. ovalispo- rus. Acknowledgement The author is grateful to Carl Stenholm’s foundation for supporting the investigation of the Storpissan nature re- serve. In a subsequent paper the region of Omberg will be Breatedumore thoroughly. References Cunningham, G.H. 1963. - The Thelephoraceae of Australia and NeWweeleal land. N./@.mUeD.) SCI ..51NGUSEI. RES DU LI. 145. pp. 142-143. Donk, M.A. 1956. - Notes on resupinate Hymenomycetes - III. Fungus 26: 3-24. Jackson, H.S. 1948. = Studies of Canadian Thelephoraceae. |. Some new species of Peniophora. Can. Journ. Res. Ep Pa 45S) SeWateh Ue Jilich, W. 1979. - Studies in resupinate Basidiomycetes - VI. On some new taxa. Persoonia 10(3): 332-333. MYCOTAXON Vole DieweNO wel ep pee le 4126 Apral-June sy lost BE ee ee eee NOTES ON CORTICIACEAE (BAS EDIOMYER TES aie Three new combinations in Hypochniciellum. Kurt Hjortstam Malaregatan 12 S-441 35 Alingsas, Sweden SUMMARY The genus Hypochniciellum with the type spe- cies Leptosporomyces ovoideus Jiilich was de- scribed by Hjortstam and Ryvarden (1980) and characterized by pellicular to membranaceous fruitbodies and thick-walled, cyanophilous spores. Three new combinations are proposed here: H. cremeoisabellinum, H. molle, and H. subilla- queatum. The generic circumseription is slightly emended to include also species with spores greyish but not distinctly blue in Mel- ZeOT Seereagen ti. INTRODUCTION When Eriksson and Ryvarden (1976) treated the genera Lepto- sporomyces and Leucogyrophana they discussed several simi- larities between Leptosporomyces ovoideus Julich and Leuco- gyrophana cremeoisabellina (EitschsomPart.y. ame hbicmcenm Parm., and L. subillagueata (Litsch.) Jiilich. The authors also expressed) theypossibility of joinindgmechesspeci ese nea separate genus. Later on Ginns (1978) circumscribed the genus Leucogyropha- na and excluded the three species mentioned above without any suggestions as to their generic position. Consequently, the species needa place in the family Corticiaceae, and in my opinion Hypochniciellum seems to be suitable. Without any doubt the thickness of the spore-wall is a cha- racter to consider, uniting the species. In spite of the fact that the spores of H. ovoideum are non-amyloid in com- parision with the other species, which have a more or less greyish reaction, the species have many characters in comm- on The genus Amyloathelia Hjortst. (1979) is in many respects not appropriate for the species here referred to Hypochni- ciellum. All species in that genus have strongly amyloid spores and thinner spore-walls. WAS) Key to the species’ of Hypochniciellum 1. Cystidia present, basal hyphae with thickened Vici isreme OO tae — OMI MEW. Cis tad. Gets lade teers te sei sede s «cs Soe OL Le 1. Cystidia absent, basal hyphae thin-walled We Mey CSS) Ueto s Coho 6 Sh COD.S 6 tas DOI ae 2 POD OLE Sa Oli 7a UMenl OL Oiev.t-tsloretet o-b3h01m.) which is compatiblewwithethervaria] tion observed in the living isolates. This similarity in conidial size clearly supports the conclusion that C. banttanum and C. trtechotdes are conspecific, even though the conidia in the exsiccatum tend to be larger than those in the living cultures. There is no doubt that Banti's isolate had the ability tOmerowsat3/°Gu(0).n Inetact, Bantisroundstiataniomrungus was pathogenic for rabbits (7), whose body temperature is approximately 40°C. Since there is no living culture of Banti's isolate, it is impossible to determine whether it could grow at 42-43°C. Borelli (4) demonstrated that isolates of C. banttanum can grow at this high temperature. The thermotolerance and neurotropism in man of the original isolates of C. banttanum and C. trtchotdes add additional support to the conspecificity of these two taxa. Another controversial point is the reported color of the human brain lesions. According to Banti's description (1), the cerebral nodules were ..."of a deep brown color, giving one the impression of a melanotic sarcoma." Ac- cording to Emmons et al. (5), the lesions caused by C. trtchotdes occur as gray abscesses. Judging by what is published in the literature, this aspect of the disease is variable. For example, in a Venezuelan case (2), the lesions were described as gray-blackish. In the necropsy protocol of Banti's case, which was reproduced by Stigliani (7), the lesions were described as a ..."dirty gray... from which a thick, mucous thread-like, grayish-greenish liquid came out...". The latter text was most likely dictated by Banti during the performance of the necropsy. Since the color of the lesions could have darkened within a few hours, this may have resulted in the different descriptions of the lesion color. The contention by Emmons et al. (5) that the pathological aspects of the two infections are Significantly different enough to maintain C. banttanum and C. trichotdes as separate taxa is unjustifiable. 155 It is apparent that C. trichotdes and C. banttanum are conspecific. The arguments put forth by Emmons et al. for maintaining these as separate taxa are not compelling. Cladosportum banttanum can be characterized as growing at temperatures up to 42-43°C and by producing long, sparsely branched chains of blastoconidia consisting of as many as 35 or more conidia. The conidia are smooth, 1l-celled (rarely 2-celled), oval to oblong-ellipsoidal, 3.0 X 6.4um (1.8-7.3 X 3.7-14.6um), and pale brown. The conidia arise from hyphae or simple septate conidiophores that are pale brown in color. The nomenclature is: Cladosportum banttanum (Sacc.) Borelli, Riv. Anat. Patol. OnGol ear Oo lOnmeLo OUR =EOMU LG EDC LLAMA ECACC ANIM Y.COlem LO: O20 mL ole (basionym) = Cladosportum trtchotdes Emmons in Am. J. Clin. Pathol. 22:540-541, 1952. ACKNOWLEDGEMENTS The authors wish to thank Dr. Donald P. Rogers for reviewing our manuscript, Professor Sergio Chiesa for loan of the herbarium material maintained in the Saccardoan Herbarium, and Dr. Dennis Dixon for loan of his slide culture preparations and living cultures. LITERATURE CITED ame baiGinmG alo lle SOpLasuneCcasOndisO1td omicosiscere— brale. Atti Accad. Med.-Fis. Fiorent. p. 49. Mo yey se lLOLUS es Soprasun -caso7dis0idaomicos: cerebrale. Lo Sperimentale 65:341. ae Barletta.) wandeA.sAnCULOMOLLe Cameo lame GlLados sporiosis profunda. Mycopathol. Mycol. Appl. 15:422-428. 3. Bintord Ge. so R.th-) lhompson, M. ha sGOtnameand. G- W. Emmons. 1952. Mycotic brain abscess due to Cladosporium trtchotdes, a new species. Am. J. Clinweratiolse22:D30—542. 136 Borelli, D. 1960. Torula banttana, agente di un granuloma cerebrale. Riv. Anat. Patol. Oncol. yO loaO22. EMMOUGS ComWeG emi DLULOLG me bem UCZ moni cme. Kwon-Chung. 1977. Medical Mycology. 3rd. Ed. Lea and Febiger, Philadelphia. pp. 471-482. Saccardo, Py. A. 1912.) Notae mycologicae. lie fungi ex Gallia, Abyssinia, Japonia, Mexico, Canada, Amer. bor. et centr. "Anns Mycol .910:510—-3227 Stigliani, R. 1961. Particolarita istologiche della torulosi encefalica nei preparati originali Bantiani della prima Osservazione conosciuta. Arch. Vecchi, Firenze 36:329-343. MYCOTAXON Foleex Ube aNowl le ppenl 572144 April-June 1981 A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA: PART 2, HYALOSCYPHACEAE SUBF. ARACHNOPEZ1ZOIDEAE* RICHARD P. KORF Plant Pathology Herbarium, Cornell University benacass Newmy OFkw1 GS 538USA "Much like a subtle spider which doth sit In the middle of her web, which spreadeth wide." Sir John Davies THE IMMORTALITY OF THE SOUL Order HELOTIALES suborder HYMENOSCYPHINEAE Family HYALOSCYPHACEAE Nannf. 1932 Subfamily ARACHNOPEZIZOIDEAE Korf 1978 Tribe Arachnopezizeae Nannf. in Korf emend. Korf 1978 ONE KNOWN MACARONESIAN GENUS ARACHNOPEZIZA Fuckel 1870 emend. Korf 1952 Key to the known Macaronesian species 1. Ascospores 5-7-septate at maturity. 3. A. aurata 1'. Ascospores O-1-3-septate at maturity ...--..seseee- 2 2(1'). Ascospores O-1(-2)-septate at maturity; apothecia on husks of chestnut (Casta- gteks))) “Welbbaety Gkoo gs oongupocdon.: 6 Dou oGo ue 3 2'(1'). Ascospores mostly 3-septate at maturity; apothecia on wood, leaves, duff, acorns 3(2). Hairs thin-walled, tapering, usually 1l-septate, * This flora will appear in an irregular order, with Part 1, containing the introductory material, one of the last to be published. Reprints of individual parts will not be a- vailable. References will appear in the final part. For a geographical definition of Macaronesia and other special features of this flora, see the note on page 144. 138 not constricted at the septum. 1. A. aranea f. aranea 3'(2). Hairs thicker-walled, multiseptate, constricted Lnemseptar 2. A. aranea f. monilipila 4(2'). Apothecia 1-3 mm _ diam., ascospores 1LOR2=—135 2(= 162 le Kae? 2-3 amie 4. A. aurelia ii( 2" j)2esApothecia, less, than O.5emmedian sare 5(4'). Apical cells of hairs without cyanophilic con- tents; ascospores (13.5-)18.3-22(-26) x 2.9-3.7 ym; paraphysis apices variously deformed. 5. A. obtusipila 5'(4'). Apical cells of hairs with cyanophilic contents; ascospores (14.6-)16.1-25.7 x 2.2-4.4 ym; para- physis apices filiform, not deformed. 6. A. zonulata 1. Arachnopeziza aranea (De Not.) Boud., Icones Mycol. Expl. Pl., sér. 3: 2. 1906 forma aranea. = Lachnum aranea (De Not.) Lindau, in Engler & Prantls Nat. Pflanzenfam: 11 (130)- ADB. kelley Arachnoscypha aranea (De Not.) Boud. ex Den- nis weMycolsg Ra pe o288o me lo49.. tl RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978), Korf (1952). PREVIOUS MACARONESIAN RECORDS: None. SA, TYPE LOCALITY: Italy. Qa | KNOWN MACARONESIAN DISTRIBUTION inks be reo A ae eee Madeira. CUP-MM 1542. ’ ’ CANARY ISLANDS. yeni 1542 a ua La Palma. CUP-MM 870. SUBSTRATA: On cupules (burrs) of Castanea _ sativa. Notes: I saw one 2-septate ascospore in my mounts (il- lustrated), but typically the spores are either continu- OUSS.Or sl-septate- Probably common wherever chestnut grows, but difficult to find among the hairs on the in- 39 eressUlil aces ol (hemmspinyeenusks. wal cannot jagree with Dennis, who separated this species in a distinct genus, Arachnoscypha Boud., of which it is the type species, but which Boudier himself later abandoned. In my earli- er monograph (Korf, 1952), I missed Lindau's transfer to Lachnum, which should be added to the synonymy eivienmthere. 2. Arachnopeziza aranea (De Not.) Boud. f. MONILIPILA KOGieaeieeEnhOVc PREVIOUS MACARONESIAN RECORDS: None. A Arachnopeziza aranea f. aranea pilis multisep- tatis miniliformibus dif- Gent. 0 Differing from the type cular Se Ee form in having multi- (} septate rather than 1- TM Lene SeptateCmelallS,mrands =bDy the constrictions at the septa, giving the moni— TiO Gime S DCC Umma tLOmmaLhe hairs. ae es A. aranea f. moni- lipila, ascus, pa- raphysis apex, 7 spores, subi- Oe CliG ah mn OU mesh mer OO Cl aml) mellelile Delta mm aMc Kohn, on cupules of Castanea sativa, Castanea grove below Pousada Vinhaticos, Madeira, Portugal, Wie Oe KNOWN MACARONESIAN DISTRIBUTION MADERIA. Madeira. CUP-MM 1545 (holotype). SUBSTRATA: On spiny burrs of Castanea sativa. Notes: The asci arise from repeating croziers, and the éctal excipular cells are quite large. The new form may represent merely a growth-stage of the species. 3. Arachnopeziza aurata Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 304. 1870. RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978), Nhat. (AESIEVAIh 140 PREVIOUS MACARONESIAN RECORDS: *Dennis) corals (19/7). iY Pe BOCAS TYee Germany KNOWN MACARONESIAN DISTRIBUTION LV oy Dai ie Vic CANARY ISLANDS. Gomera. CUP-MM 1348, 1368(TFC). Hierro. CUP-MM 1446(TFC), 1450. La Palma. CUP-MM 898. Tenerife. CUP-MM 223, 230, 246, 256 ec0C CUE), eacD Ube )e Sele mC, SykAg Wasrl * AZORES. *Terceira. “Dennis & al., Ws SUBSTRATA: (On pwooduand: thunk:of Briccl nates manent arborea, on bark of Myrica faya, wood Ao GS2 Dele ; SPS FTO : CUP-MM 1145, of Castanea sativa, and undetermined Saisie decorticated wood and bark. Notes. This is surely the most common species of the genus in Macaronesia, as it tends also to be in North Americar andein ws burope. 4. Arachnopeziza aurelia (Pers. : Fr.) Fuckel, Jahrb. Nassauischen Vereins Naturk. 23-24: 303. 1870. RECENT TAXONOMIC TREATMENTS: Dennis (1949, 1978), Korf (1952). PREVIOUS MACARONESIAN RECORDS: None. iY Bre EOGAL IY =r cance. H KNOWN MACARONESIAN | DESERT BUss LON CANARY ISLANDS. A. aurelia, 7 as- Hierro. CUP-MM 1459 (OSC, cospores, CUP-MM eG F459.) =x 1000; SUBSTRATA: On fruits and husks of Castanea sativa and other vegetable debris. Notes: In North America and Europe this is most common on cupules and leaves of Quercus. 141 5. Arachnopeziza obtusipila Grelet, Amateur Champignons Olt) see corme oO eneemende Kot aMycologiaw43:@213% 1951. RECENT TAXONOMIC TREATMENTS: oul O51 O52). PREVIOUS MACARONESIAN RECORDS: *Baagge & al. (1972) as "inopercu- ate discomycet.. fer LOCALITY = Erance. KNOWN MACARONESIAN DISTRIBUTION *MADEIRA. *Madeira. *CUP-MM 2502(C). SUBSTRATA: On _ decorticated wood of Pinus pinaster. Notes aul ame inagebted™ tothe stu— dents of the University of Copen- hagen who collected this (Baagge baal. ,o/2)8 ands for calling “the Specimenmto my attention for iden— tirication. A. obtusipila, as- cus, paraphysis, 5 ascospores, hate. CUP-MM 2502, xeeLOOO: 6. Arachnopeziza zonulata (Rolland) Boud., Hist. classi- He, Ghicteleynyen Wiley jig We. ISON RECENT TAXONOMIC TREATMENTS: Korf (1952). PREVIOUS MACARONESIAN RECORDS: None. TY PEMLOGCALITY: Corsica (France). KNOWN MACARONESIAN DISTRIBUTION CANARY ISLANDS. Tenerife. CUP—-MM 1222. SUBSTRATA: On wood of Pinus canariensis. Notes: In the absence of type material, I (Korf, 1952) Btreated this as a doubtful species. If I am correct in interpreting this as Rolland's species, it is only criti- 142 cally distinct from A. obtusipila. Malencon and Bertault (1958) proposed varietal status for Grelet's species un- der A. zonulata, but their combination was not validly published. I have made 3 trips to Rolland's type locali- ty near Corte, in Corsica, but did not find any species matching his description. A. zonulata, section through the apothecial margin with 2 hair bases, 4 ascospores, paraphysis apex, ascus with J+ pore, 3 hairs in cotton-blue, CUP-MM 1222, x 1000. 143 Tribe Polydesmieae Korf 1978 ONE KNOWN MACARONESIAN GENUS Pol DEOMIAREBOUdTCH am BULlA es soOC. se Mycol. = Prance™ 1- 9113. 1885. Key to the known Macaronesian species Pm ScOSpores al—3—septate sat mmaturity,.9.0-13.5 x 2.2— Be dh pola 1. P. fructicola 1®. Ascospores 3-septate at maturity, (13.9-)14.6-20.5 (-22.0) x 3.7-5.1 pm. 2. P. pruinosa 1. Polydesmia fructicola Korf, Mycotaxon 7: 475. 1978. RECENT TAXONOMIC TREATMENTS Kort CUSH Ee PREVIOUS MACARONESI- AN RECORDS: “Konrf (1978), +tBeltrdn Tejera (1980). ree LOCAL LT.Y: Ma- deira. KNOWN MACARONESIAN DISTRIBUTION *MADEIRA. *Madeira. *CUP- Uf MM 1504 (holotype) 7 (TFC), *1533, *1551, Vv *1556, *1618, *1619, #2333, *2361, *2362, — Dini San 2200 ea 2c9 | *CANARY ISLANDS. *Tenerife. *CUP-MM ae 1306. SUBSTRATA: On pedun- P. fructicola, ascus with J+ cles, pods, twigs and pore, ascus with spores, para- wood of Acacia spp., physes, 4 upper ascospores, bark and capsules of CUP-MM 1504; 4 lower spores Eucalyptus spp., un- CUP-MM 2361; all x 1000. mdetermined bark . and branchlet. 144 2. Polydesmia pruinosa (Jerd. in Berk. & BreveeBoudien, Bull. Soc. Mycol. France 1: 113. 1885. RECENT TAXONOMIC TREATMENTS: Dennis (1978), Korf (1978). PREV LOUS MACARONESIAN RE- CORDS: *Korf (1978), t+tBeltran Tejera (1980). KNOWN MACARONESIAN DISTRIBUTION *MADEIRA. *Madeira. *CUP-MM 1481(TFC), 1627) ay (OL ee LOO *CANARY ISLANDS. *La Palma. *CUP-MM 639 (le Ga ee / OO. SUBSTRATA: 2On = pyrenomycetes, on pyrenomycetes immersed in wood, on branch of Acacia sp. SPECIAL FEATURES OF THIS FLORA Since, Party 1; thes introductory »material, 1s to appear later, one reviewer of this part confessed he did not know where Macarone- sia is, and that he could not find it in any of the atlases he consulted! Thus: Macaronesia: the Atlantic island chain comprising the archipelagos of the Azores, Madeira, Salvage Islands, Canary Islands, and Cape Verde Islands. Etymology: from Greek, uakapos (fortunate) + vfigcos (is- land): "Les Tfles Fortunes," 'The Fortunate . Islands."" (Want to guess how often these P. pruinosa, 4 eee papers will be cited as Macronesia instead spores, paraphysis of Macaronesia?) apex, CUP-MM 1481, Substrates follow O. Eriksson, A. Hansen SC LOO. & P. Sunding. 1974. Flora of Macaronesia. Check-list of vascular plants, 1974. Uni- versity of Umea, Sweden. iv. + 66 pp. Distributional symbols: Papers with previous Macaronesian records citing spe- cimen data are indicated by one or more asterisks (* ** ***) and the cor- responding specimens examined have been coded to those asterisks with their CUP-MM numbers; where such specimens cannot be located or were not avail- able on loan, the symbol ''n.v." follows brief specimen data. Papers citing taxa without giving specimen data, or merely repeating previous records, are indicated by a dagger or daggers (t+ tt ttt). Herbarium abbreviations are those of the Index Herbariorum, ed. 6, and up- dates appearing periodically in the journal Taxon. Without the generous financial assistance of the National Science Foundation (Grant DEB75-23557) this preliminary flora would never have been undertaken. af MYCOTAXON WOR. XLT GNow Ll, pp. 145=149 April-June 1981 A PRELIMINARY DISCOMYCETE FLORA OF MACARONESIA: PART 3, HYALOSCYPHACEAE SUBF. TRICHOSCYPHELLOIDEAE* LINDA M. KOHN Department of Botany, Clemson University GlemsonwaeoouthsGaro linay79631,.USA and Plant Pathology Herbarium, Cornell University Tthacas. New oYork 14853 USA "A Hair perhaps divides the False and True." Omar Khayyam (tr. Edward Fitzgerald] RUBAIYAT, Stanza 19 Order HELOTIALES Suborder HYMENOSCYPHINEAE Family HYALOSCYPHACEAE Nannf. 1932 Subfamily TRICHOSCYPHELLOIDEAE Nannf. 1932 ONE KNOWN MACARONESIAN GENUS LACHNELLULA Karsten 1884 emend. Dennis 1962 Key to the known Macaronesian species 1. Apothecia white, with a bright orange hymenium, on living trees and cut ends of stumps of Pittospo- rum spp. 1. L. pittospori subsp. azorica ier Apothecia ~ogreenish-glaucus ' or grey ‘to’ nearly BIS gononéocodsocaeto pd Oans Ao Sabon Coan oon anh fs 2(1'). Apothecia grey to black. 2. L. pulveracea 2'(1'). Apothecia greenish-glaucus, strongly pli- Sater 3. L. viridi-glauca * The parts of this flora, under the editorship of Richard RB. Korf, will appear in irregular order. Reprints of indivi- dual parts will not be available for distribution. 146 iz Lachnellula pittospori Kohn, Mycotaxon 12: 278. 1980, subsp. AZORICA Kohn, subsp. nov. RECENT TAXONOMIC TREATMENTS: Kohn (1980). \ PREVIOUS MACARONESIAN RECORDS: None. Cum subspecie typica Lachnellu- DIN \\ lae pittospori omnio conveni- ens sed pilos erectos apice tu- midos (in subspecie typica non praesentes) ad marginem prae- bens. Hi pili ex cellula apica- et cellulam alteram supra eam primam formantem. Huius subspe- ciei ;novae ,distributio jam con— nita ad Insulas Azoricas limi- tata est. Holotypus: CUP-MM 2140. li saepe pariunt processum fi- \ liformem aliquando tumescentum y SS S55 L. pittospori subsp. azo- rica, 5 smMarcinal hairs, 3 ascospores, CUP-MM DIAZ. xe LOOGR Agreeing with the _ descrip-— tion of L. pittospori subsp. pittospori in all respects ex- Cepl, Lon ney presence solerc— rect, “hairs with iswollen vapices vatwihewmaroinwenotspLe— sent 1m) suUDSp. #=pltlLospori.s | bnese® ahains se oLvenmaproctce from the apical. cell a filiform process, which in some cases “swells to form “a second swollen sicell vabovemsthe first. The distribution of the new subspecies is so far as known limited to the Azores Islands. The type sub- Species was described from Bermuda. HOBO. Bem geal short mals cM meh Oils @ NoPE KOLO men ny mR OS ss Man aeOle. ctUMDuL Ot meh tTLOSDOGUINGmS Dt mash | Det ha mm mene ill 2m Lombas north of Caveira, Flores, Azores, ROctug a lye tos iv.1978. (CUP-MM 2140.) KNOWN MACARONESIAN DISTRIBUTION AZORES: Flores. CUP-MM 2138 (TFC), 2140 (holotype). Terceira. CUP-MM 2009 (TFC). SUBSTRATA: On Pittosporum spp., (stumps, cut areas of living trees, and rotted wood of standing trees). 147 2. Lachnellula pulveracea (Alb. & Schw. : Fr.) Den- nis perersooniar 222154281962" RECENT TAXONOMIC TREATMENTS: Dennis (1949), Héhnel M917). PREVIOUS MACARONESIAN RECORDS: *Dennis & al. (1977). ive eLOCALIDY: Germany. KNOWN MACARONESIAN DISTRIBUTION BA ZORES: *Terceira. *CUP-MM 1681(K), 2011, 2382, *Dennis Sra ees 1 Viel Sane MADEIRA. Madeira. CUP-MM 1560, 1621, 1623. CANARY ISLANDS. Gomera. CUP-MM 1343, 1354(TFC). Hierro. CUP-MM 1424, 1444(TFC), 1477. La Palma. CUP-MM 825. Tenerife. CUP—MM 87, 248, 538, 1212(TEC). SUBOERATA: On twig. of Aca— cia sp., of Ilex perado sub- Sp. platyphylla, of Ulex eu- (i ropeus and Ulex sp., on Stemivor Rubus®’sp-, ~and “on eractermined "twigs, “bark, wood, cut stumps, branches and roots. Notes: This distinctive spe- cies, collected on various woody substrates, is char- acterized by apothecia that ? turn violet in KOH and red- aly piche ineiodine, was noted [N\Q by Dennis (1949) and by Héhnel (1917: No. 1020). As |, pulveracea, ascus, mohnelweraiso: noted, “often nvei hair in KOH- ; paraphysis, the asci are immature. The phloxine-glycerine, hair apothecial structure is typi- in lactic-blue (CUP-MM cally Trichoscyphelloideus, 87); 5 upper spores mitnieed completely. celatini— (CUP-MM 1477); 5 lower zea’ long—celled ‘éctal excipu-— spores (CUP—MM- 1354); lum, unlike that in Hyalo- Allee LOO = scyphoideae. In lactic acid- 148 cotton-blue the granulations on the hairs are no longer visible. Raitviir (1970) treated it as a doubtful or ex- cluded species of the genus Lachnellula, and Dharne (1965) did not recognize it in the genus. Rehm's Ascomy- ceten No. 1580, issued as Dasyscypha coerulescens var. dealbata Rehm, was examined and found to be conspeci- fic, confirming Dennis's (1949) synonymy. 3. Lachnellula VIRIDI-GLAUCA Kohn, sp. nov. PREVIOUS MACARONESIAN RECORDS: None. Apothecia gregaria vel fasci- culata, viridi-glauca, sessili- a, piulis -albissvestita se protun— de cupulata, receptaculi parie- tibus penitus plicatis praeser- tim in apotheciis immaturis; re- ceptaculum O.25-0.50 mm in Gram.) scOommutatio. xanthochrouca in 2% KOH eveniens. Excipulum medullare “ex textura intricata hyalina formatum, ss ce lu visee2—3 L. viridi-glauca, hair, ascus and paraphysis, me latis. WExcipudum = ectalege ex @ textura oblita formantum hyali- CUP-MM 850, x 1000. na intertexta in gelatina copiosa flava contenta, cellulis 3-6 pm latis, exterioribus pilos edentibus. Pili hyalini, erecti, septati,-grosse granulosi, 35=45 x 2 pm.,Asci elava— ti, 36-45 x 5-6 pm , ex uncis iteratis enati, 8-spori, api- ce magnopere incrassato, pori canalis pariete J-— cum/sine usu KOH antecedente. Ascospori ellipsoidei vel interdum al- lantoidei, 2-multi-guttulati, 6-7 x 2-2.5 pum. Paraphyses filiformes, septatae, ramosae, ascos longitudine aequantes, 1 pm latae, in gelatina contente. Holotypus: CUP-—MM 850. Apothecia gregarious to fasciculate, greenish-glaucus, sessile, clothed in white hairs, deeply cupulate, walls of receptacle deeply plicate, especially in young apothe- cla; receptacle 0.25-0.5 mm diam;.xanthochroic reaction in 2% KOH. Medullary excipulum of hyaline textura intri- Cata, 7 cells™ 2-35 jim wide. jiictal excipuliummof yhiva liner interwoven textura oblita bound in a copious, yellowish gel, cells 3-6 pm wide, outer cells giving rise to hairs. Hairs hyaline, erect, septate, coarsely granulate, 35- fagy Re. PR MWTE Asci clavate, arising from repeating cro- ziers, 8-spored, apex greatly thickened, pore channel wall J- with or without KOH pretreatment, 36-45 x 5-6 pm. Ascospores ellipsoid or,occasionally allantoid, 2- 149 multiguttulate, 6-7 x 2.0-2.5 wm. Paraphyses filiform, Bep.dleyem branched, = samen iength “as asci, 1 yam wide, pound= ity ‘gel: ool te meivete uh Oll, aN. GaebDenison, UsM. Kohn. & M.A. Sherwood, on wood of ?Castanea sativa Mill., near mine entrance at Km mark 13, road between Buenavista and Piel asco,eeLassralma,. Canary —lslands, Spain,” 18.1.1976. (CUP-—MM 850.) KNOWN MACARONESIAN DISTRIBUTION CANARY ISLANDS. La Palma. CUP-MM 850 [holotype] (TFC, isotype). SUBSTRATA: On wood (of ?Castanea sativa). Notes: The greenish-glaucus color, the deeply plicate receptacle (giving young apothecia the appearance of quae bundtmcake!). andy the xanthochroice reaction in 2% KOH makes this species especially distinctive. Neverthe- less, the ectal excipulum composed of textura oblita and the granulate hairs show Trichoscyphelloideus affi- mities.) Note’ that young “apothecia often have no asci and bear a copious gel layer above the hymenium. MYCOTAXON Volar Xt SaNoO wee pp alo 1.0 April-June, 1981 RESINOMYCENA GEN. NOV. (AGARICALES), AN ALLY OF HYDROPUS, MYCENA AND BAEOSPORA S.A. REDHEAD Btosystemattes Research Instttute, Research Branch, Agrtculture Canada, Wn. Saunders Bldg., C.E.F., Ottawa, Ontario, Canada KI1A 0C6 & R. SINGER Field Museum of Natural Htstory, Roosevelt Road at Lake Shore Drtve, Chicago, Illinois, U.S.A., 60605 ABSTRACT A new genus, Restnomycena, is proposed to accommodate Agarteus rhododendrt, the tyve, Mycena kalalochensts, and three new species, R. brunnescens, R. montana and R. acadtensis. At present the genus is known with certainty only from North America. The relationships to the genera Baeospora, Hydropus and Mycena are discussed. Agaricus rhododendrt Peck and Mycena kalalochensts A.H. Smith along with three previously undescribed taxa form a clearly defined taxon characterized by amyloid spores, pseudoamyloid tramal tissues and a turf-like pileal epicutis composed mainly of resin secreting oleocystidia. In her monograph of the northeastern Marasmius species, Gilliam (1976), excluded M. resinosus Peck (= A. rhododendrt) and suggested that a new genus might need to be proposed for it. A.H. Smith (1947) excluded both Omphalta rhododendrit (Peck) Sacc. and M. resinosus from Mycena although at the same time describing M. kalalochensis as a new Species of Mycena. These epithets were not dealt with in The Agaricales in Modern Taxonomy, (Singer 1975). In this last publication the species described here clearly key out to the Tricholomataceae trtbus Myceneae and less clearly to the choice between Hydropus and Mycena. However, the delimitations of both genera exclude the group. Thus the recognition of a distinct genus is a viable option. Alternatively emendation of Hydropus or Mycena oe from the concepts accepted in The Agaricales tn Modern Taxonomy could be proposed. The emendation of Baeospora by the inclusion of Baeospora pallida Singer (1977), a species with close but not crowded lamellae and white carpophores, is cause for considering this last genus aS a possible depository for A. rhododendri and allies. As detailed at the end of this paper, the erection of a new genus most clearly resolves the problems surrounding the placement of these species. Emendation of existing genera would make them too heterogeneous. Restnomycena Redhead & Singer, gen. nov. Habitus omphaltodeus vel marasmiotdeus, superficetbus + restnosts ex oleocystidits (frequenter tntermixtts elementis filamentosts vel dendrotdets). Lamellis adnati vel subdecurrenttbus, confertis vel subdistanttbus (haud confertissimis), ad actem oleocysttdits numerosts tnstructis. Stipe aequalts, subearttlagineo, steco subcorneo, + restnaceo. Tramate debtltter vel sat mantfeste pseudoamylotdeo (parttbus subgelatinascenttbus exceptts); tramate hymenophoralt regulart. Sports hyalints, amylotdets, levtbus, acyanophilts. Typus: Agartcus rhododendrt Peck. Basidiomes omphaloid or marasmioid, resinous, covered with Oleocystidia intermixed with dendroid hyphae, white or dully pigmented. Lamellae adnate to slightly decurrent, close to subdistant but not crowded, with resinous edges formed by numerous oleocystidia. Stipe equal, cartilaginous but drying to a horny consistency, resinous, covered by scattered or clumped oleocystidia. Tramal tissues weakly to strongly pseudoamyloid where walls not slightly gelatinized, but frequently some inamyloid hyphae intermixed. Hymenophoral (lamellar) trama regular. Spores smooth, amyloid, hyaline, acyanophilic. Type species: Agaricus rhododendrt Peck. A key to Restnomycena species Pee ELUMPEULODS balce cisco cee e's eatatéle che eee sets cients ets .- Marasmtus rhododendrt (see discussion) A. From eastern North America ..... a iale siete tele maven 6 conte 5 ib retois esse eeia oe AE Aeron western North America” ....c+ccce es a teroters ates es etatare ¢ Bienes ae ticle D B. Pileus brown; hyphae incrusted with B. Pileus white or whitish; hyphae lacking Pigmented Aincrustation ....ccccccccesovecs Braletere afst tiene aie eis e aie a c C. Lamellae close; spores small, He An. 5 Ko Ano. (455) 1M. «ole oe RA nee CEO . R. rhododendrt C. Lamellae moderately spaced to subdistant; spores large, 9.4-12.8 X 4-5um ......---- eee Ee ITE Ph acddterners u5z D. Pileocystidia mainly capitate, secondarily septate, mostly erect; on angiosperm litter ..........6.. R. kalalochensts D. Pileocystidia clavate but rarely capitate, rarely if at all secondarily septate and often collapsed to a repent condition; on coniferous litter .......... R. montana Restnomycena rhododendri (Peck) Redhead & Singer, comb. nov. Figs. 1-5. Agartcus rhododendri Peck (1875: 94) Omphaltta rhododendrt (Peck) Saccardo (1887: 335) Omphalopsts rhododendri (Peck) Murrill (1916: 311) Marasmius decurrens Peck (1872: 77) nom. illeg. non. M. decurrens Montag. (1854: 118) = Marasmtus restnosus Peck (1883: 181) nom. nov. for M. decurrens Pk. Marasmius resinosus var. niveus Peck (1903: 38) = Marasmius restnosus var. candtdtssimus Peck (1905: 40) nom. nov. tlleg. for M. resinosus var. niveus Pk. WU va i] tt PILEUS: 4-15(-19) mm wide, convex becoming plano-convex to plane, usually depressed centrally, occasionally umbilicate, rarely subumbonate, white to yellow white, opaque to vaguely striate marginally becoming obscurely corrugated-striate on some and sometimes obscurely concentrically ridged, dry to tacky or slightly viscid, micaceous when dry; edges incurved at first, uneven with age; context tough-pliant, concolourous; odor and taste not distinctive. LAMELLAE: adnate to subdecurrent or arcuate-decurrent, whitish, moderately narrow, close, often becoming forked in places or developing ladder-like anastomoses; edges crenulate, in some specimens beaded with resin. STIPE: 12-50 nm long, 0.5-1 mm wide, equal or slightly enlarged above, tough and pliant, drying to a cormeus texture, whitish, varying from glutinous to tacky or dry and glistening from resinous cells in scattered beads, fistulose, with a silky radiating white basal disc or subiculun. PILEAL EPICUTIS: a layer of polymorphic, suberect, later decumbent and tangled cystidia ranging from cylindrical narrowly clavate or capitate oleocystidia to narrow dendroid nonresinous forms with some intergradation, 40-60 X 3-9.5um, hyaline, thin-walled, often filled or partially filled with vacuolate to homogeneous oily contents and covered with similar exudates which dry as slightly yellowish and hardened masses. PILEAL TRAMA: obscurely duplex, with slightly broader hyphae above, 5-15um diam., than below, 3-10um diam.; hyphae subparallel, clamped, smooth, walls thin to slightly thickened, faintly pseudoamyloid. LAMELLAR TRAMA: similar to pileal trama, varying to slightly interwoven, more definitely pseudoamyloid. CHEILOCYSTIDIA: abundant, forming a sterile edge, similar to the cylindrical to narrowly clavate or fusoid oleopileocystidia, 55-65 X 7-7.5um. PLEUROCYSTIDIA: rare or absent, similar to the cheilocystidia. BASIDIA: 19-22 X 5-5.2yum, narrowly cylindrical to obscurely utriform, 4-spored, clamped. BASIDIOSPORES: 5.4-8.5 X 2.4-4.1(-4.5) um often being predominantly at one or the other end of the range, mostly ellipsoidal or broadly cylindrical, inequilateral in profile, prominently apiculate, smooth, hyaline, thin-walled, amyloid. STIPE HYPHAE: parallel, 3-5ym diam. in the cortex, up to 10um diam. in the medulla, mostly prominently pseudoamyloid in the medulla, walls uneven and slightly thickened. CAULOCYSTIDIA: similar to the pileocystidia, often clustered. BASAL MYCELIUM: interwoven, 2-3um diam., smooth, inamyloid, clamped, hyaline, walls refractive and slightly thickened. HABITAT, HABIT and SUBSTRATES: scattered to subcespitose on leaf litter, small twigs or bits of wood and woody fruits or husks from Rhododendron, Quercus, Fagus, Castanea, Carya and sometimes Pinus when mixed with Quereus in eastern hardwood forests. COLLECTIONS EXAMINED: (only collectors initials cited after first full citation): CANADA: Ontario: London, Sept. 27, 1896, Aug. 18, 1897, Sept. 8, 1915, J. Dearness (DAOM); and probably London area, sept. 11, 1903 and October 23, 1926, J.D. (DACM); Toronto, July 12, 1932, H.S. Jackson (DAQM 50112, ex TRIC 3381). U.S.A.: Georgia: Rabun Co., Rabun Bald, Sept. 3, 1947, Walters 153 (MICH). J2linots: River Forest, bank of Desplaines R., July 1, 1902, E.T. & S.A. Harper 542 (F). Kentucky: Harlan, Sept. 4, 1916, C.H. Kauffman (MICH). Michtgan: Lenawee Co., Cleveland L., Omsted State game area, pepe. 9, 1970, W. Patrick (M. Gilliam 939; MICH); Livingston Co., George Reserve, July 27, 1970, J. Williams (M.G. 834; MICH), Pinckney, Bide 195) ,eAstH. omith 68235 (MICH), Oakland '\Co-,, Proud L., Oct. 10, PI7O 70M. G.) 992) (MICH), Haven’ Hill, Aug. 8, 1972, A.H.S. 81375 (MICH); MiesrtLemaweCOl. me Ali ALDOL, sUULV 920, 0.912, C.HaKe, OCt. /,,1931, (A-HsSey irae 935 8A... 11699, Aug. 922, 1937,,A.Heo. /1/5, Aug. 11, 1960, Mensa enol5/3, July 11, 1970, C. Nimke (M.G. 693, 701), July 30, 1970, C. Nimke 49 (MICH), Gorman L., Aug. 20, 1972, A.H.S. 81620 (MICH), Halfmoon L., July 26, 1970, S.J. Mazzer 6186 (MICH), Manchester, mtv 4,9 1935, A.H.S. 1470 (MICH), Sharon Hollow, Aug. 4, 1960, R.L. Shaffer 2513, Aug. 19, 1970, M. Gilliam 929, Sept. 16, 1970, M.G. 952 (MICH), Silver L. area, Aug. 22, 1960, R.L.S. 2646 (MICH), Waterloo Rec. area, Sept. 3, 1968, F. Hoseney 1094, July 10, 1970, F. Hoseney Mec mol), uly 22,.1970, M.G../86, July 2,,1971, F.H. 1800 (MICH), Winnewana, July 11, 1970, M. Gilliam 648 (MICH). Minnesota: Rice Co., Wheeling Two., Nerstrand State Park, Aug. 29, 1965, M.G. Weaver 1243 (MICH). Worth Carolina: July 11, 1924, J.V. Couch 7380 (MICH), Swain Co., Flat Cr., Aug. 1, 1937, L.R. Hesler & A.J. Sharp (MICH). New York: Albany, rural cemetery, July-Aug. 1872, C.H. Peck [type of M. decurrens] (NYS); East Worchester, July, C.H.P. (NYS); Forestburgh, Sept. 1874, C.H.P. [type of A. rhododendri] (NYS); Genesee Co., Bergen Swamp, Sept. 4, 1972, H.S. Vishniac (MICH); Greenbush, Aug., C.H.P. (NYS); Ithaca, Aug. 11, 1904, C.H.K. (MICH), Aug. 1935, H.C. Beardslee Jr. 35074 (MICH); Port Jefferson, Aug. 6, C.H.P. [type of M. restnosus var. niveus] (NYS), Schuyler Co., Hector land rice area, Aug. 19, 1972, feSeVea(MICH). Ohio: Lane; July 13, 1922, H.C.B. Jr. (MICH); Portage Co., West Branch State Park, July 8, 1972, M. Gilliam 1493 (MICH) . Pennsylvania: Mt. Gretna, Sept. 5, 1926, C.H.K. (MICH) ; Media, June 29, 1940, P.M. Rea & Woodbury (MICH). Tennessee: Great Smoky Mts. Park, Cades Cove, Aug. 18, 1938, A.H.S. 10374, Indian Gap, Aug. 29, 1938, A.H.S. 10625, Aug. 7, 1942, L.R.H. 14476, Laurel falls ert eA 5, 01936,0A.H.S. 9909 (MICH). Restnomycena rhododendri was first described by Peck (1872) as Marasmius decurrens. He apparently collected a similar fungus in LSS Figs. 1-5. Restnomycena rhododendrt (DAOM, ex Dearness Oct. 23, 1926). 1, basidiospores. 2, basidia. 3, pileocystidia. 4, caulocystidia. 5, cheilocystidia. WMHs) Greenbush in 1869, #20, but lost the specimen before describing it (J. Haines, pers. comm.). Peck's validating description of A. decurrens indicated a dark taxon with a grayish or tawny pileus and a gray stipe. These colours are not apparent on the type of A. decurrens, microscopically or macroscopically, possibly indicating that more than one taxon was involved when the description was written. However, as the name is now lectotypified by Gilliam (/.c.), it applies to the pale form described in this paper. Peck (1883) not Saccardo (1887) as is often cited, later proposed the new name M. resitnosus for M. decurrens which was a later homonym of M. decurrens Mont. However, Peck (1875) had in the meantime unknowingly described it a second time as Agaricus rhododendrt which is, therefore, the earliest valid and legitimate name. His variety niveus is herein treated as being con-varietal with the type variety as it is now typified. The proposal of the new name M. restnosus var. candtdisstmus to replace M. restnosus var. niveus is superfluous under present nomenclatural rules, but it should be kept in mind that Peck was following the Rochester Code. Smith (1947) was the first to recognize that Agartecus rhododendrt and Marasmius restnosus were congeneric, although he reported the spores of A. rhododendrt to be smaller, 3-4.5 X 2.5um, than for M. restnosus. Bigelow (1970) treated the two as a conspecific but offered no evidence. In our studies spores measuring 6.5-7.2 X 3-3.2um were found on the type of A. rhododendri which in other respects is typical for M. restnosus. Thus the two are treated as conspecific. Hesler's (1959) report of inamyloid spores for the type of M. restnosus is evidently in error as we concur with Gilliam (1976) that amyloid spores are present. Restnomycena rhododendrt appears to be restricted to the eastern deciduous forest where it occurs on litter of the dominating hardwood trees. It is the largest of the Restnomycena species and the one most commonly collected. Restnomycena kalalochensts (Smith) Redhead & Singer, comb. nov. Figs. 6-10. = Mycena kalalochensts A.H. Smith (1947: 99) "Pileus 3-8 mm. broad, convex, remaining broadly convex, margin incurved at first, spreading in age, chalk White and appearing pruinose under a lens at first, glabrous and uneven in age but remaining chalky, slightly sulcate at maturity, not hygrophanous; flesh membranous and pliant (but not reviving), odor not distinctive, taste not recorded; lamellae adnate, broad, distant, 10-12 reach the stipe, two tiers of lamellulae, white over all, edges pruinose; stipe 3-7 mm. -long less than 0.5 mm. thick, equal or the base flanged slightly, strigose, the remainder pruinose like the pileus, chalky white over all." Smith (1947: 99-100). PILEAL EPICUTIS: an erect to suberect tangled turf of polymorphic cystidia, 25-50 X 3-8um, varying from cylindrical to narrowly clavate, usually capitate to subcapitate, often secondarily septate, scantily resinous oleocystidia to variously branched to Se narrow dendroid and antler-like forms, which lack resinous exudates or contents, walls thin, hyaline, smooth, clamped basally. PILEAL TRAMA: somewhat duplex with slightly broader hyphae above, 5-15um diam., than below, 5-10u diam.; hyphae faintly pseudoamyloid initially but becoming more intense after several days in permanent Hoyer's-Melzer's reagent, clamped, smooth, thin-walled, subparallel. LAMELLAR TRAMA: hyphae similar to the pileus trama hyphae. CHEILOCYSTIDIA: abundant, forming a sterile edge, less variable than the pileocystidia, mostly narrowly cylindrical to slightly clavate, occasionally forked or branched, occasionally with an apical finger-like elongation, 31-48 X 5-6.5um, scantily resinous. BASIDIA: 24-25 X 7.5-8.0um, clavate to obscurely utriform, 4-spored, clamped. BASIDIOSPORES: 7.8-10.8 X 3.8-4.8um, ellipsoid to obscurely fusoid or occasionally obovoid, inequilaterally flattened in profile, smooth, thin-walled, amyloid, hyaline, with a prominent apiculus. STIPE HYPHAE: parallel, 5-15um diam., smooth, pseudoamyloid, hyaline, with the broadest hyphae in the medulla. CAULOCYSTIDIA: similar to the pileocystidia but only scantily resinous like the cheilocystidia. BASAT, MYCELIUM: not studied. HABITAT, HABIT and SUBSTRATES: Scattered on litter of Rubus, Alnus rubra Bong. and grasses in the coastal forest zone of western North America. COLLECTIONS EXAMINED: CANADA: Brtttsh Columbia: North Vancouver, Capilano Canyon, Oct. 8, 1973, S.A.R. #AS 9 (DAOM 166524), Vancouver, Warvemts Cpe uuner2, LOAN S.Avk. 3425 (UBC) .0 UsoeAas) SCaLtrornia: Humbolt Co., Big Lagoon Park, Dec. 18 & 19, 1956, A.H.S. 56787, 50033,5 50823, Prairie Cr. Park, Dec.99, 1956, A.H-S. 56520 (MICH). Oregon: Seaside, Sept. 21, 1944, W. Gruber & A.H.S. 19020 (MICH). Washtngton: Jefferson Co., Kalaloch, April 30, 1939, A.H.S. 13035 [type] (MICH). A.H. Smith (1947) reported slightly larger spores, 8-11 X 5-6um, an inamyloid lamellar trama and a lack of incrustations on the pileocystidia. In our studies resinous exudates and dextrinoid tramal tissues were found on the type although the iodine reaction was not as prominent as that for the stipe. In addition, the resinous exudates tend to become clarified and dissolve in Melzer's reagent more so than in KOH 3% aqueous sol. Resinomycena kalalochensts differs from R. rhododendrt by distant lamellae, smaller basidiomes, larger spores, and more scant resinous exudates, in addition to the differences in geographic range and to a lesser extent substrates. Restnomycena montana Redhead & Singer, sp. nov. Figs. 11-14,26. Pileus 3.5-5.5 mm latus, convexus vel depressus, candidus. Lamellae arcuatae, candidae. Sttpes 9-23 mm long., 0.4-0.6 mm erassus, candidus. Sporae 8-9.3 X 2.8-4.5um, amylotdeae, elltpsotdeae. Ptleocysttdia 45-67 X 2.5-7\m, restnosa, clavata vel dendrotdea ut nonrestnosa. Chetlocystidta 42-52 X 6-6.2um, restnosa, elavata vel strangulata. Tramate pseudoamylotdeo. a | | ; 10 Figs. 6-10. R. kalalochensts (DAOM 166524). 6, caulocystidia. c 7, basidiospores. 8, cheilocystidia. 9, basidia. 10, pileocystidia. 160 Figs. 11-14. FR. montana (DAOM 178214). 11, basidia. 12, pileocystidia. 13, cheilocystidia. 14, basidiospores. Lot Holotypus: DAOM 178215. Donald Station, Rocky Mt. trench, B.C., Canada, Sept. 21, 1980, S.A. Redhead 3977. PILEUS: 3.5-5.5 mm wide, convex and slightly depressed centrally, vaguely striate to chalky white, micaceous, obscurely corrugated-striate, developing greyish casts when dry; context white, pliant; odor not distinctive. LAMELLAE: arcuate decurrent, moderately spaced, white, with micaceous edges; lamellulae in 2 tiers. STIPE: 9-23 mm long, 0.4-0.6 mm wide, equal, cargilaginous or tougher, white, finely powdered overall. PILEAL EPICUTIS: a suberect to tangled turf of polymorphic cystidia 45-67 X 2.5-7um, varying from a majority of elongated clavate oleocystidia to filiform or dendroid nonresinous elements. PILEAL TRAMA: obscurely duplex, with slightly broader hyphae above, 3-llum diam., than below 3-5yum diam.; hyphae subparallel, faintly pseudoamyloid at first, smooth, clamped, thin to slightly thick-walled, hyaline. LAMELLAR TRAMA: hyphae similar to the lower pileal trama hyphae. CHEILOCYSTIDIA: abundant, forming a sterile edge, 42-52 X 6-6.2um, narrowly clavate to cylindrical and slightly strangulate, occaSionally with a short apical elongation, with scant to abundant resin exudates. BASIDIA: 20-21 X 6.8-7.2um, clavate subcapitate, 4-spored, clammed. BASIDIOSPORES: 8-9.3 X 2.8-4.5um, cylindrical to narrowly ellipsoidal, slightly inequilateral in profile, amyloid, smooth, hyaline, with a prominent apiculus. STIPE HYPHAE: parallel, pseudoamyloid especially on the slightly broader medulla hyphae, 4-13um diam., smooth, thin to slightly thick-walled, clamped. CAULOCYSTIDIA: scattered to abundant and often clustered, similar to the pileocystidia. BASAL MYCELIUM: not studied. HABITAT, HABIT and SUBSTRATES: scattered to gregarious on coniferous needles, cone debris and small twig fragments in western Subalpine, Coastal and Columbian coniferous forests of North America. COLLECTIONS EXAMINED: CANADA: Brittsh Columbta: North Vancouver, Mt. Seymour, 3500' alt., Sept. 17, 1973, S.A.R. #AJ 3 (DAOM 178218); Sept. 22, 1973, S.A.R. #AI 24 (DAOM 178214); Columbia R. valley in Rocky Mt. trench at Donald Stn. on Hwy. 1 crossing of river, Sept. 21, 1980; S.A.R. 3977 [type] (DAOM 178215). U.S.A.: Washington: Lower Elwha, July 3, 1939, A.H.S. 14732 (MICH). Restnomycena montana has longer spores than Rk. rhododendrt, occurs on coniferous debris and usually develops a greyish cast on drying. Geographically the two are widely separated. Resinomycena kalalochensis differs by its more capitate and shorter pileocystidia Which are often secondarily septate and scarcely resinous. Also the latter has smaller basidiomes which have not been observed to develop a grey cast and it occurs on angiosperm debris. Restnomycena acadtensts Redhead & Singer sp. nov. Figs. 20-25. Pileus 2-10 mm latus, convexus denum depressus, candidus. Lamellae arcuate, ecandidae. Stipes 10-40 (-70) mm long., 0.2-0.8 mm crassus, candidus. Sporae 9.4-12.8 X 4-5um, anyloideae, elltpsotdeae. Ptleocystidta 30-58 X 3-9um, restnosa, clavata vel nonrestnosa ut 162 dendrotdea. Chetlocystidta 38-40 X 6-8um, aeque ptleocysttdia. Tramate pseudoamylotdeo. Holotypus: DAOM 166073. Kouchibouguac Natl. Park, N.B., Canada Sept. 24, 1977, J.E. & S.A. Redhead 2558. PILEUS: 2-10 mm wide, convex becoming depressed to cyathiform, translucent-striate but readily becoming opaque on partial drying, white to whitish, occasionally tinted fawn on the disc, rugose with age, moist to dry; edges incurved initially; context white, pliant; odor and taste not distinctive. LAMELLAE: arcuate-decurrent, becoming more distinctly decurrent, white, subdistant to moderately spaced; edges crenulate and often beaded; lamellulae in 2 tiers. STIPE: 10-40 (-70) mm long, 0.2-0.8 mm wide, equal or tapering up, white or with ochreous to rosy buff tints basally, glutinous when wet, powdered when dry, often attached to the substrate by a small radiating silky white mycelial pad. PILEAL EPICUTIS: a sSuberect turf and later repent tangle of polymorphic cystidia 30-58 X 3-9um, varying from short to long clavate oleocystidial forms to narrow filiform to dendroid nonoleocystidial forms with some intergradation, thin-walled, smooth, hyaline, with copious to moderate resinous exudates. PILEAL TRAMA: obscurely duplex with the broadest forms in the upper half, hyphae subparallel, 5-llum, smooth, hyaline, faintly pseudoamyloid at first, clamped. LAMELLAR TRAMA: hyphae mostly parallel, 3-5um diam., pseudoamyloid. CHEILOCYSTIDIA: abundant forming a sterile edge, similar to the pileocystidia but rarely of the branched forms, mostly clavate, 38-40 X 6-8um. BASIDIA: 20-32 X 7-8um, (2-)4-spored, clavate, clamped. BASIDIOSPORES: 9.4-12.8 X 4-5yum, ellipsoidal to cylindrical or obscurely reniform to pip-shaped, inequilaterally flattened in profile, with a prominent apiculus hyaline, white in mass, amyloid. STIPE HYPHAE: 3-10um diam., with narrower hyphae in the cortex than in the medulla, strongly pseudoamyloid, smooth, parallel, hyaline, clamped. CAULOCYSTIDIA: usually clustered, similar to the pileocystidia but less often with long pedicels and more often secondarily septate. BASAL MYCELIUM: slightly interwoven, agglutinated, 3-10um diam., pseudoamyloid. HABITAT, HABIT and SUBSTRATES: scattered on small broken pieces of woody angiosperm debris in predominantly Acadian coniferous forests of larch or spruce, fir and pine. COLLECTIONS EXAMINED: CANADA: Wew Brunswick: Kouchibouguac Natl. Park, Aug. 11, 1977, R.L. Milikin (DAOM 169619), Sept. 24, 1977, J.E. & S.A.R. 2558 [type] (DAQM 166073), Oct. 9, 1978, S.A.R. 2753 (DAOM) . Restnomycena acadtensts can be distinguished from R. rhododendri by the larger spores, more distant lamellae and the more flaccid flesh. Restnomycena kalalochensts differs by its small stature, its more erect turf of capitate and scarcely resinous, secondarily septate pileocystidia and its geographic range. Restnomycena montana differs by its smaller usually narrower spores, smaller basidia, a coniferous substrate and its geographic range. The glutinous to viscid stipes are a result of copious resinous 163 exudates. These dry down to hardened masses in less humid situations creating a powdered appearance to the stipes. In herbarium specimens these masses become yellow with age. Restnomycena brunnescens Redhead & Singer, sp. nov. Figs. 15-19. Pitleus 6 mm latus, planoconvexus ut depressus, rugulosus, brunneus. Lamellae adnatae vel subdecurrentes, bubalineae vel ochraceae. Sttpe 20 mm tlong., 0.5 mm crassus, concolorus. Sporae 6.5-8 X 3.3-4um, amylotdeae, elltpsotdeae. Ptleocystidia 55-90 xX 3-1lum, restnosa, clavata vel strangulata, vel nonrestnosa ut dendrotdea. Chetlocystidia 36-50 X 10-14um, resinosa, clavata vel capttata, rarts nonrestnosa ut dendrotdea. Tramate pseudoamylotdeo. Hototypus: DAOM 165884. Kouchibouguac Natl. Park, N.B., Canada, July 13, 1977, J.E. & S.A. Redhead 2336. PILEUS: 6 mm wide, plano-convex and depressed centrally, rugose, tacky, scarcely translucent on edges, greyish sepia centrally, fulvous with ochreous to buff margins; context membranous, thin, concolorous above; odor not distinctive. LAMELIAEF: adnate to subdecurrent, moderately spaced, buff to ochreous with whitish crenulate edges, sometimes forked; one tier of lamellulae. STIPE: 20 X 0.5 mm, concolorous with the pileus with the darkest pigments below, with a striated powdered appearance from dried resin, micaceous, attached to the substrate by a radiating white mycelial pad. PILEAL EPICUTIS: a layer of tangled polymorphic cystidia 55-90 X 3-llum, varying from clavate to fusoid often strangulate oleocystidia exuding copious resinous exudates to filiform often branched dendroid nonresinous forms. PILEUS TRAMA: duplex, with hyphae 10-15yum diam. subparallel, thin-walled, pseudoamyloid, yellowish brown and prominently incrusted with brownish resin-like materials in flat patches above, 5-10um diam., hyaline, walls thin or pronounced below. LAMELLAR TRAMA: hyphae similar to those of the lower pileus trama. CHEILOCYSTIDIA: abundant, forming a sterile edge which on drying is a continuous hardened resinous line, 36-50 X 10-14um, clavate and prominently capitate, hyaline, thin-walled. BASIDIA: 27-29 X 5-6um, narrowly clavate (2-)4-spored, clamped. BASIDIOSPORES: 6.5-8 X 3.3-4yum, ellipsoid to obscurely obovoid or fusoid, hyaline, smooth, thin-walled, amyloid, with a prominent apiculus. STIPE HYPHAE: parallel, 3-5ym diam. in the cortex with pigments similar to the pileus tramal hyphae, 10-15um diam. in the medulla, pseudoamyloid, clamped. CAULOCYSTIDIA: clustered, similar to the cheilocystidia but less swollen apically, occasionally forked, intermixed with a few dendroid filiform elements. BASAL MYCELIUM: not studied. HABITAT, HABIT and SUBSTRATE: solitary on fallen leaf of Rhododendron canadense (L.) Torr. in a dense Acadian coniferous forest. COLLECTION EXAMINED: CANADA: WNew Brunswick: Kouchibouguac Natl. Park, July 13, 1977, J.E. & S.A.R. 2336 [type] (DAOM 165884). Resinomycena brunnescens is readily distinguished from all other species in the genus by its darkly pigmented pileus and stipe. The 165 Figs. 15-19. R. brunnescens (DAOM 165884). 15, basidia. 16, cheilocystidia. 17, caulocystidia. 18, basidiospores. 19, pileocystidia. 167 a - oq N ie) oO’ ~ Yo) ma e a a f i sz Y g NO 8 Bh hed f A vaae oa = oO = ea ~ AN 1 oI ONN 4 Ve) AN wo foo) coe ef PELE ees 3B qa o~ a6 g ~~ as eBags 2 = © O se) < tS el ia ON Aq ora Sls ne 20, basidiospores. 23, cheilocystid Scales = 15um. Figs. 20-25. 168 incrusted hyphae responsible for this darkened appearance are very distinctive microscopically compared to the nonincrusted hyphae in other Restnomycena species. In addition, R. brunnescens is characterized by very prominently capitate cheilocystidia which are inflated more than in any of the other species. As noted above, Peck's original description of Marasmtus decurrens is of a brown fungus. The existence of a brown species of Restnomycena in eastern North America suggests that Peck may have collected both R. brunnescens and R. rhododendrt. However, this cannot be confirmed now. GENERAL DISCUSSION The three most probable generic depositories for Resinomycena if the taxon is not recognized as a distinct genus are Hydropus, Mycena and Baeospora, there being a foregone conclusion that the species are excluded from the genus Marasmtus because of amyloid spores. The genus Baeospora as it was defined in The Agartcales tn Modern Taxonomy (Singer, 1975) represents a small cohesive genus characterized by a number of characters, but in particular, small amyloid spores, inamyloid tramal tissues and crowded lamellae. Restnomycena rhododendrt is the species most similar to a Baeospora in that it has close lamellae and small amyloid spores, however, the tramal tissues of the pileus and stipe are pseudoamyloid unlike any Baeospora species. Also the massed large oleocystidia mixed with dendroid elements on the pileus form a structure unlike any epicuticular formation found in Baeospora, including B. pallida. When all species of Resitnomycena are taken into account, species such as R#. acadtensts and R. kalalochensits, exhibit characters not found in Baeospora such as distant, slightly decurrent lamellae and large spores. As a result of these combinations of characters Restnomycena cannot readily be incorporated into Baeospora. The genus Hydropus as defined by Singer (1975) contains species having numerous pileo-, cheilo- and caulocystidia and in some species pseudocystidia with oily contents. The spores can be amyloid and the tramal tissues can be weakly pseudoamyloid. Again the abundant Oleocystidia on the pilei, stipes and lamellae of Restnomycena species, not found in Aydropus, in combination with intermixed dendroid elements form a tissue not found in Aydropus and clearly separate the genera. The genus Mycena is treated as delimited earlier (Singer 1975). It includes species with amyloid spores, pseudoamyloid tramal tissues and oleocystidia as does Hydropus. The majority of Mycena species have an epicutis consisting of a repent layer of diverticulate narrow hyphae. Deviations occur in some sections of the genus where smooth repent filamentous hyphae replace the diverticulate elements. Those species which had smooth epicuticular hyphae bearing obtuse cystidia were removed to the genus Hydropus. The most aberrant type of epicutis for a species admitted to Mycena by Singer (1975) is found in Mycena rorida (Fr.) Kuhner and its allies. ‘These species have a hymeniform layer of sphaeropedunculate cells and, apparently missed by previous authors, in M. roritda, intermixed oleocystidia. Previously (Singer 1975), it was indicated that these species were related to Hydropus and might need to be removed from Mycena. However, the strongly pseudoamyloid tramal tissues and a gelatinized stipe cortex weighed in favor of retaining the species in Mycena. Here again, the well developed layer of usually clavate oleocystidia and narrow dendroid elements found in Restnomycena is not typical of any section of Mycena and represents a large divergence from a typical Mycena type of epicutis. Furthermore, the typical Mycenae have characteristically broad (broader than Resinomycena) hypodermal and tramal hyphae; the hyphal cells often being short and voluminous. In a less restrictive delimitation of Mycena such as by Smith (1947) or Kuhner (1938), Restnomycena would be included. From the above discussion it can be seen that Baeospora, Hydropus and Mycena are open to emendation to encompass Resinomycena species but only by introducing a species group having an unusual combination of characters peripheral to the core of the genera involved. Instead the proposal to recognize a distinct genus seems more justified. Restnomycena species are obviously closely related to each other. They are readily distinguished from Baeospora, Hydropus and Mycena by the abundance of resinous oleocystidia intermixed with dendrohyphidia. The segregation of Restnomycena species maintains the integrity of the older genera, leaving them less heterogeneous and, therefore, more recognizable. There is no doubt that Resinomycena is closely related to Baeospora, Mycena and Hydropus. Obviously one solution would be to merge all three genera and add Resinomycena. Again there seems to be more merit in recognizing each of these genera including Restnomycena. Finally, Marasmius rhododendrt Singer (1947, 1936), a species described from Spain in the subalpine zone of the Pyrenees on Rhododendron ferrugineum L. is probably congeneric. ‘The type (F) now consists of stipes only. Spores obtained from the type were amyloid although weakly in some and up to 12.5um long. The stipe hyphae are pseudoamyloid and in an outer layer more or less gelatinized (the stipe was described from fresh material as somewhat shiny and viscidulous). They bear cystidia with dried incrustations similar to that found on Restnomycena species. In the original description (1936) yellowish incrustations were mentioned on the lamellar edges of vesciculose to ampulliform cheilocystidia ("echinulatis" possibly being an error in the 1947 description unless referring to dendroid elements). The exact nature of these cystidia and the pileipellis could not be restudied in the damaged type collection and the identity of the species remains in doubt. The weakly developed collar is not found on the presently recognized Restnomycena species. This species should be restudied from fresh materials. ACKNOWLEDGEMENTS We thank Drs. J. Haines (NYS), R.J. Bandoni (UBC), R.L. Shaffer, R. Fogel and A.H. Smith (MICH) for the use of herbarium materials and facilities. Drs. J. Ginns and J. Ammirati offered appreciated reviews and Mr. K.W. Spicer provided technical assistance. Parks Canada supplied collecting permits or facilities in New Brunswick and British Columbia and the Agriculture Canada research station, Kamloops, 170 provided transportation in 1980. Mrs. M. Meredith typed the final manuscript. LITERATURE CITED Bigelow, H.E. 1970. Omphaltna in North America. Mycologia G2 spelen 3 2 Gilliam, M.S. 1976. ‘The genus Marasmtus in the Northeastern United States and adjacent Canada. Mycotaxon 4: 1-144. Hesler, L.R. 1959. Southeastern Agaricales, IV. J. Tenn. Acad. Sci. 34: 167-171. Kuhner, R. 1938. Le genre Mycena. Encyl. Mycol. 10: 1-710. Montagne, J.F.C. 1854. Cryptogamia Guyanensis seu plantarum cellularium in Guyana gallica annis 1835-49 acl. Leprieur collectarum enumeratio universalis. Ann. Sci. nat. bot., sér. 4, 1: 91-144. Murrill, W.A. 1916. Omphalopsts. N. Amer. Flora 9(5): 310-318. Peck, C.H. 1872. Report of the botanist. Ann. Rep. N.Y. State Mus. Nat. Hist. 24: 41-108. . 1875. Report of the botanist. Ann. Rep. N.Y. State Mus. NatwHist. ruc] s/o iG. ~ L883.) -Orders]16.).Fungive Dp. 1/4235 eines eDay ame plants of Buffalo and its vicinity. Bull. Buffalo Soc. Nat. SCT esa (4): 65-279. . 1903. Report of the state botanist. N.Y. State Mus. Bullvo1ce Lolo. . 1905. Report of the state botanist. N.Y. State Mus. Bull. ~94:°19=50: Saccardo, P.A. 1887. Sylloge Fungorum. V. 1-1144. Patavia. Singer, R. 1936. Notes sur quelques Basidiomycétes. Rev. Mycol. (N.S) RC 2) seb =84 . 1947. Champignons de la Catalogne. Espéces observées en 1934. Collectanea Bot. 1(3): 199-246. . 1975. The Agaricales in Modern Taxonomy. 3rd. ed. J. Cramer, Vaduz. 1977. Interesting and new species of basidiomycetes from Ecuador II. Nova Hedw. 29: 1-98. Smith, A.H. 1947. North American species of Mycena. Univ. Mich. SCI eSer. we liseli MYCOTAXON VO Lee el ee NOSSEL PSG Dp Dic guile 17 4 Api JUneeLo6 SINOTERMITOMYCES, A NEW GENUS OF AMANITACEAE FROM YUNNAN, CHINA ZANG MU Kunming Institute of Botany, Academia Sinica, Kunming, Yunnan, China SUMMARY Detailed analysis has led the author to propose the termito- philous Sinotermitomyces as a new genus, including two new species, S. cavus Zang and S. carnosus Zang. Both were col- lected in Yunnan, China. Type specimens of the two new spe- cies are deposited in the Herbarium of the Kunming Institute of Botany, Academia Sinica (HKAS). The new genus Sinotermitomyces consists of two new species growing on termite combs. These edible fungi are distributed in subtropical and tropical regions of southwestern Yunnan. In Yunnan, profession- al collectors gather both Termitomyces and Sinotermitomyces in the rainy season from May to September, and dry preservation with salt and vegetable oils are used for building up a year-round supply (Cheo, 1942, 1948; Batra, 1979). The genus Termitomyces in China is commonly found southward of the Yangtze. The distribution of Sinotermitomyces is in forests of the southwestern part of Yunnan. Specimens were collected by the author under Pinus khasya Royle ex Cord., Pinus yunnanensis Fr., ande@astanopsisshystrixs (Hk. f.°6 Thoms.) 1A. DCs SINOTERMITOMYCES Zang, gen. nov. Pileus conicus vel campanulatus, subconvexus vel papillatus, siccus, glaber vel scabridus. Lamellae subliberae, liberae vel adnatae, albidae. Stipes centralis, cavus, aequalis vel sub-fusoideus, carnosus, coriaceus vel fibrillosus. Annulus superus distinctus. Basidiosporae hyalinae,; ellipticae vel obovatae, inamyloidea. Pleurocystidia ventricosa vel cy- lindrica, verrucosa vel tuberculosa. Cheilocystidia oblonga vel cylindri- Ca, laeviuscula vel verrucosa.‘ Fibulis nullis. Termiticola. Typus generis: Sinotermitomyces cavus Zang. Pileus conico-campanulate, subconvex and often with a prominent papillate apex, dry, never viscid, glabrous or scabrous. Gills free to almost free or adnate. Stipe central, often very long, variable in. thickness, hollow, equal or. subfusiform, fleshy or leathery or fibrillose. Annulus superus present, veil a pellicular veil that co- vers the gills in young specimens. Basidiospores from globose to ellipsoid, smooth, hyaline, inamyloid. Pleurocystidia present, cylin- dric or some fusoid-ventricose, covered with wart-—-like outgrowths. Cheilocystidia oblong to cylindric, smooth to warted. Clamp connec- tions absent. The primordia develop in the holes of Nypogeous termi- ez taria. This new genus can be distinguished from the genus Termitomyces Heim by the following characters: the small, subconvex pileus with prominent papillate apex; presence of veil remains on stipe and pileus; stipe leathery or fibrillose and hollow; pleurocystidia with sturdy, short processes (Heim, 1942; Singer, 1949). 1. Sinotermitomyces CAVUS Zang, sp. nov. (FIGS. 1-7) Pileus 1-2.5 cm., conicus vel subconvexus, siccus, glaber, primo flavi- dus vel eburneus, demum fulvo-umbrinus vel rufo-—brunneus. Lamellae sub- liberae vel adnatae, albidae vel eburneae, Stipes 20-30 cm. longus, 5- 12 mm. crassus, aequalis, cavus, tubulosus, coriaceus vel fibrillosus, apice cicatricatus, sursam scabroso-furfuraceous, basim versus discoide- us. Annulus superus distinctus.» Basidiosporae 2.4-5 x 3.5-9 pm, hyali- nae, globosae, ovoideae, ellipsoideae, laeves, inamyloideae. Basidia clavata, 15-25 x 10-14 pm, 4-sporigera. Pleurocystidia 12-20 x 25-45 pm, cylindrica vel clavata, verrucosa vel tuberculosa. Cheilocystidia 10-15 x 20-30 pm, oblonga vel cylindrica, tuberculosa. Fibulis nullis. Hab. In sylvis praecipue Pinetis yunnanensi Fr. et Pinetis khasyae Royle ex Cord. nido Termitidarum (Odontotermes) erumpente. Yunnan: Tengchung County, Tuan-Tian village, La-ba-qing. On nest of termites, in swampy grassland, under the pine woods above 2100 m. alt., 8. Vill. 19802° Li’ Xing-jiange 11. (Typus,; HKAS +6533);) tengehung County, Pu-Chuani-village,e17. Vil. 197908 Maw Xi—xiongiele com HKAcm40.i2. 4613); Shweli County, Deng Ga, in pine woods, 13. VIII. 1980. Li Xing- jiang 22. (HKAS 6568); Mangshi. 10. VIII. 1980. Zang Mu 06545 (HKAS 6545). Pileus 1-2.5 cm broad, conical or subconvex, dry, glabrous, surface ivory white or pale yellow becoming yellowish-umber or reddish- brown towards the center, glabrous. Gills almost free or adnate, white to ivory white. Stipe 20-30 x 0.5-1.2 cm, equal, hollow, tube- like, leathery to fibrillose, upwards often covered with bran-like scales or scabrous and with a discoid base. Veil persistent as a thick, membranous leathery annulus, which is striate above. Basidio- spores 2.4-5 x 3.5-9 pm, hyaline, globose, ovoid to ellipsoid, smooth, inamyloid. Basidia 15-25 x 10-14 wm, clavate, 4-spored. Pleurocystidia 12-20 x 25-45 pm, cylindric to clavate, covered with irregular wart-like outgrowths. Cheilocystidia 10-15 x 20-30 pum, oblong to cylindric, roughened. Clamp connections absent. 2. Sinotermitomyces CARNOSUS Zang, sp. nov. (FIGS. 8-10) Pileus 4-6.5 cm., conicus vel campanulatus, demum plani-convexus, sic- cus, scabridus vel subtomentosus, primo albidus vel pallidus, demum cer- vinus vel brunneus. Lamellae subliberae vel adnatae, albidae. Stipes 15-20 cm. longus, 1-2.5 cm. crassus, subfusoideus, carnosus, cavus, con- colorous, pseudorhizophorus. Annulus membranaceus, superus instructus. Basidiosporae 3-6 x 4.9-8 ym, hyalinae, globosae, ovoideae, ellipsoid- eae, laeves, inamyloideae. Basidia 9.8-12 x 14-18 wpm, clavata, 4-spori- gera. Pleurocystidia 10-18 x 24-48 pm, lageniformia vel fusiformia, exas- perata vel verrucosa. Cheilocystidia 12-18 x 18-34 pm, rotunda, ovata, oblonga, laeviscula vel exasperata. Fibulatae adsunt. Hab. In sylvis praecipue Castanopsitis hystricis (Hk. f. & Thoms.) A. DC. alt 1600-2100 m. nido Termitidarum erumpente. Yunnan: Tsangyuan Autonomous County of Wa Nationality. On termites' nest. 30. VIII. 1980. Zang Mu 06752 (Typus, HKAS 6752). \ | : tl FIGS. 1-10, Sinotermitomyces spp. (Jn. Scavus Zanes (HKAS 10533).50 91, te Carpophores+ 2, .cheilocystidia;.3, pleurocystidia; 5, basidia and basidiospores; 6, the discoid base of stipe. 8-10, 5S. (HKAS 6752). 8, carpophores; 9,, cheilocystidia, spores; 10, pleurocystidia. carnosus Zang basidia, and basidio- 173 174 Pileus 4-6.5 cm broad, conical, campanulate to plano-convex, dry, scabrous or subtomentose, surface white or pale white to waxy yel- low or brown. Gills almost free or adnate, white. Stipes 15-20 x 1-2.5 cm, subfusiform, hollow, more fleshy, uniform in colour, pseu- dorhiza present. Annulus superior, fragments adhering to stipe. Basi- diospores 3-6 x 4.9-8 ym, hyaline, globose, ovoid to ellipsoid, smooth, inamyloid. Basidia 9.8-12 x 14-18 wm, clavate, 4-spored. Pleurocystidia 10-18 x 24-48 ym, flask-shaped or fusiform, rough- ened with wart-like outgrowths. Cheilocystidia 12-18 x 18-34 ym, almost round, broadly oblong, almost smooth to roughened. Clamp connections absent. LITERATURE CITED BATRA, L. R., (ed.). 1979. Insect-Fungus Symbiosis, pp. 142-145, 148- 153% Allanheld; Osmun & Cos, Montclair. CHEOQ VC. eG. 1942. A study of Collybia albuminosa (Berk.) Petch, the termite-growing fungus in its connection with Aegerita duthiei Berk. Science Record 1: 243-248. ° 1948. Notes on fungus-growing termites in Yunnan, China. Lloydia 11: 139-147. HEIM, R. 1942. Les champignons des termitiéres. Nouveaux aspects d'un probléme de biologie et de systématique générales. Rev. Sci. 80: 69-86. SINGER, R. 1949. The Agaricales in modern taxonomy. Lilloa 22: 396- 398. , MYCOTAXON ViOuLeweE tel Le aeN Of SD Dimer iio = OO April-June 1981 TYPE eo LUDT ES oLNe THE: POLYPORACHAR 13. SPECIES SDESCRIBED IBY Js ho LEVELELE. by LEIF RYVARDEN Botanical Laboratory University of Oslo P.O. BOX L045. blindern Oslo 3, Norway >. Us Meehan Of the 145 polypores described by J. H. Léveillé 12 are accepted, 83 are synonyms, 6 names are in- valid, 95 are an uncertain status, smost of them Ga-— moderma sp.-, while 39 types were not. found. The names of many of the missing types are accepted as synonyms based on notes by previous mycologists. abercombination Tyromyces dissectus is proposed. epemracvver wpec eseand sl riachaptumeperrottetiiocare described in detail. The French mycologist J. H. Léveillé was very industri- ous and described many fungi of which 145 are polypores. His basis was the collections in the Paris herbarium be- Sides which he made a journey to Leiden and was allowed to take samples from the rich East-—Asian collections of Jung- Dunn vanGrZovVlancer. Most, Ole hasecoLlectionsrvare ptoday jin the Paris Herbarium (PC), but many of the polypore types are also in the Leiden herbarium (L.). Isotypes are at Kew (K), Stockholm (S), the Farlow herbarium (FH) and the Na- tional Frunecus Collections (BRD). Léveillé made very scanty labels, but often he fastened his specimens to the sheet with a red silky band attached both to the specimen and the sheet with a red lacquer. This made it possible to study both sides of the specimen with- out, loosening it’ from the sheet. this red lacquershas in many cases made 1t possible? to trace some of his types when label and specimen have been separated, which was the case in many collections. Never- theless, many of his types have been lost. However, both Lloyd. (1912) and Bresadola (1916 & 1920) studied many of his specimens before they disappeared and it is possible to settle their identity. Léveillé described polypores in a number of genera, and they are treated in the genus in which Léveillé originally placed them. Within each genus yO the species are placed alphabetically according to their specific epithets. After the name there is a reference to where it was published and an indication of the herbarium in which the lectotype and eventual isotypes can be found. The label is then cited in inverted commas and, if the text indicates no type locality, this 18s added in brackets. If theitype was found ‘to be 4 synonym this, is (indicated by = followed by a correct citation of the name in question. When the species has been accepted, it is cited in its proper genus with a reference to a recent description; or described in detail if no modern description seems to exist. DAEDALEA Fr. D. aulaxina 1844:197. The type has not been found (Java). = Lenzites vespacea (Pers.) Ryv. teste Bresadola (1916:230). Dem havica lous 31 OG IeCP Gr "Java". = Lenzites acuta Berk. D. fuliginosa 1844:199. The type has not been found (Mauritius). Den lurida, L644 OSS (PO® Vsotype dans). Wav aie=s lerign tesa cuta sober. Deamecrozonall oto dom ule "Zollinger no 2060" (Java) = Lenzites acuta Berk. D. plumbea 1846:142. The type has not been found (Noveboracum = New York, WShae DY pruinosa 18845198 (PC) isotype in s)- MING pacinico «= Lenzitesmacuta perk. D. splendens 1844:197. The type has not been found (Sumatra). D. Violacea 18462142. The type is apparently lost (Cuba). FAVOLUS Fr. Pee lOGa POsUsmlO ir OU ime CrCl. "Manille" (The Phillipines). = P. phillipinensis Berk. 2p ah RAVES eee Aol (COG jee "Bresil, Rio de Janeiro, Mars 1836" = F. brasilensis CRs) br F. granulosus 1863:286 (PC, isotype in K). "Chachi alt. 2600 m, Sept...1860 Lindig 2919" -(Colombia). = Panel lus pusillus - (bev. ) Duras.) & =Mlliler. F. guadeloupensis 1846:144 (PC). "Guadeloupe, M. 1‘'Herminier". The type does not represent a true polypore and has probably been fleshy and gelatinous when fresh. Today it is resinous and horny. Pileus glabrous and pustulate, dark brown, pores shallow and angular, 1-2 mm wide, dir- ty to dark brown. Context dark brown and very dense. Hy- phal system monomitic with clamps, variable in diameter, in parts inflated, 2-8 um wide, but difficult to sepa- rate. Spores and basidia not seen. The species is un- known to me. Lh me junghuhnii 1844:202 (L, isotypes in PC & 8S). DAGMUrUNCOStinmeinsulagbanvan’ ‘(rhe Phillipines) «= P. hillipinensis Berk. Meemultiplexelouu.203 (PCs) isotype in Kix S)-. "Java". = F. spatulatus (Jungh.) Lév. Pemepeltatus 18447203) (L, isotype in PC). WZ0lvinger=no. 707" (Java). =F. brastlensis (Fr.) Fr. Heetcener) Vo44; 202 (Lb). "Sumatra". = F. spatulatus (Jungh.) Lév. F. tenuissimus 1 wc Ue.s Type not found (Mauritius). GLOEOPORUS Mont. G. leptopilus 1844:194. Type has not been found (Surinam). Gey pusililus 1844 :195. I have not found the type, but the species has generally been accepted as a poroid agaric. Modern descriptions can be found in Singer (1945:224) who placed the species in Dictyopanus, and in Burdsall and Miller (1978:85) who transferred the species to Panellus. HEXAGONIA Fr. Hee plume ie 1 OA 1 99N (PC). "Java, Blume". = H. tenuis (Hook.) Fr. H. cingulata 1844:200. The type has not been located (Java). ie cyclophora 1846:143° (PC). Mane te (Tahiti ).0.= sHeetenuits (CHook.) fr. H. dregeana 1846:143 (PC). NC. Bea. (Cape. of Good) Hope) Dreg. no. l/b.""= HH. tenuis GHook eer. H. glabra 1846:143 (PC). "Bombay (India), Polydore Roux". = Lenzites acuta Berk. H. molkenboeri 1844:260 (L). "Java, Junghuhn". = Lenzites vespacea (Pers.) Ryv. The synonymy was also noted by Bresadola (1916:231). H. pulchella 1844:200 (L). VJavalvmw= oH se tenlusm (HOOK... Er. Pe itabacina, 1654-178 (PC, isotype in kK). "Coll. Zollinger" (Java). = Cyclomyces setiporus (Berk.) Pate LENZITES Fr. L. berkleyi 1846:122 (PC, isotypes in S & BPI). "Grand bassin Saint Jean (Louisiana, USA) Mougeot no 36". =i betulina (Pr.). Fr. The type came from New York, but I have not found it. Bresadola (1920-66) had apparently examined it and indi- cated the synonymy given above. The cited specimen is selected as neotype until the lectotype reappears. LEG Deectiiatae cud no I No type has been found. No type locality was given. L. guilleminiana 1846:122. "Bresil meridional, Guillemin no. 1339" (PC). = Stipto- BB llum erubescens (Berk.) Ryv. junghubnii 1844:180 (PC). "ad truncos Java" Léveillé's hand. "Pour moi identique a halts betulina L. juvenile" Bresadola's handwriting. =“eebetulina /Crrw Jr. L. murina TOA 122) (PC te sO type. inno: - ‘Java, Korthals". Selected as neotype. The type locality is given assouma tra, DULL =nNO.such) CollecVion hasaueen found. = Trametes menziesii? :(Berk.) Ryv. Leamyriophyl lal 166322029 (PC): Myovaes déeiJ. Triana 1651-57. -Novo (Granata’, (Colombia. elegans (Fr.) Pat. Ce Se LOUUT U7 OCPC Ya SOcLY pPeSmlibeNy sombre "Java, ad truncos". = Lenzites vespacea (Pers.) Ryv. L. platypoda Loud eeCO es CEO. '"Manille (The Phillipines) ™.: = L. elegans (Fr. ) Par. LOM Ceml USO or Loot scr Cnr "Guadeloupe". = L. elegans (Fr.) Pat. POLYPORUS | Er’: P. apalus 1848:124. No type has been found. (Rentilly, France). Pemaonormasi bout 06". The type has not been found (Java). = P. sanguinaria Kl. teste Lloyd. Clo lO). Plactleatus=! G462137 00? C eisotypesin BPr). NZOdveinger no. 2055! “lava i=. trametes modes tamer re) Ryv. P. albomarginatus ©LG4W:.191 (PCy isotypes IniBPl kel). No text on label, except the name written by Léveillé. (Java). This is an accepted species in Pyrofomes as P. albomarginatus (Lév.) Ryv. For a description, see Ry- varden & Johansen (1980:529). Pe Janisopilus loud oT EGRO.) Tsotypeswin suas worl). "Java, Blume". = Coriolopsis sanguinaria (K1l.)’Teng-. Pega pposttus el oloo iy Tip Geico t yneom Tien BE eo) "Zoll. no. 19. Auf faulen Baumen in Wald bei Tjiboda (Java) Mai 1842". This is an accepted species in Phelli- nus' as P. appositus (Lév.) Pat. For a description, see Ryvarden & Johansen (1980:141). Poeatypus; Lond Cer c) "Guadeloupe, M. L'Herminier". = Trametes modesta (Fr.) Ryv. P. auriculaeformis 1844:194 (L, isotypes in BPI and “s)* "Java". = Phellinus senex (Nees & Mont.) Imaz. Peeplumelelold Looe Ty Wi soOtypes in bOrand Beh). "Java, Zoll. no. 11" = Trametes menziesii (Berk.) Ryv. P. bonplandensis 18463301, (PC. dsotynée an sBPL). "Am equatoriale". = Trametes membranaceus (Fr.) Kreisel. Pp. botryoides 16464120. (Sa sotypemin Beis) "Wragg. orig. ex." (in Bresadola's hand) (North America). = Globifomes graveolens (Schw.) Murr. 9 Paepracypusslcolo: 1? 7a(PChelsotypes ineBPT )’. "Guadeloupe, M. L'Herminier, Février 1843". = Trametes marianna (Pers.) Ryv. fan coil imorphuse 16467133 7( PC). "Voyage de M. Perville N.O. de Madagascar 1841." Tp tee seangacceptedaspecves@inyPhellinus: aseP. vcalli= morphus (Lév.) Ryv. For a description, see Ryvarden & Johansen (1980:145). Percaltochnrovice O44 Ol, The type has not been found, and no type locality was given. Precandicanse| 005205. (PC. sOUY per ine hy. "Tequendama (Colombia), alt. 2600 m, Sept 1856, Lindig Noemecoe Te = ri chantumep i Pormis, (Free ins Kl.)) RYV. Per candadulus 1846: 301" (PCRGisotype an’ Kk). "Caesarodunum" (?) = Trametes cervina (Schw.) Bres. as already noted by D. Reid on the sheet. PB. chryseus 1846:301 (PC). "Nouvelle Grenade (Colombia), M. Justin Goudot 1844. ThVsm Svaneacceprednspecicceiteriel linus, as 72 .1cory— seus (Lév.) Ryv. For a description, see Ryvarden & Jo- hansenw 0603151"). P. cinerascens 1844:184. The type has not been located (Java). P. cineraceus 1846:139. The type has not been seen by me, but Bresadola (1916: 223, as "cinereus") reports the species to be a synonym Ol irametess hirsuta er.) bis P. cohaerens 1846:132 (L, isotype in PC, BPI, FH and S). "Zollinger no. 13" (Java). = Coriolopsis asper (Jungh.) Teng. as already indicated on the sheet in herb. PC by Bresadola. Ree conrmertuswmlo4s Olu (CPG. Lsotype: Inv BPL). "In truncos Java". = Coriolopsis asper (Jungh.) Teng. P. eonnexus, 1846:135 (PC). "Brasil, Rio de Janeiro. M. Gaudichaud 1831-33". As already indicated by Bresadola (1920:68) the type collection is mixed. One specimen is Coriolopsis cape- rata (Berk.) Murr. while another is Phellinus gilvus (Schw.)@ Pat. The description is so vaguel)-thatwit may cover both specimens, and thus, the name has to be re- jected. Pe coniacenus, 16462137. The type has not been found (Nelli Gherry, India). The name is invalid as a homonym of P. coriaceus Endl. 1830. Fe Cornmugacvus) 1o46ee41 36. (li) "Java". Invalid name, being a homonym of P. corrugatus Pers. 1826. The type of the invalid name is a specimen of Coriolopsis asper (Jungh.) Teng. Pee CONVO tUS! O44 MOOMCPC. sisotypes: in L,eSte NYoand BRL). "Java, Junghuhn". = Trametes menziesii (Berk.) Ryv. P.. cyathiformis 1844:181. The type has not been found (Haiti). Peademidofiiie boue:;02) (Ss, Pisotypesin BPl ands Ky). "Frag. specimen orig. leg. Demidoff, comm. Patouillard". This is an accepted species in Pyrofomes as P. demidof- ble lev ve hOLlL ik sPoug. efor avdescription,) see Ryvar— den (1978:399). 180 P. dermatodes, see Trametes dermatodes. Paeaiii vatuse lout OlmLsy easotypeuines). "java, ex religui Junghuhnii". = TraméetessmenZiesas (Berk. ) eRY¥V. P. disciformis, DOuts 1937 Thep type has not been found! (Mauritius). Pevdissectusml S46. 13 9P¢PC):: "Herbier du Chile, donné par M. Gay". This isja Tyromyces species, and) the followings compi— nation is proposed: Tyromyces dissectus (Lév.) Ryv. comb. nov. Basionym: Polyporus dissectus Lév. Ann. Sci. Nac emcees! VOU De So ml otbe Fruitbody pileate, dimidiate with a contracted base, ADOULE 2JDRxXE oabi cma Dasemabouc Wi cmuwidem ¢—Osnmatoick, fragile. ‘Pileus applanate, margin fimbriate tovdastinct— ly Split orelobeds surface white to dirtyelicn@ Drown, smooth, azonate, finely velutinate to almost glabrous in parts and weakly wrinkled radially. Pore surface light brown, pores 4-5 per mm, entire to slightly in- cised, tubes 2 mm deep, white. Context ochraceous, 1 mm TOM Cutts | Hyphal system monomitic, hyphae 2-5 um wide and with clamps, partly gelatinized in 2.5 2% KOHes in) Melzerewith walls about 0.5 um thick, sparingly branched. Some hy- phae have a dense yellowish protoplasm, distinct both in KOH and Melzer, 2-6 um wide and with scattered clamps, these hyphae may be interpreted as oleiferous or gloeo- pleurous. Cystidia and basidia not seen. Spores subcy- lindrical to oblong ellipsoid, hvaline; thin-walled and non—amyloid,. 4=-5.x 2-2.5 um. As long.as onlyethe: typeris known, thesmacroscopical characteristics Nave fo, besused with care. This taxon may be the same as T. floriformis (Quel. ) Bond. & sing lheiremicroscoplecal eharacveristicsaare 1dentical and? the, fruitbodles arewsimilar. 1) flordior= mis as collectedisins Buropel ie usualy morespurer wove. buty themdarkerUcolours of tnemty pe Ole l.edLesecLusias certainly due to bad preservation after it was collected in Chile and then sent to France. Pe dozyenus 1646 si23. The type has not been found (Java). Pe elatus 4] O46;129. The type has not been found (Guadeloupe). PV nex tensiswlo40e lo Um PCO): "Guadeloupe, L'Herminier". This is an accepted species in Phellinus as P. extensus (Lév.) Pat. For a descrip- tion, see Ryvarden & Johansen (1980:158). Pe. fastuosus, LO44eA 90 Nina teotype anbPl). Jotnpapore!. Tne species Tseacceptedeunarhne: Linusewacgmr « fastuosus (Lév.) Ryv. For a description, see Ryvarden & Johansen (1980:159). Peiruscelius, Lo54- 7 Flow Jeouypecwlneronanda la. "Insul. Flores prope Balie". = Coriolopsis asper (Jungh. ) Teng. P.tarusenus. O46: Lavoe POE soLyne in "ikl. "Zollinger no. 1454" (Java). The type is a specimen of what Berkeley later described as P. setiporus,:a species 181 which usually is placed in Cyclomyces. The type species Cie bem obict isto. tUscusenr © 1033 making it illeriti- mate to transfer Léveillé's name to Cyclomyces. However, should anyone want to place the taxon behind Léveillé's and Berkeley's names in a genus other than Cyclomyces, then Léveillé's name has priority. eeeeaudichaudii 1846p. 178) (PC, Wsotypes' in S & BPI). "Singapore 1839". = Trametes menziesii (Berk.) Ryv. ecavyanus 1 O46r1 ofr (PC) Meniules Mem GlomGay..o30%.8 =—8Trametes marianna (Pers. ) Ryv. P. gibberulosus 1846:139 (PC, isotypes in K & FH). "Guyana, Batara Reidweg, Wegel no. 568". = Trametes vil- HOSa HOOK) RY Ve Pemecossypinus PG4sGe 124 (PC, isotype in’ BPI). "Ad truncos in Voges (France)". = Tyromyces caesius Chr Munim ie uademoupensis Post onl34(PC, isotypes tn. Sie® NY). Wevuadeloupe, M.i;'Herminier". =| Fomitopsis supina (Fr: ) Ryv. as already indicated by Bresadola on the. original label. Pemnasvoerl1 i OU 190N ti, isotypes in PCek. BP): Java. =—sPhellinusspectinatus (K1l.)) Quel. as indicated on the sheet by Bresadola~ Prenasse titi wl Ouse 187% The type has not been found (Java). P. heteromorphus 1846:123. "Guyana". = Amauroderma schomburgkii (Mont. & Berk.) Torr., as already indicated by Furtado (1968:268). erenymenius 1663°263° (PC, isotype in’ kK). chien Mee I OOROU LI etl co0"*CColombial., = Coriolopsis brunneo-leuca (Berk.) Ryv. The type: is unfortunately sterile. I have not seen the species fertile from the neotropics, and the dextri- noid reaction found in the skeletal hyphae of many Afri- can specimens was not seen in Léveillé's specimen. It may be that two macroscopically similar species are in- volved. Fertile material from the neotropics may shed some light on the problem. Mem nouc na tus OuOMLNORC PG isotypes: in FH & BPI). BeeruoOtvetyanelid Ghnerry, India orientabist.—= Trichap-— CuUMme DL OrmissnGrreeun Kile)” RyVes Be kKickxianus 18482122. The typeshas not been found (America). Eereorthalsi i091 844-9190 3h): "Sumatra, Korthals". = Phellinus senex (Nees & Mont.) Imaz. feerlcnicwmicloc lesm (PC. isotype in’ BPL)* "America, ad truncos". = Coriolopsis polyzona (Pers.) Ryv. Pomlenziteus 1654-17>°(L, isotype in PC). "Zollinger 975 (Sumatra)". = Lenzites acuta Berk. P. leucomelas 1846:140. The type has not been found. Invalid name, non P. leuco- melas Pers. 1825. eo indivi so3e263 (PC, asotype in’ K)). "Tepuendama (Colombia) Canoas alt 2600. Sept. 1860". LBZ = Trametes cfr. modesta (Fr. ) Ryv. The collection is fer= tile with spores 6.5-8 x 3-3.5 um. These measurements are in accordance with those of Fidalgo & Fidalgo (1968:27). The spores of the paleotropical taxon called T. modesta and macroscopically identical with that of the neotropics, are seemingly smaller. However, a richly fertile specimen has never been found and no sporeprint has been made. P.. dongipes® 184621249 (PC; isotype, in’ BPL)« "Guyane francaise". This is an accepted species and-the type of Haddowia, Stey. in the Ganodermataceae. For a description, see Ryvarden & Johansen (1980:93) with fur- ther references. P. macropus 1848:122. The type has not been found (Java). P. mangiferae 1846:130 (PC). "Voayge de M. Perville. N. East de Mahe (Tahiti) 1841". The species belongs in Ganoderma and its final status has to be decided when the genus has been revised. Premanubprelatus foots lyauUler "Zollinger 974" (Sumatra). = Microporus scopulosus (Berk.) Ryv. Pema ren onc sme) od Os 00n The type has not been found (Marguesas Islands). Paminastoporus Lolobel ;omure).. "Singapore, Février 1832". This is a Ganoderma species im thesG. =huctdum= complex. =Ltsestacvus nas stom bercetered when the genus is revised. P. megaloma 1846:128. The type has not been found (New York, USA). In American literature it is commonly assumed that this is the same as Ganoderma applanatum, see Overholts (1953:99). From - ther déeseription itsistvery probabil yeso-. P. melanevis1046.131) (linen sotypes dneoes eBrion. "Zollinger no. 2085 (Java)". = Coriolopsis caperata (Berk.) Murr. P. melanaleucus 1846:141. The type has not been found (Réunion). Pe imicrocyclius 1049 +168 “CLs wsotypes ine POsiKeeeNY. mb bie)e "Java". = Cyclomyces tabcinus (Mont.) Pat. Peomicrolona aaa; 183. The type has not been found (Manilla, The Phillipines). At Kew there is a specimen: "Phillipines 1908, Curran, cum. typo comparato’™ ink, Bresadola's hand. This isva specimen of Microporus affinis (Nees & Blume ex Fr.) Kunt. The specimen mentioned above is selected as neo- typemuntile authenticemateriadwisefounds, P. Mmorutzianus) 1 o40s1 301 Cla. "Zollinger 360 (Java). = Trametes marianna (Pers.) Ryv. Pi. murdinus! 16444185 (i sasotypes an, POsanderi)s "Java". = Trametes menziesii (Berk.) Ryv. P. nephelodes 1846:125. The type has not been found (Paramaribo, Brazil). In the Stockholm herbarium there is a specimen: "182 P. nephelodes Lév. = Polyp. flabelliform. Kl. transiens Ja- va v. Héhnel" Bresadola's hand. The specimen is Micro- porus affinis (Blume et Nees ex Fr.) Kunt. However, as the genus Microporus is not known from South America, b8sS the specimen from Java cannot be selected as neotype of P. nephelodes. P. nordmannii 1842:93. The type has not been found (Korbek, USSR). Fries (1874:581) refers the species to Heterobasidion annosum (Fr...) Bref. but it is not stated whether he had seen the type or based his: suggested synonymy on the descrip- tion alone. The latter is rather vague, but could apply tOVasresupinate specimen ofvH. annosum. P. notopus 1844:194 (PC). "Java". = Trametes modesta (Fr.) Ryv. Pemostreabusmilodo lo 7m PC erisotype IneBel). "Bresil, Rio de Janeiro, M. Gaudichaud 1831-33". = Tra- metes cubensis (Mont.) Sacc. P. pala 1844:183. The type has not been found (Surinam). Peepeetunculus! O46413 acre). "Indes orientalis, Nelli-Gherry (India)". = Trametes versicolor (Fr.) Pil. P. perpusillus 1844:191 (PC). "Am. Boreal". = Fomitopsis scutellata (Schw.) Bond. & Sing. ar P. phaeus 1846:132. The type has not been found (Sri Lanka). Peeplacopusraldtbalotas )i "Java, Junghuhn". This is a Ganoderma sp. in the Gano- derma lucidum complex. Its status has to be settled when the genus is revised. P. platypilus 1844:192. The type has not been located (Java). Pee pkumbeus: 10462136 (PC, Asotype inwS). "Guadeloupe, M. L'Herminier". = Rigidoporus fusco-linea- CUSMULECS ony VE P. rhodophaeus 1844:190 (S). "Java". This is an accepted species in Fomitopsis as F. rhodophaeus (Lév.) Imaz. For a description, see Ry- varden & Johansen (1980: 340). P. rigidus 1844:189 (L). "Java, Zollinger no. 732". = Nigroporus durus (Jungh.) Ryv. Rea Pudis a. C4643 30 (PC). "Guadeloupe, M. L'Herminier". Invalid name, non P. rudis Berk. 1839. The type of the invalid name is Fomitopsis BUDS BOE eh VV. P. rugulosus 1844:189 (L). NNow57'" CJavaja(leg. Junshubn?,). i= Coriolopsis sangui— naria (Kl.) Teng. P. seclerodermus 1846:129 (PC). "Guadeloupe, L'Herminier". = Fomes fasciatus (Fr.) Cooke. P. ‘scleropodius .1846.123 .(BPI.). "Tle Bourbon (Réunion)". = Lignosus sacer (Fr.) Ryv. P. sericellus 1846:125. The type has not been located (Sri Lanka). Pawetoeroides 104u lee (PCeisotype in kt). "Sumatra". This is an accepted species in Coltricia as G. sideroides (Lév.) Teng., for a description, see Ry- varden & Johansen (1980:112). Léveillé named also Coll. 184 Zollinger No. 90 as P. sideroides, but this collection is a specimen of Phylloporia chrysita (Berk.) Ryv. Dup- licates of this collection have been seen in herb. L, Seer rigger On and mb inks P.-sordidus- 16442192 (PC) isotype ins). "Guadeloupe, L'Herminier". = Trametes modesta (Fr.) Ryv. P. splendens 1844:187 (PC, isotype in L). "Java". = Trametes modesta (Fr.) Ryv. P. spurcus ,1946°1350(PCy isotypesvin’ S)- "Guadeloupe, L'Herminier". = Phellinus gilvus (Schw.) Parts P. stevenii 1842:91. The type has not been seen. Lloyd (1915:285) indicates it to be Ganoderma applanatum based on the original des- cription and plate. Donk (1974:67) agreed, and I sub- scribe to the conclusion. Pe subf lavusau c4or 300m (PC)E "Nouvelle Orleans, Salle". = T. versicolor (Fr.) Pil. P. swartzianus 1846:132. The type has not been found (Jamaica). Po tegularise1 040-1 319eGPC mn SOLy peal tmnt). "Zollinger 2054 (Java)". = Trametes scabrosa (Pers.) Cunningh. Piemtvenaxelo40 >: 139% The type has not been located (Teguendama, Colombia). P.. tener 16465139 CPC)e "Guadeloupe, M. L'Herminier". = Trametes villosa (Hook.) Ryv. P. tenuissimus 1844:188. The type has not been found (Java, leg. Korthals). Petestaceus 91 S46" 126g(PC)™ "Brasilia meridionalis, Dupré, 1842". This is a Ganoder- ma sp. in the G. lucidum complex. Its status has to be settled when the genus has been revised. P. trachodes 1844:192. The type has not been found (Java). Pemetbricolor slot be 34e The type has not been found (Bahia, Brazil). Petunia conus 640-15 deer cee "Zollinger No. 2069 (Java)". = Coriolopsis sanguinaria (Kis mere np. Pamitbris ti smlo4b tc OmGRpCeaeisot ypesinakhy wZoLVnger |\No-=2035° (ava )feuinvaliduname,, nontr.ntris— tis Pers. 1825. The type of Léveillé's invalid species is a specimen of Nigroporus vinosus (Berk.) Murr. P. unguiformis 1846: 13% (PON@isotypemain fH). "Zollinger No. 1437, Tjiboya (Java)". = Rigidoporus mi- croporus (Fr.) Overeem. Peoevolneratusml cn veloc ec. "Ad truncos, Java". = Coriolopsis sanguinaria (K1.) Teng. P. zollingerianus 1846:131 (PC, isotypes in S and FH). "Zollinger 1386 (Java)". = Trichaptum byssogenus (Jungh.) Ryv. 185 SISTOTREMA Fr. S. ochroleucum 1846:145 (PC). "Bombay (India), Polydore Roux" = Lenzites acuta Berk. TRAMETES Fr. ieeacuta 1LoW4s196. The type has not been located (Sumatra). Specimens deter- mineduby sbréesadolavin herb. So and Bri eare all Coriolop— sis strumosa (Fr.) Ryv., and this was probably also Lé- veillés concept. Tercrassanouy soya ko). "Madagascar". = Hexagonia hirta (Fr.) Fr. T. dermatodes 1844:196 (PC). "Voyage de M. Gaudichaud sur la Bointé. 1836-37. Manille (The Phillipines)". = Oxyporus cervino-gilvus (Jungh. ) Ryv. as already indicated by Reid. (1953:135). Pei ncanagi cud] 06m Bre "Manille (The Phillipines)". This is an accepted species in Trametes, for a description, see Ryvarden & Johansen (1980:565). ieeperrottetiiy 16441 95 (PC). "Guyane francaise M. Poileau" (printed on the label). "Java, M. Perrottet", unknown handwriting. The name, however, is written by Léveillé. There has been a mixing of labels or facts. The type represents a South American taxon which has never been found or reported from the paleotropics. The type is ap- parently the same specimen which was used by Montagne and Berkeley when they described Trametes trichomallus in 1849. The latter is the type of Trichaptum Murr. Trichaptum perrottetii (Lév.) Ryv. NOOW sted ee DOL elo 23 (an Lope. Fruitbody applanate, sessile, semicircular to elongated shelf-like, mostly broadly-attached, usually not decur- rent on the substratum, 5-15 cm long, 3-/ cm wide and to 8 mm thick (tomentum not measured), tough and flex- ible. Pileus with a dense, strigose to villose or hir- sute layer of forked hairs, dark brown, becoming darker towards the base and more greyish towards the margin, azonate or weakly-zonate, up to 10 mm thick at the base. Margin entire and sharp. Pore surface snuff brown, pores angular to round, first entire and thin-walled, 2-3 per mm, in older specimens often incised, coalescing and in parts sinuous to daedaleoid, in the latter case up to 2 mm wide and several mm long. Tubes deep brown, 2-5 mm deep. Context very thin, 0.1-0.4 mm, brown to dark och- raceous. Hyphal system dimitic, generative hyphae thin- walled, hyaline and with clamps, 2-4 um wide. Skeletal hyphae abundant, thick-walled to solid, mostly yellowish to light brown, 3-5 um wide. Cystidia abundant in the hymenium, clavate to ventricose with a tapering apex, smooth or with an apical crown of crystals, 10-18 um long, slightly projecting, basidia clavate 12-15 x 4-6 um and with 4 sterigmata. Spores subcylindrical to ob- long ellipsoid, smooth, thin-walled and non-amyloid, 5-7 x 2-3(3.5) um. On deciduous wood. Specimens have been examined from USA, Cuba, British Honduras, Guatemala, Costa Rica, Colombia, Brazil, French Guyana and Bolivia. Tee vel tba var hod delOG™. The type has not been found (Sumatra). ACKNOWLEDGMENTS The staff of the herbaria in Stockholm, Leiden, Paris, Kew, New York, Harvard and Beltsville are most kindly thanked for their cooperation during my type studies. The Norwegian agency for International Development has given fi- nancial support for which I am very grateful. R. Winter has suggested improvements in the English text. REFERENCES Bursdalil eH. Heir. SaMiler.) Oserk.n OO rl ereevaluuatdormon Panellus and Dictyopanus. Beiheft. Nova Hedw. 51:79-91. Donk, M. A. 1974: Check list of European polypores. Verhand. Afd. Naturkund. Kon. Nederl. Akad. Wetensch. 2 ser. no 62. Pidalgo, eM. ies Pak. kwridal goles 0.01960 sePokvuporacescmiicom Venezuela I. Mem. New York Bot. Gard. 17, no 2:1-34. Fries, E. 1874: Hymenomycetes Europaei. Uppsala. Furtado, J. S. 1968: Revisao do genero Amauroderma (Polypo- raceae). Sao Paulo. Priv. ed. Léveillé, J. H. 1842: Observations medicale et enumeration des plantes, p. 35-237 in Demidoff, A. (ed.) Voyage dans la Russie meridionale et la Crime. Vol. 2. Léveillé, J. H. 1844: Champtgnons exotigques._ Ann. Sei. ‘Nat. Det een. lem eO Giclee. Léveillé; J. H. 1846: Description des champignons de l'her- bier vdu Museumsede) Parts Ann.) ooCl Na Camel es my Oem ea oy a Léveillé, J. H. 1846b: Fungi, p. 164-204 in Gaudichaud, M. (ed.) Voyage autour du Monde sur la corvette La Bonité. Botanidue.9 V0. ous lk. Léveillé, J. H. 1848: Fragments mycologiges. Ann. Sci. Nat. Ser. 3. Vols. '9:119-144. LEevetie gc) eh. sLood ce hung igeip. Bobo ZOlsme riper mtd s) Systematische Verzeichnis der im indischen Archipel in Jabren 1842-48 gesammelter sowie der aus Japan emp- fangenen Pflanzen. Léeves ese J eH. e166 3 ceehunet Sine teriang eM. wnieues eel anchors J. E. (ed.) Florae nova-granatensis. Ann. Sci. Nat. Sere UmVol. 20262-3800) Lloyd, C. G 1912: The polyporoid types of Léveillé at Lei- den. Meded. Rijks Herb. Leiden 9:1-5. Lloyd, C. G. 1915: Synopsis of the genus Fomes. Lloyd Mycol. Writ. 24 eo ld —206. Reid, D. 1953: New or interesting records of Australasian Basidiomycetes III. Kew Bull. 12:127-143. Singer, R. 1945: The Laschia-complex (Basidiomycetes). Lloydia 8:170-230. MYCOTAXON VOT AL ELE Nomal ppl 87-190 April-June 1981 THE PUBLICATION DATE OF ARENDHOLZ'S THESIS ON LEAF-INHABITING HELOTIALES WOLF-RUDIGER ARENDHOLZ Biologie, Universitat Kaiserslautern 6750 Kaiserslautern, West Germany and RICHARD P. KORF Plant Pathology Herbarium, Cornell University ithacay. New!) York 14853 7USA SUMMARY July 7, 1979 is shown to be the date of effective pub- lication of W.-R. Arendholz's doctorate thesis, 'Mor- phologisch-taxonomische Untersuchungen an blattbewohn- enden Ascomyceten aus der Ordnung der Helotiales," and the date of valid publication of the new taxa and new combinations proposed therein. Tiveesta usmiote pUDlICationj#o!l BWs—Rk ae Arendholz!s | Pheb: thesis, entitled "Morphologisch-taxonomische Untersuch- ungen an blattbewohnenden Ascomyceten aus der Ord- nung der Helotiales'' was assumed to be "unpublished" Pree KOnimalcdenkOnneulLjooymemmmorder tOmprotectthesstatus Of VArendholz.s. new names, they reported) one, as an as yet unpublished new species referred’ to the genus ~w slhey ‘shad received in’ the mails from’ Arendholz a copy of his thesis (Arendholz, 1979), a soft-bound volume having the appearance of being some sort of photo-copy of the original typewritten thesis on deposit with the Universitat Hamburg. The purpose of this note is to call attention to the fact that over 180 copies of this thesis were actually distri- buted. eandy inateiny no, way can, the» new, names, therein be ignored or treated as not validly published. On July 7, 1979, 139 copies were mailed by Arendholz to the 188 Staats— und Universitdtsbibliothek, Hamburg, which sub- sequently distributed them to other libraries, and that date is held here, to constitute the (date otmdistri bution CONSLITUTINGeellectivempublications sundeissticasbnicomdil onal Code of Botanical Nomenclature (Stafleu et al., LOU Seem Gt Cc lome2 J .61% Twenty further copies were sent tO the = Wibranyemol mache? SINStitutiethuL ee |CemelneeeDOLduik ders Universitat Hamburg ss andysanother a 2 5mund cee il) samec scientists in different countries. Approximately) 20 addi— tional @ copies: pancees Ea nvweALendn Olzcm DOS SCSSoIOUmm OULU of a photo-offset lithography press run of some 200 cOpLes. While it seems unfortunate. that.the copies that were dis-— tributed) Camry, ino indications of sthesexactsddatesOlmars ume Dution, —nor Vveven wthats themvolume sconsuitMies samspubDlica— tion in the sense of the Code of Nomenclature, recent articles on “effective” publication »under VArticles® 29-3 lmot the ~Codestleavem nowdouDl ming our nincde ta teethometiesis was jcifectively, published men! Nemes fOurematliCleSmal imal acon 29(4), the August 1980 issue, support our _ position [Brummit (1980); Hara and Eichler (1980); Nicholson (1980); Weresub and McNeil (1980) ]. Since thessthesisieiS = printed byes pnoto-Ofisetmlithograpiye it must apparently be accepted as printed matter (Art. 29): “... it may safely be assumed that matter pro- duced’ *' by 9a conventional Gtypewriter, « Consisting Olgsone Lope copy, plusmone. to several carbon scoples. =i sanetncol. sidered sto) be, printed. On ties other ihand = copyeos ginal ly produced on a typewriter and then reproduced by off- set lithography (as.for example the important taxonomic journal Mitteilungen der Botanischen Staatssammlung Miinchen) must certainly be accepted as printed matter" (Brummitts 1980-87 =4/0)8 The, Codes (Art 9291) requires sthateithes printeds matter bes@diptributedmtomthemoceneralmpublicwoumdteslcastmtompo— tanical institutions with libraries accessible to botan- iStSeeoercna | Ly. eNed (Liem tiem Un DelmOtmcOD1es mmo tmL Tc number of libraries: (nor even that they be located in more than one country) is stipulated in the Code, "and presumably in an extreme case a printed work distribu- ted to as few as two libraries should be taken as ef- fectively published" (Brummitt, 1980: 479). In the case in point, the requirements for effective pub- lication would have been fulfilled by mere deposition of one copy of the printed thesis in the "Staats— und Universitdtbibliothek" and one in the library of the '"In- 189 sxiiuteeitir “Allgemeine ~—Botanik.” As noted above, many more than two copies were distributed to several libra- ries and scientists worldwide. Nicholson's (1980). key Similarly keys out this thesis as "effectively published." Establishing an exact date of publication of any prin- heaesworkt is icritical for nomenclatural purposes. For journals, the practice is to use’ the actual date of mail- ing of each issue as publication date [this is the procedure with MYCOTAXON, as it is also with MYCOLO- GIA (C.T. Rogerson, Managing Editor, pers. comm.) ]. Similarly, commercial books are published on the date they are actually offered and available for purchase, not that on which they actually reach the public or li- braries" (Brummitt, 1989: 480). Since the bulk of the copies of Arendholz's printed thesis were mailed on Puy on? je) 19/9, tom the Staats—) und Universitatsbibliothek, HagiDunewathats date shouldmnow be “accepted as the date of publication of the thesis for all nomenclatural pur- poses. Various proposals by Hara and Eichler (1980) and by McNeil (in Weresub and McNeil, 1980) focus on the kind Ol problem presented by ‘Arendholtz’s thesis with a view tO eliminating any problems in the future, with worth-— while suggestions for requiring an indication of inten- Gion to molfer material “as ~“*published’” under the provi-— PiencmoOtmiies COde,marequirements {fonsdeposition im ajspe— cific number of (named) institutions, and indication of actual date of publication (distribution). Such propo- sals will be useful in the future, but in any case will not be retroactive, but apply from 1982 onward if they are adopted at the International Botanical Congress in sydney later this year in the forms submitted. REFERENCES CITED ARENDHOLZ, W.-R. 1979. Morphologisch-taxonomische Unter- suchungen an blattbewohnenden Ascomyceten aus der Ord- nung Helotiales. Ph.D. Thesis, Univ. Hamburg, 115 pp., fom mCP ri vatelvyedistributed 7 July 19/79) BRUMMITT, R.K. 1980. Questions on effective publication. Taxon 29: 477-483. HARA, H. & H. EICHLER. 1980. Proposal (67) to amend Art. 29.1 on effective publication and recommendation to esta- blish a special committee on Art. 29. Taxon 29: 515-516. MOR AER Pec lte MM KOHN peu lO7G |) Revisionary studies “inethe Hyaloscypheae. I. On genera with "glassy" hairs. Myco- taxon 10: 503-512. 190 NICHOLSON, D.H. 1980. Key to identification of effective- ly/ineffectively published material. Taxon 29: 485-488. STAFLEUs, F-As. et al. feds.|] 1978. Internationakecodesot botanical nomenclature adopted by the twelfth Interna- tional Botanical Congress, Leningrad, July 1975. Regnum Veg. 197 :9i-xiv, 1-457. WERESUBY OLN Ke & 0 Hea McNeilo) 1980.) Et recuives publicatron under the Code of Botanical Nomenclature. Taxon 29: 471- 476. MYCOTAXON VOLT OMULE NO we ppeato1-240 April-June 1981 AN ALTERNATIVE VIEW OF CERTAIN TAXONOMIC CRITERIA USED IN THE ENTOMOPHTHORALES (ZYGOMYCETES ) RICHARD A. HUMBER USDA-SEA-AR Insect Pathology Research Untt Boyce Thompson Instttute Tower Road, Ithaca, New York 14853 SUMMARY Long-standing unresolved controversies about the various generic classifications of the entomopathogenic Entomophthorales require a review of the validity and the relative weights of the morphological and other taxonomic criteria used to construct a useful classification for this group. Major morphological criteria used in the Batko and Remaudiére classifications are evaluated here in a survey of the taxonomic distributions of these characters and the Significance of the irregularities in their distributions. The number and nature of nuclei in the primary spores, the branching of sporophores, and the mode of discharge of primary spores are regarded as the major characters suit- able for delimiting genera. The presence of rhizoids and (to a limited extent) the shapes of primary spores are Significant secondary generic characters. The presence of cystidia, mode of resting spore formation, and presence of capillary secondary sporophores are not regarded as char- acters of generic importance. The conidial or sporangial nature of primary spores is discussed, as are difficulties in the interpretation and applications of the terms 2ygo- spore and azygospore. bo2 The applications of the generic names used in the Batko and Remaudiére classifications are reviewed in light of the findings reported here. The validity of both Stronguellsea and Entomophaga is upheld, and the sense of Erynia is restricted to those species having branched sporophores and uninucleate bitunicate primary spores which do not form secondary spores on capillary sporophores. A nomenclatural problem is noted which indicates the synonymy of Zoophthora and Erynta if these genera are not retained as separate as proposed in the Remaudiére classification. Tabanomyces is regarded as a synonym of the nematophagous genus Mertstacrum. INTRODUCTION A far-reaching revision of the entomopathogenic genera of the Entomophthorales (Zygomycetes) proposed by Batko (1964a-e, 1966; Batko and Weiser 1965) has spawned consid- erable controversy among the students of these fungi. Even though there is now wide agreement that it is unsatisfac- tory to continue classifying approximately 150 diverse species in the single genus Entomophthora Fresenius, there has been no agreement regarding the appropriateness of the criteria used by Batko to restrict Entomophthora and to establish segregate genera or of the circumscriptions of these genera. King and Humber (1981) offer a basic reconsideration of the major morphological characters of this group and discuss their taxonomic significance. While King and Hun- ber evaluate the basic strengths. and weaknesses of the Batko classification, they consider that no definitive re- working of the genera would be appropriate without a final resolution of the long-standing problem of delimiting Contdtobolus Brefeld from Entomophthora sensu lato. More recently, however, a major reworking of Batko's generic classification by Remaudiére and Hennebert (1980) and Remaudiére and Keller (1980) — hereafter referred to as the Remaudiére classification — resulted in generic assignments for some species which led to no manifest in- crease in the biological homogeneity of the recognized genera. Many taxonomic decisions in the Remaudiére classi- fication, and particularly those regarding the validity and Eos importance of various characters used to construct classi- fications of these fungi, were presented without explana- mroueend, in tury, have triggered further controversy and alternate views on classification (Ben-Ze'ev and Kenneth 1981a; Humber 198la). The time is appropriate to examine thoroughly those morphological and developmental characters which have been employed in the modern classifications proposed by Batko and by Remaudiére and his colleagues. The values of these criteria are examined in the contexts of the available classification systems. Indeed, a sense of the relative values of their chosen criteria does emerge from a consi- deration of the major schemes of entomophthoralean genera and of the internal inconsistencies generated by the selec- tion and ranking of the criteria accepted as valid in each of these generic schemes. Despite 1ts undisputed desirability, no general agree- ment regarding the generic classification of the Entomoph- thorales will be possible until the morphological and other emicreria used ini the past or being proposed now have been critically reviewed, evaluated, and agreed upon. This paper is the first in a series discussing the merits and weaknesses of the Batko, Remaudiére, and other emergent Slassitications: (€.g., Ben-Ze'év and Kenneth 198la) in an attempt to find rational bases on which to build a generic scheme which (1) avoids the inconsistencies of the previous classifications of the Entomophthorales, (2) can accurately predict the natures of characters or aspects of the host- pathogen relationship, and (3) reflects what appear to be the evolutionary relationships among these fungi. TAXONOMIC CRITERIA: ANALYSIS AND CONCLUSIONS PRIMARY SPORES Primary Spores: Conidia or Sporangia? Throughout this paper I refer to primary spores or spores rather than to the more conventional contdium or the less common sporangium or sporangtole. Contdtwn and spor- angtum are terms referring to specific developmental pat- terns; their use should be for purposes of accuracy and must be backed by sufficient demonstration of the mode of 194 sporogenesis. The reference to any asexual propagules in the Zygomycetes as contdta has been questioned by the stu- dents of the Hyphomycetes and Coelomycetes, although there is no reason to believe a prtort that conidia cannot be formed by these or any other lower fungi. If appropriate studies of the origins and fates of the various wall layers of spores (and sporophores) during sporogenesis and germi- nation prove that,indeed, they are conidia (or sporangia), then these propagules should be referred to as such. Simi- lar studies are also needed for the panoply of secondary spore types produced in the Entomophthorales. Spores containing single nuclei and borne on branched (or simple, in Strongwellsea Batko §& Weiser emend. Humber) sporophores have caused much interpretational difficulty. These spores have a layer which, in liquid, lifts away from the spore surface and balloons outward. This has been re- garded variously as a gelatinous coat (é.g., Entomophthora gloeospora Vuillemin 1886) or, more frequently, as evidence for the sporangial nature of these fungi by regarding the detached layer as a sporangial wall enclosing a single Sporangiospore (Thaxter 1888; Strong et al. 1960; Batko 1964b,, 1974)... This typéesot primarysspore seems, Lompcscn= tirely restricted to the species of Zoophthora Batko, Erynta Nowakowski, and Strongwellsea. Kenneth (1977) claims that the outer wall of Entomophthora turbinata Ken- neth [= Weozygttes turbtnata (Kenn.) Rem. §& Keller, a gen- eric assignment which remains debatable] lifts away but admits (personal communication) that this feature is doubt- ful and requires further investigation. If the spore of E. turbtnata truly were bitunicate (with a separable outer wall layer), it would be the only instance of this charac- ter in a Species with multinucleate spores. Humber (1975; King and Humber 1981; Roberts and Humber 1981) notess ultrastructuralyevidences that thessporesos Strongwellsea magna Humber, which has a separable outer wall layer, cannot be a monosporic sporangiole. The spore wall has no indigenous (sporangiospore) layer that is not shared with) thes sporophorepitse lt ainesseparab les lavenmars not a distinct (sporangial) wall, but represents only a separation of the outermost layer of the sporophore wall, all layers of which are continuous over the entire spore surface except for the papilla — which is covered only by an extension of the inner wall layer of the spore and sporophore (Humber 1975). This spore might be a true coni- dium, but this cannot be confirmed without also determining LoS the fates of the wall layers during germination (see Dykstra 1974). The only evidence for the occurence of sporangia in the Entomophthorales comes from several species of Bastdto- bolus Eidam, in which there are numerous instances when the cytoplasm inside a cell (usually but not always a spore) may be divided into several independent uninucleate units; B. mtcrosporus Benjamin (1962) provides the most striking examples of this behavior. I use the terms unttuntcate and bttuntcate introduced by Remaudiére and Hennebert (1980) and Remaudiére and Kel- ler (1980) to refer to spores on which a separable outer woeelayer siseei ther absent, or present, respectively: These terms are used almost exclusively within mycology to refer to the morphology (and function) of asci in the Asco- mycotina, and constitute a primary character dividing the (bitunicate) Loculoascomycetes from all other (unitunicate) classes of this subdivision (Ainsworth et al. 1973). The use of these terms, however, should be generally accepted for the Entomophthorales since they (1) are convenient, (2) respect their etymological derivations, (3) should not be in any way confused with their applications for the asco- mycetes, and (4) in the case of bitunicate spores, would help suppress the notion that such spores are sporangial in nature. Morphology The variety of shapes of primary spores in the Ento- mophthorales has been the source of much attention, parti- cularly among those who have included nearly all entomo- pathogens of this group in the single genus Entomophthora. A series of artificial species groupings based on spore morphology have been proposed primarily to aid species identification (Lakon 1919; Hutchison 1963; Gustafsson 1965; MacLeod and Muller-Kogler 1970, 1973; MacLeod et al. 1976; Waterhouse 1975; Zimmermann 1978). None of these authors, however, intended their groupings to have taxono- mic value. Remaudiére and Keller (1980), however, chose the morphology of primary spores to be the main criterion for their generic classification. Several genera do have characters of spore morphology which are unique to themsleves. One of the strongest and most valid criticisms of the Batko classification, in fact, 196 was its separation of the species having campanulate spores (with relatively flat bases and a strongly apiculate apex) into Entomophthora sensu stricto and Cultetcola Nieuwland depending upon the absence or presence of rhizoids. Remau- diére and Keller (1980) rejected the significance of rhi- zoids for defining the genera, and redefined Entomophthora to include only species with this distinctive campanulate spore shape and associated mode of spore discharge (see MacLeod et al. 1976; Samson et al. 1979). The spores of Triplosportum (Thax.) Batko (nom. gen. conserv. prop., see Humber et al. 1981; = Weozygttes Wit- laczil) are variable in shape, but most often are round to slightly elongate and have a truncate rather than a conical or rounded papilla. As circumscribed by Batko (1964c), the spores of Cultctcola species could be either campanulate (like those of #. muscae (Cohn) Fres.) or obpyriform with a smooth apex and rounded papilla; species with the former type were restored to Entomophthora s.str. while those with the latter type of spores were transferred to Contdtobolus by Remaudiére and Keller (1980). King (1976b, 1977) regards Contdtobolus species to have round to pyriform pri- mary spores, and notes the similarity of several entomo- pathogenic Entomophthora (sensu lato) species with Contdto- bolus. Remaudiére and Keller (1980) extended King's con- cept of Contdtobolus by transferring all entomopathogenic Species with rounded to pyriform spores to that genus. The species of Entomophaga Batko have pyriform spores with a rounded apex and closely resemble the spores of Contdto- bolus species. These two genera are unambiguously differ- entiated, however, by both the morphology and stain reac- tions of their nuclei (Humber 1981b). The primary spores of Zoophthora Batko sensu Remaudi- ére and Hennebert (1980) (= Zoophthora subg. Zoophthora Batko sensu Ben-Ze'ev and Kenneth 198la) are rather cylin- drical and usually taper apically to a blunt cone or a poorly defined point; just above the conical papilla, the spore flares to a very slight (but not always apparent) shoulder. The spores of Erynta Nowakowski species (see Remaudiére and Hennebert 1980) display the greatest shape variation of any genus in the Remaudiére classification. They range from elongate fusiform with a marked curvature and a shallow rounded papilla to somewhat cylindrical to ovoid or fusiform spores with rounded apices and papillae canted away from the spore axis. The spores of Strong- wellsea species fit well within the range of morphological 197 variation found in Erynta, a fact which seems to have moti- vated Remaudiére and Keller (1980) to merge this genus into Erynta. The spores of Empusa carolintana Thaxter, which Remaudiére and Hennebert (1980) transferred to Frynta, are obovoid to pyriform, a shape corresponding more with those of Hrynta spores than with any other genus. Karyology: Number and Type of Nuclei Early studies noting the taxonomic importance of entomophthoralean nuclei have been largely ignored (Vuille- min 1895; Cavara 1899a-b), so that no surpassing importance was attached to the number of nuclei in primary spores until the proposition of Batko's classification (1964a-d, 1966; Batko and Weiser 1965), which necessarily drew upon the observations of earlier workers regarding this cyto- logical detail. One of the most significant characters used by Batko (1964b) to establish Zoophthora was that of uninucleate primary spores, a character which is readily detectable and seems to be correlated well with the other characters of this genus. Strongwuellsea Batko §& Weiser (1965) was also described as having uninucleate spores but was clearly dis- tinguished from Zoophthora by its simple sporophores and unique habit. EHrynta Nowak. was split from Zoophthora by Remaudiére and Hennebert (1980) and is circumscribed so as to allow multinucleate primary spores even though £. caro- lintana is the only such species included in E£rynta by Remaudiére and his colleagues. As discussed at length be- Low, thesmorphological icharacters of this species do not appear to belong in this genus, and its probable misclassi- fication is clearly indicated by its status as the only species with multinucleate spores in this large grouping of species with uninucleate spores. As is the case with branched sporophores (see below), there is a very low but natural incidence of abnormally developed primary spores. The spores of Zoophthora, Erynta, and Strongqwellsea receive very nearly the entire cyto- plasmic contents of the terminal cell of the sporophore. In species with branched sporophores there are, occasional- ly, larger than usual volumes of cytoplasm containing two (or three) nuclei which are isolated in a branch by its basal septum. The spore subsequently formed from this branch will be both markedly larger than normal and contain the-extra nucleus’ or nuclei. Such exceptionally rare L198 spores are clearly aberrant and cannot be used to invali- date the significance of uninucleate spores. Similarly, they do not justify the inclusion in Erynta (all of whose other species have uninucleate primary spores ) of #. caro- lintana (whose spores are always multinucleate and whose other major morphological characters do not match with the other species of Erynta). Three genera are noted by Batko (1964a-b, 1974) to have plurinucleate spores. The spores of several Entomoph- thora species (sensu Remaudiére and Keller 1980) contain 4-6 nuclei; those of £. eultets (Braun) Fres. are charac- teristically binucleate (or occasionally trinucleate) while those of E. muscae (Cohn) Fres. contain 5-8 nuclei. Trtplo- sportum (= Neozygttes) was described by Batko (1964b) to have quadrinucleate spores. Of the eight species included in Neozygttes by Remaudieére and Keller (1980), seven have 4-nucleate spores; Entomophthora turbtnata Kenneth (1977) has spores with 5-7 or more nuclei. This deviation from the quadrinucleate condition of all other species in this genus indicates that close scrutiny must be given to the generic assignment of this relatively little studied fun- gus. On first examination, the only marked similarities of E. turbinata with the other species of Triplosportum are its simple sporophores and ovoid resting spores with a jet black ‘epispore. Batko; (1974 )eincorrectly minds catedmsnae the primary spores of Massospora Peck are binucleate; in fact, they vary from 1-6 (or more) nuclei, but most common- ly contain 2-3 nucle (Sopers19/455 193i). The number of nuclei in spores of Cultetcola species (as originally described by Batko 1964c-d) varied from pluri- to multinucleate, but really represent a bimodal distribution with either a few nuclei or many in the pri- mary spores. This bimodal variation perfectly parallels the dispersal of Cultctcola species for other reasons to either Entomophthora s.str. (plurinucleate) or Contdtobolus (multinucleate) . The taxonomic importance of nuclear morphology has not been appreciated until now even though some authors such as Batko (1964a-b, 1966; Batko and Weiser 1965) have drawn at- tention to the staining of nuclei in lactophenol/cotton or aniline blue or other stains, or to their general size and appearance, but never realized their porential taxonomic value. The large nuclei of Bastdtobolus species, for exam- ple, have been the subjects of numerous cytological studies. MINS Humber (1981b) finds that nuclei of Contdiobolus spe- cies (sensu King 1976b) have relatively small nuclei with a prominent central nucleolus but no obvious heterochromatin; these nuclei remain unstained or only very weakly differen- tiated in lactophenol/aniline blue, aceto-orcein, or a nun- ber of other nuclear stains. The species of Entomophthora and most of the entomopathogenic genera have large nuclei with prominent heterochromatin but no large central nucleo- lus; these nuclei may be differentiated in lactophenol/ani- line blue and are strongly and rapidly differentiated in aceto-orcein and other nuclear stains. This karyological criterion has proven to be of inestimable value for diag- nostic work, particularly for distinguishing species of Contdtobolus from Entomophaga (whose primary spores are nearly identical in shape) or other genera with readily differentiated nuclei, and in determining incidences of double infections in insects when one of the fungi is a species of Contdiobolus. Taxonomic Significance of Primary Spores Batko's recognition of the generic significance of the number of nuclei (one, a few, or many) in primary spores has been widely embraced. The Remaudiére classification, however, necessarily disregarded this character in order to Poace thesgreatestygeneric value on the shapes (of primary, spores. As will be made more apparent below, this depen- dence upon spore shapes is misplaced and leads to what seem to be several misclassifications which offer no improvement over earlier artificial schemes based on spore shapes and proposed without taxonomic or phylogenetic significance. Nuclear cytology affords the only consistent and read- ily observed criteria which adequately distinguish the spe- cies of Contdiobolus from Entomophthora and its segregates (Humber 1981b). These criteria also uphold the validity of Entomophaga Batko for FE. grylit (Fres.) Batko and several other species. Entomophthora obscura Hall §& Dunn and £. thaxtertana (Petch) Hall §& Bell were assigned to Entomo- phaga (Batko 1964d) but are now regarded as synonyms (see below); nuclear cytology confirms the transfer of Ento- mophthora obscura to Contdiobolus by Remaudiére and Keller (1980). The unitunicate or bitunicate nature of primary spores appears to be perfectly correlated with the nuclear number. With the sole unconfirmed exception of EF. turbinata, all 200 known bitunicate spores also contain a single large nucle- us. Except for two species of Massospora in which uni- nucleate primary spores can occur (Soper 1974), all uni- tunicate spores are pluri- or multinucleate. It is possi- ble that this distinction in wall structures of primary spores may assume a greater significance as more entomoph- thoralean fungi are found and described. Among the characters related to primary spores in the Entomophthorales, then, generic value should be accorded to ae sleast the number and’ nature of the Nuclersitisthesspore: The shape and uni- or bitunicate nature of the primary spore should always be noted. However, while various states of these characters may coincide perfectly with the generic limits defined by other criteria, no reason is ap- parent at this time why either of them should be accorded generic value equal to nuclear number and morphology. PRIMARY SPOROPHORES Morphology and Branching The branching of sporophores was one of the first cri- teria used to divide the entomopathogenic Entomophthorales into two genera (Brefeld 1873, 1877; Nowakowski 1883), and has continued to be one of the leading characters used by Batko (1964a-c, 1974; Batko and Weiser 1965) to construct a contemporary classification. The historical significance of sporophore branching as a character with generic impor- tance demands careful review of both the reasons for and the effects of its de-emphasis by Remaudiére and Keller (1980). It is necessary first to examine the systematic distribution of simple and branched sporophores and then to evaluate the exceptions to the normal state in cach genus. For any practical purpose, the sporophores are always simple in Entomophthora (although in £. cultets several Sporophores may arise from each hyphal body, thus giving an impression of branching), Massospora, Tritplosporium, and Completorta Lohde, a parasite of fern prothallia. Tabano- myces Couch et al. (1979; from tabanid fly larvae) and Mertstacrum Drechsler (1940; from nematodes) produce an up- right, unbranched sporogenous hypha which becomes septate and produces forcibly discharged spores on lateral (or ter- minal) papillae. Ballocephala Drechsler (1951; from tardi- 201 grades) also produces an upright, unbranched sporogenous hypha on which small lateral cells are formed from which the spores are budded and forcibly discharged. Sporophores in species of Ancyltstes Pfitzer (Berdan 1938; from desmid algae) and Bastdtobolus (Drechsler 1964) may occasionally branch; those of the 27 Contdtobolus species recognized by King (1976b, 1977) also may, on rare occasion, be bi- or trifurcate (King and Humber 1981). Among the species transferred to Contdtobolus by Remaudiére and Keller (1980) the sporophores of Empusa aptculata Thaxter and EF. major Thaxter may show some branching on rare occasion (see Thax- COnmelooG wert Cf 500) Dut alesal Mos UsaiWayS*Sample.s aneTe .1s no significant branching in any Entomophaga species. All species of Zoophthora s.str. (Remaudiére and Hen- nebert 1980; Keller 1980; Ben-Ze'ev and Kenneth 1981la-b) have digitately branched sporophores with the possible ex- ception of Z. crassttuntcata Keller (1980) in which the sporophores appear in histological sections to be mostly simple.* All but three Erynta species (Remaudiére and Hen- nebert 1980; Remaudiére and Keller 1980; Kramer 1981) have digitately branched sporophores. Within the bounds of the usual biological variation, it is not surprising that some species with simple sporo- phores occasionally have a small number of sporophores that show a low order of branching. Whatever branching occurs in these genera is usually basal rather than apical as in Zoophthora and Erynta (where the apical, digitate branching pattern usually results in the formation of a tightly interwoven hymenium). The infrequent occurences of branched sporophores in genera characterized by simple sporophores or of occasional simple sporophores in genera characterized by branched sporophores in no way indicates that this character should be de-emphasized or discarded. The sporophores of Zoophthora and Erynta (apart from the three exceptions discussed below) do exhibit a variable degree of branching. Unbranched sporophores might occa- * Keller's (1980) characterization of the sporophores of Z. crassttuntcata as mostly simple seems to be based en- tirely on histological sections and should, therefore, be accepted as provisional until the publication of micrographs of carefully dissected pieces of hymenium showing whole sporophores of this species. 202 sionally be found on specimens from this large group of species in which nearly all sporophores are digitately branched. Similarly, species such as Z. crassttuntcata may exist in which (digitately?) branched sporophores are less numerous than simple ones, but the affinities of these species with Zoophthora or Erynta will always be apparent from the natures of those branched sporophores, their pri- mary spores, and other associated characters. The natural degree of plasticity in the branching of sporophores in species of Zoophthora and Erynta does not imply, however, any real probability of finding species such as Z. radtcans (Bref.) Batko (= Entomophthora sphaero- sperma Fres.) or Erynta neoaphtdts Rem. & Henneb. (= Ento- mophthora aphtdis Hoffman sensu Thaxter) which form obvi- ously digitate sporophores to yield an exceptional specimen in which a majority (much less all) of the sporophores are simple. It is even less feasible that a strain of such a Species exists which produces branched sporophores when in- fecting most hosts but which forms only simple ones on one or more other hosts. The “uniform occurence of digitatessporophnoresain Zoophthora sensu Remaudiére & Hennebert — including those (digitately?) branched sporophores of Z. crassttuntcata — leaves only the two species of Strongwellsea and EF. caroli- ntana, the three exceptional species placed in Erynta by Remaudiére and his colleagues, to account for the de-empha- sis by Remaudiére and Keller (1980) of sporophore branching aSpageenericrcr!lernion:,. It is important to examine why Remaudiére and Keller (1980) supposed that a significant probability exists for finding branched sporophores in the two Strongwellsea spe- cies. It is these fungi which seem to form the basis for their de-emphasis of sporophore branching. The statement that branched sporophores are "not always present" in Stronguellsea, their primary justification for their opin- ion, seems to draw solely upon Strong et al. (1960) who note that the sporophores are "rarely branched" but who also admit the difficulty of tracing complete structures in their prepared slides. Batko and Weiser (1965) used some of these slides and correctly described the sporophores of S. castrans Batko §& Weiser (1965) to be unbranched; they neither illustrated nor noted any exception to this simple State. Humber (1975, 1976) also found the sporophores of S. magna Humber to be always simple, and included this 203 characterization in the emended generic description of Strongquellsea. The uniformly unbranched nature of sporo- phores in this genus reflects the means by which additional sporogenous hyphae interpolate themselves into and thus continually expand the surface area of the fungal ball in the host fly's abdomen (Humber 1975). It has already been noted that #. caroltntana (with multinucleate and unitunicate spores, simple sporophores, and lack of rhizoids or cystidia) shares no major charac- teristics with Hrynta species except for spore shape; the SimnplessporophnoressOtetnisesnecies do notconstitute any Significant exception to the common state of digitately branched sporophores in Erynta. However, some minor degree of branching of £. caroltntana sporophores may occur t1nstde the host body, but the emergent sporophores are uniformly unbranched on the surface of affected craneflies (Thaxter 1888). Both Giard (1888, as EF. arrenoectona Giard) and Keller (1978) also find the sporophores of FE. carolintana to be simple. Several aspects of sporophore morphology — the pre- sence and pattern of branching and the overall shape of the sporogenous cell or sporphore apex — should be considered to be important, but generic significance has usually been placed only on whether a sporophore is simple or branched. The sporophores of all Entomophthora species (in the strict sense) are markedly swollen and clavate below the spore, but this is the only genus in which this character is uniformly Dresent a nero 1s58a ess consiscent tendency, to produce clavate sporophores in species of Zoophthora or Erynta. In most other species and genera of the order, the apical por- tion of the sporophore is relatively cylindrical with only a slight tendency to become clavate. Srinivasan et al. (1964) proposed to separate Contdto- bolus from Entomophthora on the basis of sporophore shape. They suggested that Contdtobolus species have micronemous (hypha-like, indeterminate) sporophores while those of Entomophthora are macronemous (thicker than and distinctly differentiated from the vegetative hyphae, and of determi- nate length). This criterion was devised to separate Contdtobolus species from Entomophthora muscae (the type of its genus) and paid little attention to the many other spe- cies in Entomophthora sensu lato. In the more modern taxo- nomic systems considered here, the criterion proposed by Srinivasan et al. (1964) is applicable only to the few spe- 204 cies of Entomophthora s.str. (Remaudiére and Keller 1980); it is also untenable, however, since Contdtobolus adtaere- tus Drechsler (see King and Humber 1981, Fig. 4f) and some other Contdtobolus species have markedly differentiated (macronemous) sporophores. Several genera can, in fact, be distinguished in part by the exact morphology of the sporophore. Bastdtobolus species have a more or less prominent swelling of the sporophore immediately below the spore, and the sporophore is markedly narrowed at the point where the spore is at- tached. Species of Meristacrum, Tabanomyces, and Ballo- cephala form several spores on each unbranched erect sporo- genous hypha. The mode of spore production in Mertstacrum and Tabanomiyjces 1S identical, and these genera are here considered to be synonymous (see below). The spores of Ballocephala are produced on globose to elongate lateral cells produced sympodially at the growing apex of the sporogenous hypha (Drechsler 1951; Richardson 1970; Pohlad and Bernard 1978). Three species placed in Contdtobolus by Remaudiére and Keller (1980) — &. apteulata, E. major, and E. paptllata Thaxter — differ from all other species of that genus {in the sense of King 1976b, 1977) by having an extended and neck-like narrowing of the sporophore apex; this morphological difference suggests that ally characters of these fungi should be compared carefully with those of. Contdtobolus before accepting their placement in this par- ticular genus. Taxonomic Significance o1° Primary vporopnores The presence of branched or simple sporophores was re- garded as a primary generic character in the Batko classi- fication but was de-emphasized (and effectively rejected) in the Remaudiére classification, seemingly to justify the synonymy of Strongwellsea (in which the sporophores are al- ways simple) with Hrynta (in which the usual case is for sporophores to be digitately branched). This de-emphasis and the elevation of spore morphology to a primary generic character by Remaudiére and Keller (1980) further confused the classification by placing £. caroltntana (with simple sporophores) in Erynta and Entomophthora carpenttert Giard (apparently with branched sporophores, see below) in Contdtobolus. As has been advocated by Ben-Ze'ev and Kenneth (1981a) sporophore branching should be re-instituted as a prime ZOS generic character. This restoration sweeps aside the ap- parent misclassifications of the Remaudiére scheme by re- turning Strongqwellsea to its rightful status as a separate genus (Humber 1975, 1976, 198la), and leaving the place- ments of £. caroltntana and EF. carpenttert undetermined andgineneed Of sturtner study. For the entomopathogenic genera and Contdtobolus, the most taxonomically significant aspect of primary sporo- PaOLVessISeODLyY whetheruchicy are simpte OF Drancned, There does seem to be a significant difference, however, between a low level of basal branching (as in Contdtobolus and E. earolintana) and the apically digitate branching restricted to the species of Zoophthora and Erynta. The means by which the primary spore or spores are produced on the sporophore and the morphology of the sporophore apex also assume taxonomic significance when considering the complete spectrum of entomophthoralean genera. MODE OF FORCIBLE DISCHARGE OF PRIMARY SPORES Three distinct mechanisms for the forcible discharge of primary spores are known from the Entomophthorales. Two Ouethesc mechanisms appear to be entirely restricted. to Single clearly defined genera while the predominant mecha- nism operates in nearly all the remaining genera. Primary spores are forcibly discharged in all generally recognized meonera Of this order except for Massosporda, in which spores and hyphal bodies are passively dispersed from the disinte- grating abdomens of the living host cicadas. All of the diverse types Of secondary spores are also forcibly dis- charged except for those capillispores (see below), produced on long, narrow capillary sporophores. All species of Entomophthora sensu stricto (Remaudiére and Keller 1980) have campanulate and apiculate primary spores which are forcibly discharged on a stream of cyto- plasm and vacuolar sap squirted from the sporophore as the wall layer(s) securing the primary spore to the sporophore Break. ine SpOreritsel: Nas a tliat or Sliehtly convex base rather than a distinct papilla, and appears to play no active role in the discharge process. Spores discharged in this manner are accompanied throughout their trajectories 206 by a voluminous drop of the protoplasm which serves to af- fix the spore to the surface on which it lands, and forms a characteristic radially striate corona around the pri- mary spore upon drying. The primary spores of all species of the non-entomo- genous genus Bastdtobolus are borne on sporophores with a pronounced swelling immediately below the primary spore. Discharge occurs with a circumscissile rupture of the lower portion of the swelling; the spore and swollen upper por- tion of the sporophore fly away as the hydrostatic pressure in the upper portion of the sporophore is released back- wards like a small rocket (Ingold 1934). Ingold's rocket analogy is strengthened by the fact that it is often possi- ble to obtain the firing of a "'second-stage" as -the small conical projection of the Bastdtobolus sporophore into the spore itself may evert during flight, thus giving a slight push against the sporophore fragment and assuring a some- what longer trajectory for the spore than for the sporo- phore fragment from which the spore becomes detached. With the exception of Massospora, all other genera of the Entomophthorales apparently have primary spores dis- charged by the eversion of the spore's papilla against the sporophore (Gallaud 1905; Couch 1939). This mechanism de- pends upon the establishment of high turgor pressures in both the sporophore and spore prior to the rounding off of the turgid cell (Ingold 1971). The sporophore wall breaks at the point of spore attachment, and the spore papilla everts rapidly from its original position pointed into the spore and pushes the spore away from the turgid sporophore. The shock of this eversion against the pressurized water column of the sporophore sometimes causes a rupture of the sporophore tip and exudation of a water droplet at the time of discharge (@€.g., Page and Humber 1973). Despite the lack of detailed understandings of these discharge mechanisms and of the involvements of the various wall layers of the sporophores and spores, it is apparent that spore discharge is a major event in the life history of an entomophthoralean fungus requiring the expression of a considerable portion of the genome. In view of the sig- nificance of this process for the fungus and the restricted distributions of the three diverse mechanisms, it is appro- priate to recognize the mode of forcible discharge of pri- mary spores as having taxonomic value at least on the generic level. 207 MODE OF SECONDARY SPORULATION The general ability of entomophthoralean fungi to produce one or more types of secondary spores from primary spores is one of the most unusual features of this group of fungi. Among other fungal taxa, the Sporobolomycetac- eae (heterobasidiomycetous "mirror"' yeasts), many ascomy- cetes whose ascospores may bud repeatedly in the ascus, and some other entomopathogens such as a few species of Septobastdtum Patouillard (Couch 1938) and the hyphomycet- ous genera Mutogone Thaxter (1920) and Termtartopsts Blackwell, Samson & Kimbrough (1980) may produce secondary spores. In none of these groups, however, has secondary sporulation assumed such a significant role as in the Entomophthorales. It is becoming increasingly apparent, for instance, that the secondary spore types are equally asmantrective -as primary spores or that primary, spores serve as dispersive units while the secondary spores pro- duced from them may even be the major infective units (Carner 1976; Carner and Canerday 1968; Kramer 1980; Nemoto and Aoki 1975; Nemoto et al. 1979; Selhime and Muma 1966; Tsintsadze and Vartapetov 1976; Wilding 1970). The morphological diversity (Batko 1974; King and Hum- Deve ISi)wandebiologicaimimportances of ythese;sporestypes suggests that they might have taxonomic value above the Specitrce eve li@um netactietwouty pes. ofesecondaryaspore forms — microspores and capillispores — have formed the Dasesmtonitaxonomic divisions at, Subgeneric or generic levels in the Entomophthorales. Microspores are produced when the primary spores of some Contdtobolus and Bastdtobolus species produce a few to several dozen small secondary spores which are forcibly discharged and act individually as asexual propagules. The formation of microspores by C. coronatus (Costantin) Batko was first used to distinguish the genus Delacrotxta Sacc. §& Sydow; Tyrrell and MacLeod (1972) proposed that Delacrotxta should be regarded as a subgenus of Contdtobolus to accom- modate all species of that genus capable of forming micro- spores. Capillispores (despite a profusion of names which are applied to them) are passively detached spores produced Singly atop a thin and elongate (capillary) sporophore. Batko (1966) proposed to divide Zoophthora among four sub- 208 genera wil all species producing almond-shaped to elongate secondary spores on capillary sporophores to be restricted to the subgenus Zoophthora Batko. Remaudiére and Hennebert (1980) restricted the genus Zoophthora to those species producing capillaries and capillispores and transferred all species remaining in the other three subgenera proposed by Batko (1966) to Erynta Nowakowski. In addition to their presence in all species of Zoophthora subg.Zoophthora sensu Ben-Ze'ev §& Kenneth (198la) or Zoophthora Batko sensu Remaudiére & Hennebert (1980), capillispores are produced by all species of Triplosporium sensu Batko (but not Weozy- gites sensu Remaudiére §& Keller 1980), Mertstaecrum (inclu- ding Tabanomyces; see below), and some species of Contdto- bolus and Bastdtobolus. Batko (1974) considers the pre- sence of capillary ‘sporophores® to reflect an ancestray character of the Entomophthorales and rejects the possibi- lity of thei r=separate orig insmine theses cenerds Dymeovolt- tionary convergence. If the Remaudiére classification were prefectly even- handed: in ats=adoption=of avgenericecri terionspasedson capillispores, then Entomophthora turbtnata Kenneth (1977), which produces no capillispores, should not have been in- cluded in Weozygttes by Remaudiére and Keller (1980), and both Contdtobolus and Bastdtobolus should have been split.. It is curious that Remaudiére and his colleagues apparently to not accept the parallel notion of splitting Contdtobolus because of the presence of microspores in some species of this genus. Ben-Zéetev and) Kenneth | (199la)* correctly rejects the generic emphasis placed on capillispores by Remaudiére and Hennebert (1980), and propose to use microspores and capil- lispores as a character with subgeneric validity as did Batko (1966) and Tyrrell and MacLeod (1972) before them. However, it is neither advisable nor even possible to apply criteria based on specialized secondary spore forms to all genera in which they might occur since Bastdtobolus mtcro- sporus forms both microspores and capillispores (Benjamin 19625 209 RHI ZOIDS HUStOn ca lebenspecuLvenands laxonomic, Dist nipuoien Batko's (1964a-c, 1974) placement of generic value on the presence of rhizoids has been one of the most trouble- some aspects of his classification of the entomopathogenic Entomophthorales. His adoption of this criterion narrowed his circumscription of Entomophthora s.str. and led him to disperse the few species now regarded to belong in this genus (Remaudiére and Keller 1980) among two genera. This misplaced emphasis also led him to propose Cultetcola and Entomophaga, separated by the respective presence or ab- sence of rhizoids, genera whose validity and circumscrip- Clonsmhavemremainedasuspect, duemtonthe: general controversy OVermthiesvabidity, of rhazor1dseand) LOmbatkO Seinabii Ety to provide a definitive separation of Contdtobolus from Ento- mophthora sensu lato. Humbersetealmei(1977)mere-evaluated and'rej;ected) the Significance of rhizoids described for Entomophthora vtru- lenta Hall §& Dunn (1957; = Conidtobolus thrombotdes Drech- sler; see Latgé et al. 1980). The rhizoids described for this fungus are now regarded to have belonged to another entomophthoralean fungus affecting the same aphids but which produced no spores or whose spores were undetected. Remaudiére and Keller (1980) and King and Humber (1981) independently discuss those species which usually form rhizoids but occasionally fail to do so. Note that elmmeunese eXCeNtLONnsarepresents cine: absenccuOb (Ore tarlure SOutindjmrii zoids in species normally producing «them; they never involve observations of rhizoids in species not known to form them. Remaudiére and Keller (1980) believe that no Weiehtasnouldsbpesputeonsthe presence of rhizo1ds because PNeitiepTes ence mane cercainespecices. gis NOtwalwaysecOnstanun On the contrary, King and Humber (1981) regard the presence of rhizoids to be taxonomically significant while their ab- sence is not a dependable criterion.* Ben-Ze'ev and Kenneth (198la) regard the presence of rhizoids to be taxonomically Significant at the subgeneric level. Brobyn and Wilding (1977), in the most thorough consideration of the develop- Brera toi. a lechanvesmotsthestex uniintventronallyaattered this#opinion tows tate sthateche absence: of rhizoids us not taxonomically significant. 210 ment of rhizoids and cystidia yet published, uphold the presence of rhizoids to be a taxonomically significant character. It is necessary to consider the general’ situation to know if any confidence should be placed on the presence (or absence) of rhizoids as a taxonomic characters. Even though no absolute scale is available to quantify one's level of confidence, it is necessary to weigh how well a species is known: How many times has it been found? In how many sites? From how many hosts? Are rhizoids always present or always absent? An examination of the taxonomic distribution of rhizoids among the entomophthoralean spe- cies reveals that many have never been found with them present. In only a few species, particularly Zoophthora radtcans (= Entomophthora sphaerosperma), is the presence of rhizoids usual but inconstant. The absence of rhizoids might be explained in some instances by improper handling of specimens, but it is also possible that their formation On some insects might be’ a function of that particular host (Remaudiére and Keller 1980; King and Humber 1981). GENERA WITHOUT RHIZOIDS. Rhizoids are completely ab- sent or have not been found in several genera. “These in- clude Trtplosportum (at least 7 species), Entomophaga (at - least 4 species although the circumscriptiom of this genus remains in dispute), Tabanomyces (1 species; = Mertstacrum), Massospora (13 species), and Strongwellsea (2 species). None of the non-entomopathogenic genera — Bastdtobolus, Aneyltstes, Completorta, Mertstacrum, and Ballocephala — produce rhizoids. ENTOMOPHTHORA (SENSU STRICTO). Two of the six species of Entomophthora s.str. (Remaudiére and Keller 1980) pro- duce rhizoids. This is the least constant appearance of this structure in any entomophthoralean genus. It can be Said with assurance that rhizoids do not occur in the type species, HE. muscae, since this is the most frequently ob- served of all species in the order. Both #. cultets (Braun) Fres. and £. planehontana Cornu produce abundant rhizoids on hosts bearing forcibly discharged primary spores, but these anchoring structures may not occur when resting spores are formed. Both species are cosmopolitan in dis- tribution and affect a wide variety of dipterans and aphids (respectively). All of the remaining species — EF. webert Lakon ex Samson (a little known species from a neuropteran), E. thrtptdum Samson §& al. (1979; known only from thrips in Dutch greenhouses), and #. erupta (Dustan) Hall (which is known from several North American and European sites and hosts) produce no rhizoids and disperse their primary spores from living, mobile hosts. The same behavior occurs in the species of Massospora and Stronguellsea. CONIDIOBOLUS. Among the 27 morphologically homogene- OUSmSDCCLCSMLeCOUnIZeadmDY Kinge 10 76a-b,607 /)eenone as known to produce rhizoids; very few Contdtobolus species are currently known to be entomopathogenic. Remaudiére and Keller (1980) transferred most entomophthoroid species with round to pyriform spores from Entomophaga (which was cir- cumscribed to exclude species with rhizoids) and those spe- cies from Cultcicola with round spores to Contdtobolus. All other Cultctecola species have campanulate spores and were restored to Entomophthora by Remaudiére and Keller (1980). Among the 38 Contdtobolus species listed by Remaudiére and Keller (1980), four produce rhizoids: Empusa aptculata ax arena jormanax wake spapt lara. Thax..and LnLomopn— thora carpentiert Giard (1888). The first three species were placed in Cultctcola by Batko (1964c-d) because of enelwerhi Zo dseandmultanucleate round spores borne on ysim- ple (or occasionally branched) sporophores. Their pro- nounced similarity to Contdtobolus species belies two obvi- ous morphological differences from all other Contdtobolus Species: the consistent formation of a few stout rhizoids ending in strongly differentiated holdfasts, and the nearly cylindrical, collar-like narrowing of the sporophore apex. Close study of these species will be required to verify if they actually belong in Contdtobolus as suspected by King (1976b) and affirmed by Remaudiére and Keller (1980). The only morphological character of Entomophthora car- penttert Giard (1888) fitting Contdtobolus is the round primary spore supposed by Turian (1957) to belong to this Species. Several other characters of this fungus — the presence ‘of rhizoids,iminucleate spores, on branched sporo= phores (Turian 1957), and cystidia (Petch 1944, if Petch's belief that EF. coleopterorwn Petch and #. carpentteri are identical is correct) — correspond exactly with those of Erynta or Zoophthora in the Remaudiére classification. Entomophthora coleopterorum (Petch 1944) has narrowly oval rather than round primary spores; the mode of rhizoidal at- tachment is identical with that of F. carpentteri, a spe- Ale cies which Giard (1888) characterized only the the distinc- tive nature of its rhizoidal.attachment. In a situation resembling that described above for the "rhizoids" of C. thrombotdes, one cannot discard the possibility that Turian (1957) may have observed the round spores of a secondarily invasive Contdtobolus species on a beetle which was already infected by F. carpenttert which was able to form its very characteriStic rhizoids but nogspores.. |NOicytolovical staining or photomicrographic evidence supported Turian's contention that these round spores were uninucleate; it is possible that he mistook a,central oil droplet or vacuole for-arnucleus. The discrepancies between the, Grardseretch, and Turian concepts of #. carpenttert can be reconciled only after examining any existing herbarium material of these collections. In any event, however, the entire habit of this species refers it to Erynta or Zoophthora rather than to Contdtobolus. ZOOPHTHORA.- The presence oF eri zZOLds sUSsest Tone) ear. related with branched, sporophores bearing yuninuclcate ww. p1— tunicate primary spores — the species of Zoophthora Batko or Erynta Nowakowski. All eight Zoophthora species recog- nized by Remaudiére and Hennebert (1980) form rhizoids. By including four newly described species, 4. lanceolata Kel- ler (1980), Z. crassttunteata Keller (1980), Z. ortentalts Ben-Ze'ev & Kenneth (198la), and Z. petcht1 Ben-Ze'ev & Kenneth (19381b),. thestotalarisesstorat: leastmulmoteslmspc « cies known to produce rhizoids. Ben-Ze'ev and Kenneth (1938ia) remain uncertaim af rhizoids were: presen aonmcue few specimens of Z. ortentalts available to them. ERYNIA. Among the species listed by Remaudiére and Hennebert (1980) and Remaudiére and Keller (1980), some 21 of 25 Erynta species (excluding those of Strongwellsea) produce rhizoids. Together with Hrynta delphacts (Hori) Humber*, £. delpintana (Cavara) Humber**, and EF. tthacensts Kramer (1981), the total incidence of rhizoids rises to 22 of 28 species. The absence of rhizoids in the six Erynta - ERYNIA DELPHACIS (Hori) Humber, comb. nov., basionym: Entomophthora delphacis Hori, 1906, Entomol. Mag. (Tokyo) 8: 81. ** HRYNIA DELPINIANA (Cavara) Humber, comb. nov., basionym: Entomophthora delpiniana Cavara, 1899, Nuov. Giorn. Bot. Ita lei Ser aaa? 2 ZS Species discussed below is not considered to overturn the value of rhizoids as a secondary generic character although E. delphacts and EF. delptntana constitute the most notable Ormtiicsceexcep tions. inexteme( less suspected tnaGrhazoids were ipresent on Empusa virescens Thax., but could not be certain since he was not the original collector of the material which he PA CCTHUSCU mt OPCGSCrIventiismspecies: Erynta aquattca (Anderson § Anagnostakis) Humber* is known only from mosquito larvae and pupae floating on the surface of temporary pools in the northeastern United states. / This floating habit of the stricken hosts main= tains their position for favorable transmission of the fun- gus withouueany » need for rhizo1ds. /alhis species is mot known well enough, however, to say whether rhizoids might nOCMrOMN, for cxampley on =individuals trapped atethe receding edges of rapidly drying pools. Ben-Ze'ev and Kenneth (1979) report cystidia but no rhizoids from Zoophthora ertnacea Ben-Ze'ev §& Kenneth, but note the Similarity of rhizoids and cystidia of species referable to Zoophthora subg. Erynta Batko (1966). They muse on the possible inter-relatedness of cystidia and rhizoids as did Thaxter (1888) and Gustafsson (1965). However, Brobyn and Wilding (1977) provide effective evi- dence that the functions and course of development of these Serle cures sacemtunnelatreduinuspectes wiere sthey mothimoecur together; it must be assumed, then, that rhizoids and eystidiasare under completely Separate genetic controls. Empusa carolintana, with its multinucleate unitunicate Spores borne on simple sporophores and lack of cystidia or rhizoids, is once again shown to stand apart from Erynia in * HERYNIA AQUATICA (Anderson & Ringo ex Anderson & Anagno- stakis) Humber, comb. nov., basionym: Entomophthora aquatica Anderson §& Ringo ex Anderson & Anagnostakis, 1980, Mycotaxon 70, 350, NON Entomophthora aquattca Anderson G Ringo, 1969, J: Invertebr. Pathol%, 73, 386 (which was invalidly published without a designated type). The combination Hrynta aquatteca (Anderson & Ringo) Remaudiére & Hennebert, 1980, Mycotaxon 7/7, 30 Pan smal SOmitiva Licdwaccordinget OmiriLcles4>"ousthe International Code of Botanical Nomenclature. 214 all major characters except for the shape of its primary spores. The correct generic assignment for this fungus re- mains problematic, although its spore karyology, sporophore morphology, and mode of spore discharge more closely re- semble the species of Entomophaga than any other genus re- cognized by the Batko, Remaudiére, or Ben-Ze'ev/Kenneth Clas sutivcat tons. Erynta delpintana and Erynta delphacts may be true exceptions to the usually rhizoidal state of fungi in this genus. Cavara (1899b) is fastidious in his observations of the small, uninucleate, bitunicate primaryespores., isecon— dary apores like the primaries, digitately branched sporo- phores, and very large and occasionally branched cystidia of FE. delpintana. However, he never specifically mentions any hyphae affixing the affected flies to the damp piers of a bridge over a small stream. The flies were completely covered under a dense, dirty-white to straw-colored hyme- nium. This entire hymenial covering was easily broken away from the cadavers during attempts to detach the insects (Cavara 1899b). Except for the apparent absence of rhizoids all characters of this species matchjthe scharacteristics used by describe Zoophthora subg. Erynta Batko (1966); all species in this subgenus have rhizoids and prominent cysti- dia. Erynta delptntana is known only from Cavara's origi- nal collections, but it seems likely that examination of this material or any fresh collections would reveal the Presence ofarh@zords: Erynta delphacts is a much more important exception to the presence of mhizoi1ds thant thevotherespectesmconsiccred above. This pathogen of leaf- and planthoppers in Asian rice paddies bears a strong morphological resemblance to EZ. neoaphtdis Remaudiére § Hennebert (= Entomophthora apht- dts Hoffman sensu Thaxter) from aphids (Shimazu 1976, 1977; Remaudiére and Hennebert 1980), but differs notably in the complete lack of rhizoids on any of its hosts, including two artificially infected aphid species, Aphts gossyptt and Macrostphum akebtae (Shimazu 1977, and personal commu- Nication). #rynta delphacis produces cystidia both on its host and in culture. Even though Hori (1906) illustrates and describes spherical, yellow-white resting spores 24-28 um in diameter and containing many fatty granules, no such spores have been found in subsequent collections of this Species in Japan, the Philippines, or Indonesia, nor are they formed in culture. pases) Remaudiére and Hennebert (1980) consider the differ- ences of growth tm vitro between £. delphacts and E. neo- aphtdts to be insignificant and regard #. delphacts as a nomen dubtum. However, a comparison of a Japanese strain (Shimazu F32; our RS 134) and several Philippine isolates of E. delphacts (RS 458, RS 459, RS 461, RS 478, RS 479) from green leafhoppers, Nephotetttx virescens, with numer- our cultures of £. neoaphtdits isolated by Insect Pathology Research Unit personnel indicate that #. delphacts grows substantially more rapidly on a wider variety of media (nutritionally simple or complex), and sporulates a great dealmoresprotiitically, overrva Vongerstime sthenvany ‘culture of E. neoaphtdts we have yet observed. Until more exten- Sive studies can be made comparing these two species, it appears to be preferable to recognize H. delphacts as a legitimate species of Erynia. It should also be noted that if later studies indicate the synonymy of EF. delphacts and E. neoaphtdts, the International Code of Botanical Nomen- clature would require that £. delphacts (Hori) Humber must be the correct name for the fungus still generally known as Entomophthora aphtdts Hoffm. (sensu Thaxter). Much more pertinent information is needed about the morphology of rhizoids than the perfunctory characteriza- tion found in many descriptions that rhizoids are merely “present. \-)sAs noted by “King and Humber (1981) 3 complete descriptive information about rhizoids should include their points of emergence, dimensions (particularly thickness), abundance (few or many?), relative spatial distribution (isolated, aggregated into one or more pseudorhizomorphs, or forming a veil?), and the morphology of any terminal holdfast apparatus. Thaxter (1888), Gustafsson (1965), Ben-Ze'ev and Uziel (1979), and Ben-Ze'ev and Kenneth (198la-b) are exemplary in the clarity of their illustra- tions and characterizations of rhizoidal morphology. This descriptive information can be useful for detecting dual infections involving rhizoidal and non-rhizoidal species Since Brobyn and Wilding (1977) affirm that the morpholo- Dress OferiilzO1ds) appear to be: speciecsuspecific. At least three morphological classes of rhizoids can be recognized: Most rhizoidal species have either (1) numerous individual rhizoids which are relatively little differentiated from vegetative and may or may not have any terminal holdfast differentiation, (2) numerous hypha-like rhizoids aggregated into one or more columnar pseudorhizo- morphs, or (3) relatively few, thick rhizoids with strongly ALO differentiated (usually discoid) terminal holdfasts. No comprehensive study of the development and morphology of rhizoids or of their taxonomic distribution has yet been undertaken. The thorough studies of Brobyn and Wilding (1977) were limited to a few species affecting aphids. In view of the observations presented here, however, such a comprehensive study would be exceptionally useful. Rhirzoids as a Taxonomicy Character The Batko classification adopted the widely accepted contention of Nowakowski (1883) that the presence of rhi- zoids is a major generic character. This criterion, super- imposed on those of nuclear number in primary spores and the branching of sporophores had effects which have been regarded as unacceptable and have prevented the general acceptance of the Batko classification (Remaudiére and Keller 1980; King and Humber 1981). The Remaudiére classi- ficatiLonse ine turn. rejected ithetsien7 fi Cancemos at hespre— sence of rhizoids. It seem certain, however, that the absence of rhizoids in many genera is significant while their presence in others is an important secondary charac- cour Regardless of the level (if any) at which one may ac- cept the taxonomic importance -of rhizoidsy species present— ing exceptions to the normal rhizoidal or nonrhizoidal con- dition of their genus should be examined carefully. The classifications of Strongwellsea castrans, S. magna, and E. earolintana in Erynta and of FE: aptculata, EH. major, E. paptllata, and FE. carpentterit in Contdiobolus are important examples of possible misclassifications pointed out by the lack or possession’ of *rhizoids, CYSTIDTA Cystidia (pseudocystidia, paraphyses, or pseudopara- physes for other authors) are short to long, spear-like to columnar structures of determinate growth that project above the level of the sporogenous hymenium. Until recent- ly, little has been known of their function, but Brobyn and Wilding (1977) convincingly demonstrated that the cystidia of Hrynta neoaphtdis are penetration organs which rupture the host cuticle and aid the subsequent emergence of the PANG SPOropnoOres. mene emer gencesOtecystidlda betore the sporo- phores also has been noted in £. ertnacea (Kenneth 1977) and £. tthacensts (Kramer 1981). It seems likely that CVStldlaweservestospertorace tnesnostectticle prior to ny— menial formation in all species in which they occur; future investigators should be careful to note the order of emer- gence of cystidia and sporophores. Remaudiére and Keller (1980) indicate that, apart from Erynta and (less commonly) Zoophthora, the only other genus in which cystidia occur is Contdtobolus. Among the 38 spe- cies of Contdtobolus accepted by Remaudiére and Keller, the only ones ever mentioned to form cystidia are C. obscurus, C. thrombotdes (= EF. vitrulenta), and EF. carpenttert (as E. coleopterorum Petch 1944); as was noted above, the latter species almost certainly belongs in Hrynta or Zoophthora rather than Contdtobolus. The description of EF. virulenta (= C. thrombotdes; see Latge et al. 1980) states that “¢ystidia occur rarely” (Hall and Dunn 1957). However, as explained above for the mrhiuzords Mor this specresy no ssupsequent treatment Of 1¢ TevULComiiname~cl (nereriuzOLds@eOr CYStidias mine CyYSstt. dia" of C. thrombotdes might have been those of another entomophthoralean fungus which produced both rhizoids and cystidia (see Humber et al. 1977). Hall and Dunn (1957) might also have interpreted vegetative or sporogenous hyphae whose indeterminate growth occasionally far exceeds enewveneraleicevel, Ore tie nymen1im LONbe cySticia,. Remaudiére et al. (1979) report the formation of long cystidia by C. obscurus on aphid cadavers placed in a humid environment. I have observed similar growth of long hyphae above the hymenium of aphids affected by C. thrombotdes but always find these structures to be undifferentiated vegeta- tive hyphae which should not be regarded=as cystidia by the definition given above. The listing of Lamta (= Entomophthora) cultcts in a chart by Nowakowski (1883) as having cystidia present and rhizoids absent was corrected by Batko (1964c) as being a typographical transposition not corresponding with the Pol- Ponerextawit CirmcOLrectly mnOteSmcii Ss sSpeClesico NaVvemrnii— Zo1ds DUE NO cystidia. The genus most closely related to Zoophthora and Erynta must be Stronguellsea, with its uninucleate and bi- 218 tunicate primary spores borne on simple rather than branched sporophores (Batko and Weiser 1965; Humber 1976). Remaudiére and Keller (1980) consider the lack of rhizoids and cystidia to be insignificant criteria and use this in- formation to explain their submergence of Stronguellsea into Erynta (see Humber 198la). In fact, if cystidia func- tion only for cuticular penetration, it is not surprising that Strongwellsea species form no cystidia. These fungi produce no hymenial structures outside the host's body. The abdominal hole (the only cuticular rupture caused by the fungus) begins with a point perforation of the pleuron and enlarges progressively. This point initiation of the rupture appears to be involved with the hypertrophic reac- tion of the host cuticle which leads, in turn, to the ex- tensive lining of the lower portion of the fungal ball by the host's cuticle. It seems clear that some mechanism completely distinct from that of cystidial penetration is involved in the formation by Strongwellsea of an abdominal hole (see Humber 1975, 1976). In view of the above discussion, it srs probably jcor- rect to say that the only species in which cystidia (as determinate organs for cuticular penetration) occur are those with uninucleate, bitunicate primary spores borne on branched sporophores and classifiable in Erynta or (less commonly) Zoophthora sensu Remaudiére §& Hennebert. Unlikesforerhazoids,. thespresencesorgabcencomOtecy oti. dia has never been used to separate genera although the Batko classificaiton recognized their value as a supporting character strongly correlated with the occurence of uninuc- leate, bitunicate primary spores and branched sporophores. ASawith the other charactersediseussedshere, sexceptlousaco the usual state of thescharactersinga cenuseshouldgindicave thateae close sinspecctonsormt ne cOLTespOnucncesmO Leda lst characters of the suspect species with those of other spe- ciespin they genusuiseinvorder wealnesp resencemOcacystldl amin E. coleopterorum (Petch 1944), a presumptive synonym of Contdtobolus carpenttert (Giard) Remaudiére § Keller, of- fers a significant example of this principle. 219 RESTING SPORES aygospores or Azygospores: A Morphological/Cytological Dilemma I applaud the de-emphasis by Remaudiére and Keller (1980) of the distinction between zygospores and azygo- Spores. sineyenotercorrectlys that resting Spores are not known (or are not formed) in a series of species, and that the mode of their formation has not been observed in a Parsee propored Onsotaspecics sctia badQsproducemsres ti ngEspores, Both types of development may occur in closely related spe- cies or even within a single species (MacLeod and MUuller- Kogler 1973; Humber 1976; Remaudiere and Keller 1980; King and Humber 1981). This developmental distinction has been used only twice as a taxonomic character of generic importance. The peculiar mode of conjugation of (amoeboid?) hyphal bodies or gametangia of Empusa fresentt Nowakowski 1883 (= Neozy- gttes aphtdis Witlaczil 1885) and the resultant jet-black, ovoid zygospores formed at the point of conjugation prompt- ed Ihaxter (1888) to set this and related:species apart iin Empusa subg. Tritplosportum. This taxon was later raised to generic status by Batko (1964b), who was apparently unaware of the nomenclatural priority of the generic name Neozy- gttes Witlaczil (see Humber et al. 1981). King (1976b) notes that the double-walled resting spores formed by the species of Contdtobolus are anisogamous zygospores. Other types of conidioboloid resting spores such as chlamydo- spores and villose spores are distinguished from zygospores (or azygospores) by their single rather than double wall SCRUCTULeS. Couch (1939) notes that the zygospores of Contdtobolus species are formed in the larger of two gametangia (and are almost always produced in the axis of the parental hypha) while those of Entomophthora (sensu lato) are produced in lateral buds from the conjugating hyphae or hyphal bodies. Very few species of Entomophthora s.1. produce their zygo- spores in the hyphal axis: FE. vtrulenta (= C. thrombotdes), E. obscura Hall & Dunn [= £. tgnobtlts Hall §& Dunn = #£. thaxtertana (Petch) Hall §& Bell; see below], and Empusa diptertgena Thaxter. The former two species are now recog- nized as species of Contdtobolus by Remaudiére and Keller (1980). With the apparent exception of £. dipterigena, 220 whose zygosporogenesis is illustrated by Gustafsson (1965), the resting spores of other species of Entomophthora or its segregated are budded off from the parental hypha or hyphal body. Zygospores, the bilayered, thick-walled spores derived from a conjugation of two gametangia, have been a well established phenomenon in the Entomophthorales since their first recognition (Brefeld 1973; Nowakowski 1877, 1882, 1883; Thaxter (1888). Nearly every subsequent major discus- sion of these fungi has referred to zygospores as sexual spores and to azygospores as asexual spores (Schroter 1897; MacLeod 1956, 1963; Gustafsson 1965; Waterhouse 1973). In general morphological terms, zygosporogenesis may be called a sexual process, in which case azygosporogene- Sis must be regarded as asexual. However, in their strict sense, the terms zygospore and azygospore refer only to the mode of development of a thick-walled spore in the Zygomy- cotima (Fitzpatrick 1950; Bessey #1950 Ainsworth ol: Snell vand Dicks 19713" AlexopoulosmandsMims se! 979)e. There is a hazard in’ thinking sotm@zycosporeseas sexual and azygospores as asexual: One can be misled easily into performing an act of semantic sleight-of-hand by then ex-. pecting that the genette events of a sexual life cycle, karyogamy and meiosis, necessarily occur in a ''sexual" zygospores but cannot occur’ in an “asexual” azygosporey A related terminological confusion can occur when one refers to the resting spore as the sexual spore in the entomoph- thoralean life “cyclesasvopposedmto sthestorcib lyedrscharced primary -(or*secondary) @spores which ane, anseveryeasense- asexual spores. Nuclear events in entomophthoralean resting spores — reductions in nuclear number, karyogamy, and meiosis — remain completely uninvestigated in all but a few species. This gap in our knowledge reflects a lack of fundamental Studies on the life cycles’ in’ this group. Some cytological studies have noted the progressive reductions of nuclear numbers to two or even a single nucleus in fully mature resting spores (Vuillemin 1900a-b; Krenner 1961; Latgé 1976; Couch et al. 1979; D. McCabe and B. Martinell, unpublished). Other investigators using different species have found no obvious change in the nuclear number from the time of formation to the fully TOE mature, dormant state (Olive 1906; Riddle 1906; Goldstein 1929; Humber 1975). If the Entomophthorales follow pat- terns of karyogamy and meiosis similar to those in other Zygomycetes (Cutter 1942a-b), karyogamy should occur in the resting spore followed by meiosis in the spore just before or during its germination. No conclusive cytological evi- dence of nuclear fusions or of meiotic division has yet been found in entomophthoralean fungi despite the strong circumstantial evidence for’ their occurences (e.g¢., Couch tera dee 97.98): It must be hoped that synaptonemal com- plexes, a widely recognized ultrastructural marker for chromosomal synapsis, will be demonstrated at the time and ST LCeOMeticapresumpulVve mclOsi se inmsoncsOrsmoresmelners, of the Entomophthorales. Data regarding changes in nuclear numbers presented by Latgé (1976) support the possibility that karyogamy and meiosis occur in the resting spores of C. thrombotdes, and tiatecnesvevetatiVesnuclei, OL thisespecies must pe haploid, Albeit, Latgé feels that his data are also compatible with gametangial meioses of diploid nuclei such as occurs in the Oomycetes (see Alexopoulos and Mims 1979). Little ambigu- Peyeaboucethc interpretation OL tice! lilcecyCcie.remains, however, in view of the results of electrophoretic analyses of isozyme patterns of several strains of C. thrombotdes and Z. raditcans (May et al. 1979) and of EF. muscae, Masso- spora etcadina Peck, and Entomophaga gryllt (Fres.) Batko (Soper, May,@and*@Martanell; invpreparation): These results Suggest that the vegetative nuclei of these species are haploid, although heterokaryosis may occur in some species Si ncemmuluipleralleles;otrsome loci may, bespresent. inva mycelium. There is no evidence yet of heterothallism within the Entomophthorales. All zygosporogenesis appears to be homo- thalliceandmleavess opens atileaststwo possiblesinterpreta= tions torstheo life cycles of these: fungi: Those species whose zygospores or azygospores at some point contain only a single nucleus probably have autogamous (sexual) life cycles. Those species whose zygospores or azygospores undergo no reduction in nuclear numbers may have abandoned a sexual life cycle in favor of an apogamous (asexual) life which preserves the particular (successful) combination of characters in that species. No evidence now exists that parasexuality occurs in any species of the Entomophthor- ales. Zee The distinction between zygospores and azygospores cannot be retained as a character of taxonomic signifi- cance although the presence and nature of zygospores is a useful ancillary character in at least Triplosportum and Contdtobolus. The developmental origins of resting spores can be difficult to interpret correctly even itesuitapie material is available. Nuclear events in resting spores are of far greater importance for the fungus than whether conjugations precede sporogenesis. Karyogamy and meiosis, the critical genetic events in a sexual life cycle, appar- ently may occur in either zygospores or azygospores; similarly, other species may have apogamous life cycles regardless of whether their resting spores are zygospores or azygospores. Indeed, the full details of entomophthoralean life cycles and their breeding strategies may yet prove to be useful taxonomic characters, but not enough is currently known of these processes to draw meaningful conclusions. There is no doubt, however, that this information is of absolutely vital importance for the practical development and manipulation of these fungi for use in the microbial CONtTLO MMOL Sethe reinsectanostse Resting Spores: Problems and Interpretation of Taxonomic Traps It appears that the generic events of sexual repro- duction might occur in either zygo- or azygospores, and that even species producing ''sexual"' zygospores might be apogamous. Several further caveats and an example serve to reinforce the rejection of any taxonomic value which might be attached to the distinction between zygospores and azygospores (Remaudiére and Keller 1980). In most instances, authors simply state their obser- vation that the resting spores of a given species are zygospores or azygospores and neither illustrate nor dis- cuss the evidence supporting this conclusion. However, Sawyer (1931) notes the great difficulty in determining if conjugations occur in Z. radtcans. Humber (1975) also notes difficulties with the resting spores of Stronguellsea Species due to problems of interpreting the morphology of some hyphal bodies and to the evanescence of the emptied hyphal wall remnants which provide the only temporary evi- dence for the mode of formation of mature resting spores; determining the 'pedigree™ of these resting spores is dependent upon seeing them during a relatively narrow Periodsuuting their development. Consequently, 1t seems unwise to place full confidence in any characterization of resting spores as zygo- or azygospores unless convincing documentation is also presented. The difficulty in correctly applying these developmen- tal designations casts doubt on whether any taxonomic de- cisions should be based solely on this distinction (even though it may haye some utility for characterizing, but not Separating, some genera as noted above). Ihe practical ef- ECCUS Ompditrering ViewpOINtseorstice ValuceOLt thts distinc- tion can be seen in the following example: In studies on the identities of aphid-pathogenic Species of the Entomophthorales, Humber (1978) noted that Entomophthora tgnobtits Hall & Dunn should be used as the nomenclaturally valid name for Entomophthora thaxtertana (eeccijelial wambelipeaespecleselouwhich Hal leand Bell (1963) considered F. tgnobtlts to be identical. Empusa thaxtertana Petch (1938) was proposed for a species mis- identified by Thaxter (1888), and which was characterized as having azygospores. The resting spores of E£. tgnobtits are described by Hall and Dunn (1957) as zygospores. Hum- ber (1978) considered the reported differences in the ori- gins of resting spores to be so ambiguous as to be insigni- ficant, an opinion which is reinforced here. A later-study by Remaudiére et al. (1979) used morphological and bio- chemical data to demonstrate the synonymy of EF. thaxtert- ana and EF. obscura Hall §& Dunn (1957) [= Contdtobolus ob- seurus (Hall & Dunn) Rem. & Keller], which was described as producing azygospores. In fact, the differences between EF. thaxtertana, E. tgnobtlts, and E. obscura had always been troublesome for diagnosticians. The only major described difference be- tween #. tgnobtlts and HF. obscura is that of zygospores versus azygospores, a difference considered here, by Humber (1978), and by Remaudiére and Keller (1980) to be insigni- ficant. A more consistent and simple approach to this taxonomic problem would have been to accept EF. obscura to Hestieecorrectundice DyaVvilTtucrOL pacenpriOrityeoveras. tgnobilts rather than to regard the latter species as a nomen dubtum merely because of the described difference in the, ortegins of the resting spores in these two species (Remaudiére et al. 1979). 224 If there is to be even-handed application of the accepted taxonomic criteria and acknowledgement of the net effects of rejecting other criteria which have been applied in the past, it iS now necessary to recognize Unescorrect synonymy of Contdtobolus obscurus to be the following: Conidiobolus obseurus (Hall & Dunn) Remaudiére § Keller 1980, Mycotaxon 17, 331. = Entomophthora obscura Hall.& Dunn 1957, Hil- GAraGHae: 7,902. = Entomophaga obscura (Hall & Dunn) Batko 1964, Bulle gAcad. Polons Cia ocr OCummD Lois 12, 404. Entomophthora tgnobtlts Hall & Dunn 1957, Hil- SardiauaieeloZ. Entomophthora planchontana Cornu sensu Thaxter 1888, Mem. Boston Soc. Nat. Hist. 4, 165, NON Sensu. Petch 1948 y9hrans bri teeny COlemoOG. Pil a, = Empusa thaxtertana Petch 1938, Trans. Brit. Mycol. Soc. 21, 34. NAME INVALID WITHOUT LATIN DIAGNOSIS. [= Entomophthora thaxtertana (Petch) Hall § Bell 196355 “Jesinsect Pathol. worm Usoe| [= Entomophaga thaxtertana (Petch) Batko 1964 Bulle Acade RPalon!aeSer soci aabLOl amie 404. ] CHOOSING A CORRECT CLASSIFICATION: TRIAL BY ANOMALY One of the most severe tests for a classification SystemersetOusce NOWaner tm GecalanalGl oeticulcast suyl teal species from the group being classified. Withanethe, Ento- mophthorales, several seemingly anomalous generic assign- ments from both the Batko and Remaudiére classifications are discussed here. Howeverj;iin terms of testing the real capacities of these or any other classifications of these fungi, confirming the correlations of characters, or sug- gesting where primary taxonomic weights should be placed, Dab the most important fungus has not yet been considered. PitSetseareunuescriped=species tound Dy sbadazy 31975) ein Poland which was, obviously, not known at the time of the proposition of the Batko classification, and was not con- sidered by the Remaudiére classification. THiswrungusmathixcceolatsmiliptera.=lendipeataae) aby rhizoids to wet, decayed wood just above a stream surface in a deciduous forest in the Wielkopolski National Park, Poland. Its primary spores are Slender and curved, 45-75 (82) x 8-10.5 um, strongly tapered apically and with a frat cencdmm conical spapi | laemeeinesesciaraclers: a1mostsexace- ly match those of Erynta contca (Thax.) Rem. §& Henn., which RaSmcucCndetEClLVCdusPOLCSmlOLedmsUZCE LOCH El Ca ml eDatazy) Ss fungus) with bitunicate walls and single nuclei, borne on branched sporophores, and with cystidia and rhizoids pre- Sent. .batazy's fungus) 1s, however, clearly different from E. contea since its primary spores have unitunicate walls, contain 4-10 nuclei, and are borne on simple sporophores ; cystidia are absent. The Remaudiére classification, with its primary empha- sis on spore morphology, would place this species in Erynta even though that assignment would be impossible if one ac- cepts the emended circumscription advocated here (excluding Stronquellsea and EL. caroltntana from the genus by limiting ite COUSPEC1ES With uninucleate, bitunicate spores: borne .on branched sporophores and not producing capillispores). The Batazy fungus might be regarded as a species of Cultctcola except that Batko (1974) apparently rejected Cultetcola as a heterogeneous mixture of species now regarded to belong to Entomophthora s.str. or Contdiobolus (Remaudiére and Keller 1980; King and Humber 1981). There is no basis for including the Batazy fungus in Contdtobolus according to Phescircumscripiionssot ethicmeenusecithenestated ors implied by Batko (1974), Remaudiére and Keller (1980), here, or in Humber (1981b). It might be assumed that the Polish fungus could be allied to EF. aptculata, EF. major, and E. paptllata becauscuoL their multinucleate uni tunicatesspores wexcepe Pnatetneanuclearsmorphotogysotethe slat tersspeciesscorre- sponds to that of Contdtobolus species while that of the Polish fungus resembles Entomophthora and all of its segre- gates rather than Contdtobolus (see Humber 1981b). The generic system outlined by Ben-Ze'ev and Kenneth (1981la) would include this species in Entomophthora sensu stricto, but the mode of spore discharge in the Batazy fungus is by Ciceeversi Oneotethe papilla: 226 If, as suggested here, one rejects spore morphology as thei primarysgeneric criterion and accepts the restriction of Erynta noted above, and places generic value upon the karyology of primary spores, the morphology of the sporo- phore, and the mode of spore discharge, the only remaining sensible disposition for Batazy's fungus is in a new and (for now) monotypic genus allied to but differing from Erynta. It seems reasonable to assume that other entomoph- thoralean species may yet be found with characters like those of the one characterized by Batazy (1978). It is imperative that this Polish fungus be fully characterized and more widely publicized. This fungus seems certain to provoke much useful discussion of the taxonomic criteria which should be accepted for entomoph- thoralean fungi, and of how much weight should be accorded ComeacheOLathescacricenlar MODERN VIEWS OF ENTOMOPATHOGENIC GENERA OF THE ENTOMOPHTHORALES This study has examined the major morphological cri- teria used in entomophthoralean taxonomy, particularly as they are applied in the generic classification schemes pro- posed by Batko and by Remaudiére and his colleagues. It is apparent from all of the above considerations that the application of only three characters provides an effective separation of species into morphologically and biologically homogeneous genera. These characters are the following: 1. The number and nature of nuclei in the primary spores. Zo Ines morpho logysobathessporophores (and the mode of sporogenesis). 3. The mode of discharge of primary spores. Neither the morphology of primary spores, types of secondary spore formed, presence or absence of rhizoids, presence or absence of cystidia, nor the mode of formation of resting spores was found here to be important for de- limiting genera in the Entomophthorales. Each of these Vogt characters, however, may have utility as secondary or sup- porting characters which do not in themselves determine the PimVceSwOL sscicra = so oOme OL these Characters arevappropriate for the circumscription of subgeneric taxa (Ben-Ze'ev and Kenneth 198la). It is appropriate to summarize the difference among the Batko and Remaudiére classifications and that proposed preliminarily by Ben-Ze'ev and Kenneth (198la), and to note those adjustments which seem to be necessary to circun- Scribe these genera still more accurately according to the Gr iecTiasaccep ted ene: ENTOMOPHTHORA Fresenius. Batko's overestimation of the importance of rhizoids caused him to split a small and natural group of species (MacLeod et al. 1976; Samson et al. 1979) with simple sporophores, campanulate apiculate primary spores containing a relatively small number of nu- clei, and a characteristic mechanism of spore discharge among two genera. The application of the mechanism of spore discharge as a generic criterion confirms the narrow definition of Entomophthora proposed by Remaudiére and Kel- ler (1980) despite the de-emphasis here of the value of spore morphology. The exceptionally high degree of morpho- logical and biological homogeneity of this group of species would be manifestly lowered if the mode of spore discharge did not exclude both Empusa caroltntana Thaxter and the species found by Batazy (1978) and discussed above. Both of these species have simple sporophores and plurinucleate primary spores but are discharged by papillar eversion rather than by the "fungal cannon" mechanism of #. muscae. MASSOSPORA Peck. This venerable genus which affects gregarious cicadas (Homoptera: Cicadidae) was not treated in the Batko classification until his phylogenetic treat- ment (Batko 1974) since Massospora represents the only genus of the entomogenous Entomophthorales whose validity and circumscription (see Soper 1974, 1981) has never been questioned. The species of this genus have plurinucleate primary spores which are not forcibly discharged from the Simple sporophores lining small cavities in the abdomen of affected insects. TRIPLOSPORIUM (Thaxter) Batko, nom. gen. conserv. prop. sBatkow(1964b) taccepted Whaxter"s (18838)) belief ithat E. fresentt and similar species should be accorded generic Status. This small and natural group of species is. distin- 228 guished primarily by its unique mode of zygosporogenesis and ovoid zygospores with black (or very dark) epispores. Batko (1964b) circumscribed the genus to include species whose primary spores are quadrinucleate, borne on simple sporophores, and capable of producing capillispores as one type of secondary spore. Remaudiére and Keller (1980) ac- cept Batko's circumscription of this genus but they (1) reject the requirement for the primary spores to be quadri- nucleate in order to justify the (contentious) inclusion of E. turbtnata Kenneth (1977), and (2) replace Trtplosportum (Thaxter) Batko with the older generic name Weozygttes Wit- laezil (1885) Humber ct als (i98l))" otter severaiereasons why Trtplosportum should be conserved against the older and nomenclaturally correct name Neozygites; this nomenclatural issue will be resolved at the 13th International Botanical Congress during the summer of 1981. ENTOMOPHAGA Batko. This genus, based on Entomophthora grylit Fres., was proposed to include species wtih multi- nucleate spores and simple sporophores but without forming rhizoids. Its validity has been questioned because of its linkage to the unacceptable Cultetcola through Batko's emphasis on rhizoids and also because no satisfactory cri- terion has been proposed to distinguish unambiguously be- tween any of these species and the morpholgoically similar Contdtobolus. The lack of this criterion seemingly prompt- ed Remaudiére and Keller (1980) to emphasize spore morpho- logy by transferring all species of Entomophaga and those of Cultetcola with round rather than campanulate priamry spores to Contdtobolus.. However, the criterion of nuclear morphology noted above and discussed fully by Humber (1981b) readily delimits Contdtobolus from Entomophthora sensu Stricto and its segregate genera, and confirms the validity of Entomophaga for species with multinucleate unitunicate primary spores containing large, readily stainable nuclei, and discharged by papillar eversion from simple sporophores. ZOOPHTHORA Batko and ERYNIA Nowakowski. Batko (1964b) described Zoophthora, with Entomophthora radtcans Brefeld (= Entomophthora sphaerosperma Fresenius) as its type, for all species having uninucleate, bitunicate spores borne on branched sporophores and producing rhizoids. This genus has been accepted as a natural and valid grouping. Four subgenera were proposed by Batko (1966) based on the morphology of sporophores, rhizoids, and cystidia, and the type of secondary spores produced. Remaudiére and Henne- bert (1980) limited Zoophthora to species capable of form- 229 ing anadhesive secondary spores atop capillary sporophores (see King and Humber 1981), with all other Zoophthora spe- cies being transferred to Erynta Nowakowski (1881). The Remaudiére classification modified the sense of Batko's grouping to accommodate species with simple sporophores and even multinucleate unitunicate primary spores so long as the spore shape was similar to that in other Erynta SDSCLES For purposes of discussing the Remaudiére classifica- Cionmsanesthisepaper, at has been necessary to accept this generic separation on at least a provisional basis. Con- siderable objection can be raised, as noted above, about ehesuserotecaprl larvesporophores ease ar charactemsom generic Send ol Cancen mais sSuemoasmDecns treated pant by ,Ben~ Ze'ev and Kenneth (198la) and will be discussed more fully Dymtnese@autnlors@ineassubsequcnts paper. A nomenclatural problem reminiscent of the simultane- ous use of Empusa Cohn and Entomophthora Fresenius as taxo- nomically distinct genera has been raised by Remaudiére and Hennebert's use of both Zoophthora Batko and Erynta Nowak- owski. In 1881, Nowakowski proposed the genus Lrynta to include Entomophthora ovtspora Nowakowski (1877) and Ento- mophthora curvispora Nowakowski (1877), but later rejected this generic name in favor of Brefeld's (1877) usage of Entomophthora as taxonomically distinct from Zmpusa (Nowak- owski 1882, 1883). At the time of its description, Zooph- thora Batko (1964b) included only its type species, Ento- mophthora raditcans Brefeld. However, the inclusion in Zoophthora of FE. ovtspora and FE. curvtspora (Batko 1964d) automatically required the adoption of the earlier name. Batko (1966) was incorrect in believing that Nowakowski's later disuse of Erynta removed that name from consideration in matters of nomenclatural priority. The ultimate resolution of this nomenclatural problem with Erynta and Zoophthora depends on the outcome of the debate to establish a concensus opinion of whether the Bat- _ko or Remaudiére classification, or some modification of one of them shall be accepted for the species with bituni- cate, uninucleate spores on branched sporophores. If, on the one hand, two genera are recognized with one based on Entomophthora ovtspora, the type species chosen for Erynta by Remaudiére and Hennebert (1980), and the other on Ento- mophthora radtcans, there is no nomenclatural problem to resolve. If, on the other hand, common practice rejects 230 any generic separation and recognizes only a single genus which includes both £. ovtspora and E. radtcans, then either Hrynta would have to be adopted as the correct gene- ric name or Zoophthora might be proposed for conservation against Hrynta. It is unfortunate that such a confusing nomenclatural issue should arise once more in the Entomophthorales, but the three possible resolutions are unambiguous. My person- alsopinaon should tbe 'cleareirom this spapey. me lecOsilouEpe- lieve that the species originally classified by Batko in Zoophthora should be split between two genera separated by characters such as spore morphology and the formation of capillary sporophores. As much as it might prevent some amount of further confusion in the literature if Zoophthora Batko were conserved against Hrynta Nowakowski, I do not believe that such a proposal would be accepted for incorpo- ration in the lists of nomina conservanda in the Interna- tional Code of Botanical Nomenclature. The great prepon- derance of species proposed as or transferred to Zoophthora have already been transferred to Erynta by Remaudiére and Hennebert (1980) or here; the relative number of species required to be transferred as a consequence of conservation of a younger name against an older and nomenclaturally cor- rect one appears to be one of the major concerns of the nomenclatural committees which decide these matters. The new combinations in Hrynta proposed here are fully justi- fied and nomenclaturally correct whether one accepts the taxonomy of Batko (and his followers) or or Remaudiére and Hennebert for classityinosthe species sinsdquesuion= STRONGWELLSEA Batko § Weiser emend. Humber. Humber (1976) provided an emended generic description and valida- tion of this genus whose spores are uninucleate and bituni- cate but are borne on simple rather than branched sporo- phores, and are discharged by papillar eversion. Remau- diére and Keller (1980) provided no effective rationale for rejecting the importance of sporophore morphology or other diverse supporting data (Humber 1976) in order to synonymize Strongwellsea with Erynta. There can be little doubt from the criteria considered here to be significant at the generic level that Strongqwuellsea must be recognized aS vamseparate genus.) -Asmore extensive; support stores enls Opinion is presented by Humber (198la). Zor CULICICOLA Nieuwland. This genus is nomenclaturally superfluous since its type species, Entomophthora cultects (Braun) Fres., belongs in Entomophthora sensu stricto (Remaudiére and Keller 1980; King and Humber 1981). The species included in this genus by Batko (1964c-d) are now dispersed to Entomophthora s.str. or Contdtobolus Brefeld by Remaudiére and Keller, although some question remains whether the conidioboloid species producing rhizoids — E. aptculata, E. major, and FE. paptilata (which was not classified by Batko, 1964d, but clearly belonged in this group) — should be in Contdtobolus or in a closely allied but different genus. Batko's inclusion in Cultctcola of E. vtrulenta (= C. thrombotdes; see Latgé et al. 1980) was based on the erroneous description of this species as pro- ducing rhizoids (Humber et al. 1977). Batko regarded this as the most tentative of his pro- posed segregate genera, and he later apparently rejected Culteteola in his extended justification of his taxonomic ideas and discussion of phylogeny in the Entomophthorales (Batko 1974). Cultetcola has been the most objectionable of hisegenera tor other studients, of these«fungi1. CONIDIOBOLUS Brefeld. Even though Batko and Weiser (1965) note that resting spores of Contdtobolus are not budded off to the side as in all other genera considered in the Batko classification, they again cited the widely used but untenable "'criterion'' which supposes Contdtobolus spe- SLeSetoubeabasicallyesaprobesawhide, those) of the other genera treated were entomopathogenic. Remaudiére and Kel- ler (1980) regard the lack of a suitable criterion to de- limit Contdtobolus from morphologically similar species put in Cultetcola and Entomophaga by Batko (1964b-d), with round to pyriform, multinucleate primary spores and simple spores, to justify their inclusion in a broadly redefined Contdtobolus. As indicated here, however, nuclear cytology does pro- vide the criterion sought to delimit Contdtobolus species from those of Entomophthora and its segregates (Humber 1981b). Some of the species transferred to Contdtobolus from Entomophthora sensu lato truly are species of Contdto- bolus; these include £. virulenta (= C. thrombotdes; Latgé et al. 1980), C. obscurus, and possibly C. aptculatus, C. majon,,and.C. papiilatus (although these last three’ species may belong in a separate genus allied to Conidtobolus due to marked differences in the structure of the sporophore Boz with the presence of rhizoids as a supporting character). Other species — e@.g., EF. gryllt, FE. batkot, and EF. gigan- tea Keller — have nuclei whose morphology indicates their closer affinities to Entomophthora than to Contdtobolus (Humber 1981b). TABANOMYCES Couch, Andreeva, Laird, & Nolan. This little known genus (Couch et al. 1979) was described for the peculiar mode of germination and spore formation from entomophthoralean resting spores found in larvae of tabanid flies in the Soviet Union.; The thick-walled; coloriess, ovoid resting spores of the type species, T. milkot (Dudka & Koval) Couch §& al., were originally described as Coelomo- myces mtlkot Dudka & Koval. During the germination of the spores, the single nucleus undergoes two’ (meiotic?) “divi- Sions, and a short, thick, quadrinucleate sporophore forms. This hyphasbecomes septate eandecachwuninuctecates cel o- duces and discharges a lateral primary spore which is, in turn, capable of producing a secondary spore on a short Capillary =sporophore (Couchmet 417] 970) neseuevenecmalic: structures are essentially identical to those described for the germination of resting spores in the nematophagous genus Mertstacrum Drechsler (Davidson and Barron 1973). ThesnucleiallustratedebpyeCouchectealem Gloy ljmarce col ative> ly small and have a prominent central nucleolus; this type of nucleus resembles that of Contdtobolus (Humber 1981b) and Mertstacrum (Humber, unpublished). Remaudiére and Keller (1980) do not mention Tabano- myces, but this genus cannot be easily placed in any entomo- genous genus included in the Batko or Remaudiére classifi- cations. The remarkable similarity of Tabanomyces and Mertstacrum indicates that these genera are synonymous and that 7. milkot must be recognized as a heretofore unknown entomopathogenic species of Mertstacrum*. TARICHIUM Cohn. This genus was created for those spe- cies of the Entomophthorales known only by their resting spores (see MacLeod and Muller-Kogler 1970). Neither the * MERISTACRUM MILKOI (Dudka §& Koval) Humber, comb. nov., basionym: Coelomomyces mtlkot Dudka §& Koval in Dudka, Koval § Andreeva, 1973, Novitates Systematicae Plant- arum non Vasculartum 10, 88-91. SYNONYM: Tabanomyces mtlkot (Dudka § Koval) Couch, Andreeva, Laird § Nolan, 197975 Proce Natle Acadn ocr (USA) «cre 2090-500 255 Batko nor Remaudiére classifications fully accept this genus, but properly regard its members as having still un- determined affinities. It can be assumed that Tartchtum Species may be connected to a species producing primary spores and, thus, placed in the appropriate modern genera. A taxonomic system which accepts a narrow sense of knto- mophthora and a number of other entomopathogenic segregate genera must also accept Tartchtum as the correct provision- aieCeneric nane stOrethe specres involved ratner than the name Entomophthora (Tartchtum) as was suggested (without any formal nomenclatural status as a subgenus of Entomoph- thora) by MacLeod and Muller-Kogler (1970). Concluding Remarks on Generic Classifications Despite its manifest shortcomings, the classification proposed by Batko (1964a-e, 1966; Batko and Weiser 1965) forms the basis for all other contemporary approaches to a generic classification for the Entomophthorales. This classification is discussed in depth (although not substan- Cia tbyeanp iit ed )ymeitimecicurirstatcempt CO OUGLine) Ee course of evolution in the Entomophthorales (Batko 1974). The Remaudiére classification attempted to eliminate the flaws which Remaudiére and his colleagues perceived to pre- VentecncmcciCTraleaccepuancesO1sthespackO Classit) Catton amen generic arrangement more similar to that proposed by Batko than by Remaudiére, Hennebert, and Keller emerges here in view of the acceptance of primary spore karyology, sporo- phore morphology, and the mode of primary spore discharge as the three characters best suited for delimiting genera. Ben-Ze'ev and Kenneth (198la, 1981b, and other papers in preparation) also accept the karyology of primary spores and morphology of sporophores as characters with generic importance, but do not recognize the mechanism of spore discharge to be taxonomically significant. No formal proposal of yet another new generic classi- fication, seems appropriate at this time. This study has sought only to evaluate the characters and criteria which have been or might be used to construct generic classifi- cations of the Entomophthorales. I have extended the con- clusions of this study regarding appropriate generic cri- teria to the existing classification schemes in order to show their net effects and to stimulate further discussion and debate. 234 Each attempt to propose a new generic classification for the Entomophthorales has suffered from the effects of one or more faultily circumscribed genera and from the less obvious effects of incompleteness. It is clear from the Still incomplete catalogue of species described as or at- tributable to Entomophthora (sensu lato) (MacLeod 1963; MacLeod and Muller-Kogler 1970, 1973; MacLeod et al. 1976), that the available information about a distressing number of species is inadequate to allow a reliable classification in the narrowly defined genera comprising the contemporary classification schemes. I must also be noted that all of these latter-day efforts to reclassify the Entomophthorales have concentrated almost exclusively upon the entomopatho- genic genera: Several morphologically diverse genera are known only as saprobes or pathogens of fern gametophytes, desmide algae, tardigrades, and nematodes; at least three species of this order are also known to affect humans and other vertebrates (King 1979). These fungi are no less valid members of the Entomophthorales than those species altackinesinSects*+ 1 tSminappropMlatemtOaroUceapOU Gustine choice of criteria used to define genera or about the cor- rectness of one or another classification of the Entomoph- thorales without taking these non-entomogenous genera into full account. Despite these impediments to the proposition of a con- prehensive generic scheme, however, I believe that most concerned parties now agree that following a period of fur- thered1scussiony of taxonomi Cucti tenia wel tewit EpempoSssiL le to propose a generally acceptable, realistic, and phylo- genetically based classification for the Entomophthorales in which each genus will have a uniformly high degree of morphological and biological homogeneity. Despite the obvious difficulties in proposing such a comprehensive scheme, some comfort should be found in the thought that IF the generic scheme which is yet to emerge is accurate, then no further major adjustments of generic circumscriptions should be necessary. As new taxa are found and described, and as older taxa being held in abeyance become better known, they will be classified in the existing stable gene- ric structure or seen to differ in enough significant char- acters to warrant the erection of new genera. ZOD ACKNOWLEDGEMENTS I wish to express my profound gratitude to I. Ben- ZeMey elem Nem COUCI Rem Geek eCnnetis ahem) a khOrts ands G. Remaudiere for valuable correspondence and discussion of the taxonomic and nomenclatural issues raised here, and to IvBen-Ze'ev Jab. Kramers. DoE. McCabe; and Ra) S-tSoper FOMMUNCImecn tl calerevicwSeotetneemanuscrEpe. REFERENCES CITED AINSWORTH, G.C. 1961. Atnsworth & Bisby's Dictionary of the Fungt, Sth ed. 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The later has been found again: descriptions of a non stratose specimen from Africa and of a two-layered spe- cimen from Guadalupe are given. An abstract in English inclu- ding a detailed key to species is given in the end of the pa- per. INTRODUCTION Dans le cadre d'une étude des spécimens du genre Hymeno- chaete récoltés principalement en Afrique par J. BOIDIN d'une part et G. GILLES d'autre part, nous avons décrit précédem- ment une espéce, H. spathulata, remarquable par ses spinules a sommet spatulé (LEGER, 1980) ainsi que deux espéces, H. 4e- panabihis et H. hatpago (LEGER, 1981) a spinules pourvues de diverticules. Le présent travail a trait aux espéces dont 1'hyménium posséde des éléments stériles ornés de digitations. Jusqu'ici deux espéces présentent ce caractére: H. pinnatifida Burt et H, digitata Burt. H. pinnatifida a été retrouvé en République Centrafricaine, au Gabon et a la Guadeloupe. Une description compléte est donnée qui fait apparaitre notamment que cette espéce peut étre bistratifiée. A titre comparatif, l'étude de He OLO. 01d estereprisesebecompletce aupartim dustypesaesnUR.. Enfin, deux espéces sont décrites : H. acanthophysata nov.sp. a hyphes acanthophysoides et H. hauetslevii rov.sp. a éléments hyméniens de type pseudoacanthophyses. 242 HYMENOCHAETE PINNATIFIDA Burt ANN ee MESSOULT RbOU. sGard wnt ro Do eam iO Basidiome résupiné, étalé, adhérent, mince, 60-150- (200) pm, en petites taches plus ou moins confluentes, brun jaunatre (Munsell 1OYR 5/4 ou encore Saccardo's umber selon Ridgway) puis brun rouillé (Munsell 5YR 4/6) quand il s'éta- le plus largement, 4 marge fibrilleuse appliquée, 0,5 mn, chamois (Munsell l1OYR 7/8). Cortex (Cx, fig.1) épais de 5 4 15 um, sombre, formé d'un enchevétrement serré d'hyphes brun-rouge foncé, plus ou moins cimentées et collapsées, x 2-2,5-(3) pm, 4 paroi ép- aisse. Contexte (C, fie) assez elache, 50 a5 100simadsepaise seur, dimitique. Hyphes génératrices hyalines, x 1,5-2 pm, a paroi mince, ramifiées et cloisonnées. Hyphes squeletti- ques brun-jaunatre, x 2-2,5 pm, a paroi épaisse, non cloi- sonnées, exceptionnellement ramifiées. SpinwLessbrun) Ponce.) laspLupactal on j0mues 5. mum. non émergentes; certaines, plus rares, mesurent 40-50 x 6- 7 wm et émergent jusqu'a 20 pm. Toutes ces spinules nais- sent d'une couche sous-hyméniale sombre. Hyménium (H, fig.1) de 15 a 20 pm; rares basidioles et trés rares basides mures, 15 x 3 pm, a 4 stérigmates de 3 pm. Dendrophyses jaunes trés abondantes, longues de 12 a 22 pm et de diamétre variant de 2 4 6 pm, a4 nombreuses ramifications tres etroites (O25 pm) en petites touffes terminales pour la plupart. Trés rares hyphes paraphysoides gréles grimpant le long de certaines grandes spinules. Assez nombreuses masses cristallines volumineuses (souvent de 10- 20 x 10 pm) a4 la base de 1'hyménium. Sous-hyménium (sH, fig.1) formant une zone sombre de 20 pm environ, d'aspect identique au cortex, constituée d' hyphes plus ou moins cimentées et collapsées, x 2-3 pm, 4 paroi épaisse. Spores (sur sporée) cylindriques étroites, trés légére- ment déprimées, 3,8-4,5-(5) x 1,6-2 pm, hyalines, uninucléees (Giemsa), non amyloides, blanches en masse. ~ Description effectuée 4 partir de la récolte LY 5584, La Ma- boké, République Centrafricaine, 31 mars 1965, leg. J. Boi- din (Herb. Boidin). AUTRES RECOLTES LY 5931 sur Carapa procera, La Maboké, République Centra- fricaine, 16 septembre 1967, leg. J. Boidin. 243 (a> ¢ oP - i; Gans iy One Or aaah AP + a Bo. Se Gis Hymenochaete pinnatifida Burt (LY 5584 ) 244 LY 6041 sur support indéterminé, Boubakiti, République Centrafricaine, 27 septembre 1967, leg. J. Boidin. LY 9260 sur support indéterminé, Libreville, Gabon, 20 novembre 1978, leg. G. Gilles. ESPECES DE REFERENCES EXAMINEES H. pinnatifida Burt, New Smyrna, Florida, january 1897, CrCl iloydee SUeGlype) pete 2140. ICED rn H. pinnatifida Burt, ad Ligna emortua angtospernae in paroecta Aurelia, Algiers, 17 september 1971, leg. M.T. Dunn, det. A.L. Welden (Fungi Ludovictant ex herbarto Tula- neav/ Gb5) eevee oe LE: GAS PARTECULIER* DE LA RECOLTE EY%s091] Ce spécimen, trouvé par J. BOIDIN le 29 septembre 1976 4 la Guadeloupe se présente comme un H. pinnatifida bistra- ti BLes Chien). Tout se passe comme si un deuxiéme basidiome complet avec un cortex Cx2, un contexte C2, une zone sous-hyméniale sH2 et un hyménium H2 (pratiquement stérile) etait né d'un ba- sidiome précédent constitué (comme décrit pour LY 5584) d' un hyménium Hl], d'un sous-hyménium sHl, d'un contexte Cl et d'un cortex basal Cxl. Dans le dessin de la figure 2, la ba- se du champignon —- c'est-d-dire le cortex Cxl ainsi que les 4/5 du contexte Cl - n'est pas représentée. Le basidiome a. une €paisseur totale de 150 a 230 pm dont 100 pm pour le contexte Cl (alors que le contexte C2 n'a qu'une quinzaine de pm d'épaisseur). DISCUSSION : Les spécimens africains présentent une trés bonne concor- dance avec le type d' H. pinnatif{ida Burt examiné ainsi qu' avec la récolte américaine LY 6723 déterminée par A.L. WEL- DEN. Cependant, deux remarques s'imposent - L'assez grande variabilité de couleur des basidiomes, non seulement d'une récolte a4 l'autre mais aussi a4 l'inté- rieur d'une méme récolte. Nous distinguons trois groupes selon la couleur: gris rosatre (light cinnamon drab, vers 2,5 YR 6/2) pour LY 9260 et pour le centre de quelques taches de LY 6/233) brun; rougeatre (reddish brown, vers: sv bu) se ae le ou 4/6) pour LY 6041, LY 8091, LY 6723 pro parte et LY 5584 pro parte; brun jaunatre (yellowish brown, vers l10YR 5/4) pour LY 5584 pro parte (petites taches trés jeunes) et LY 5931. Cette couleur variable s'explique au moins en partie par le développement relatif de 1l'hyménium: ainsi, les taches de LY 6723 ot 1'hyménium est bien développé et fertile sont gris rosatre alors que les taches brun rougeatre ne présen- Vane i SO age ph BC Yt a iss ae siny Big ees ee hy br Baa) ice te ber ae of bes 3 en ie hy 7 sp is is a Bee , fi. SIBRY 25° 3 Y Axes By =| ) Jz zB u co Co Goce B eas » = x Nee oo ei “, Hate, Few, e* -n.@ A er. * ) car < ae Way & “i eas oe v Eee oy ‘Saas ae se . Bin. ae 2) “ . Aes a SITs iy x 3: ae Ue A. i / A a i Fl AQ Figure 2: partie supérieure d Hymeno- chaete pinnatifida Burt (LY 8091) 246 -tent qu'un hyménium faiblement développé, pratiquement sté- rile et constitué presque exclusivement de dendrophyses. De méme LY 9260, dont 1l'hyménium fertile est bien développé est uniformément gris rosatre. La marge, chamois a rouillé, a presque toujours une couleur plus vive que le reste de 1"hy- ménium. - Concernant la stratification, 1l'examen de la récolte LY 8091 conduit incontestablement 4H. pinnatifida Burt. Cette bistratification n'a jamais été signalée ni par E.A. BURT, ni ultérieurement lors d'autres récoltes. En résumé, H. ptnnati4ida Burt est une espéce bien carac- térisée par ses dendrophyses (a nombreuses et fines ramifi- cations)issues d'une zone sous-hyméniale sombre, par un contexte dimitique assez lache et un cortex basal sombre. La couleur de l'hyménium peut varier de facon sensible suivant 1'état de maturité tandis que la marge est toujours de cou- plus vive. L'aire de dispersion de H. pinnatif{ida s'étend non seulement a l'Afrique mais également 4 la Guadeloupe dont le spécimen montre un caractére jamais observé jusqu'ici pour cette espé- ce: une bistratification nette. Il serait intéressant de re- chercher aux Etats-Unis notamment, une présence éventuelle de stratification lors de futures recoltes. HYMENOCHAETE DIGITATA Burt Ann Missouri Bot a) Gards>= O47 OLS Basidiome résupiné, largement étalé, adhérent, épais de 700 a 800 pm, brun havane (Brussels brown 4 antique bro- wn, 7,5YR 4/4 a 5/6), 4 surface granulée-tuberculée, 4 marge trés mince, évanescente, concolore. Cortex (Cxl, fig.4) d'une quinzaine de pm, formé d'un enchevétrement serré et brun rouge sombre d'hyphes de 2,5-3 pm de diamétre, plus ou moins cimentées et collapsées, 4 pa- rol épaisse. Ce cortex se prolonge par un feutrage assez lache constituant le revétement piléique (tomentum T), d'environ 40 ym d'épais- seur. Contexte (fig. 3 et 4) stratifié, composé d'une alter- nance d'une quinzaine de couches sombres avec un nombre égal de couches plus claires. Les couches sombres (appelées Cx2 4a Cx8 et dont seules Cx7 et Cx8 sont représentées sur la figure 3) ont une teinte brun rouge foncé et présentent une consti- tution identique 4 celle du cortex basal Cxl. Les couches plus claires ont une double constitution: d'une part, les Ex i= EE ISA SY Be TR) EE qe NAHB) \ LV Be 4 1 AB r Guy Riek #0 ee os an. eg eS at hg se & G Cl wee soca 2 Ly 4 ; ass Rt Posie AID os O ( ee | PY t) 4H po ek T/ UO ALD § pa \) NY LOE UAE eH \G Q Nad LS WAT \ 4a (VY aaulnassp LR aed H7 Cx7 Figure 3: partie supérieure d’ Hymenochaete digitata Burt (Type) 248 Figure 4: base d* Hymenochaete digitata Burt (Type) hyméniums antérieurs (nommés Hl] a H7 et dont seuls H1,H/,H8 sont représentés sur les figures 3 et 4) sont bruns, formés d'une dense palissade d'hyphes de 2-3 pm de diamétre, 4 pa- roi épaissie, a articles relativement courts (7 4 10 pm), trés collapsées, auxquelles se mélent des dendrophyses 4 paroi épaissie, entiérement jaun&tres. D'autre part, au dessus de chaque hyménium, des zones trés claires sont formées d'un enchevétrement trés lache d'hyphes cloisonnées et ramifiées, de 2-3 pm de diamétre, a4 paroi mince et d'hyphes non cloi- sonnées, 2-3 pm de diamétre, a paroi épaissie (dimitisme). A la base de chaque hyménium, de nombreuses masses cristal- lines plus ou moins volumineuses. Spinules nées pour la plupart des couches trés som- bres; elles sont de deux types: trés nombreuses petites 249 spinules 20-30 x 2-3 pm, non émergentes. et grandes spinules 50-75 x 6-7 pm, moins abondantes et émergentes jusqu'a 40 pm. Hyménium (H8, fig.3) de 25 pm environ; quelques basi- dioles mais basides extrémement rares (non vues par BURT), [jexe2.5— et wa 4epetitsesteriematesades!)5enm.) Leessentiel est constitué par des sortes de pseudoacanthophyses nombreu- ses: ce sont des éléments hauts de 12 a 17 pm, 4a base ren- flée (3,5-4,5 pm) hyaline et a paroi mince, tandis que 1'ex- trémité rétrécie, jaunatre et 4 paroi épaisse porte de nom- breuses ramifications en tous sens. Certaines sont cependant déja entiérement jaunatres et a4 paroi épaissie comme celles des hyméniums antérieurs. Spores non observées. Description d'aprés le type de H. digitata Burt, El Boquete, Chiridut.ePananave Won seMaxOn «moo eC PH DISCUSSION : L'étude du type de BURT a permis d'apporter quelques pré- cisions sur cette espéce qui n'a pas fait l'objet,depuis le travail originel, d'étude détaillée. Le point le plus inté- ressant, outre la présence non encore signalée d'un tomentum et la constitution précise des différentes couches de strati- fication, est sans doute l'aspect détaillé des pseudoacantho- physes qui n'avaient été que superficiellement décrites par BURT. HYMENOCHAETE ACANTHOPHYSATA Léger nov.sp. Basidioma resupinatum, Late jacens, adherens, tenue, obscure braunneum, magis tn medio propter prutnam grtiseokum; ambi tu ex nrubiginosos fulvo, margine angustissimo, albo; trama bir- Strata, dimtica, hyphis genetricibus hyalinis, tunica tenut, x 2-3 pm, ramosis fibulatisque, ex hyphis skeletticis e fla- vis brunnets, tunica crassa, xX 2-3 pum, neque rAamosis neque Septatts constante; contice nullo; spinukis braunnets, amplrs, 40-70 x #-8,5 pm, usque ad 35 pum omergentibus ; hyphis para- physoridets gnacitibus, x 1-1,5 pm, quasdam spinulas ctrcun- dantibus; basidiis narissimi(s, 12 x 3 pum, 4 sterigmatis 3 pum Longis; peramultis hyphis acanthophysoridets, tunica crassa, x 2-2,5 pm, non vel vix emergentibus; passin crystallis in- ter hymenium praesentibus; sports cylindratis, Leviter de- pressis, 4-4,5-(5) x 1,5-1,8 pm, hyalinis, uninucleatits, haud amyloiders, in massa akbrs. HOLOLU USE: ely on (LY), legends sBoidinen« LY //41% Ini suostia- to 4ncognito, Makokou, Gabon. 250 Ethymologie : espéce ainsi nommée du fait qu'elle posséde de nombreuses hyphes acanthophysoides (acanthophyses défi- Miles) pareAeee IEALTTenml 920). DESCRIPTION DU TYPE : Basidiome résupiné, largement étalé, adhérent, mince, 125 a 200 pm, brun sombre (5YR 4/2, fulbtgtneus de Saccardo) puis plus grisatre au centre par la pruine (5YR 5/2, benzo brown R. 4 2,5YR 5/2, cinnamon drab R.). Bordure fauve rouil- 1é (7,5YR 5 a 6/6, vers antique brown R.). Marge trés étroi- te, fibrilleuse, trés appliquée, blanche. Contexte (fig.5) bistratifié, dimitique. Hyphes géné- ratrices hyalines, a paroi mince, 2-3 um de diamétre, rami- fiées et cloisonnées. Hyphes squelettiques brun jaunatre, 4 paroi épaisse, 2-3 pm de diamétre, ni ramifiées ni cloison- nées. A la base du champignon (pas de cortex), le contexte est en- chevétré peu sombre (30-50 pm) puis passe 4 une zone sous-— hyméniale plus obscure (8-15 jim) ot les hyphes sont plus fermement enchevétrées et plus ou moins cimentées. Enfin, les hyphes se redressent pour donner 1'hyménium supérieur. Spinules brunes, massives, a extrémité peu pointue mais assez souvent mucronée, 40-70 x 7-8,5 jam, émergentes jusqu'ad 35 pm. Des hyphes paraphysoides gréles (1-1,5 pm) peuvent les entourer. Hyménium : l'ancien, collapsé, comprend des basidioles fripées, longues de 6-10 pm, des spinules et des éléments acanthophysoides brun jaunatre. Ces éléments naissent de la zone sous-hyméniale sombre. L'hyménium supérieur est formé de basidioles de 6 4 10 (15) pm de haut, de spinules, de trés rares basides 12 x 3 pm, 4 4 stérigmates de 3 pm et surtout d'abondantes hyphes acanthophysoides 4 paroi épaissie, de 2 a 2,5 pm de diamétre, non ou peu émergentes (10 pm). Ces sortes d'acanthophyses a paroi mince lorsqu'elles sont jeunes, nais-— sent d'hyphes squelettiques ou génératrices. Localement, des masses cristallines se rencontrent au niveau de 1'hyménium. Spores (sur sporée) cylindriques, légérement déprimées 4-4,5-(5) x 1,5-1,8 pm, hyalines, uninucléées (Giemsa), non amyloides, blanches en masse. Sur support indéterminé, Makokou, Gabon, 5 mai 1976, leg. J. Boldin. iHolotype »:) LY 9/741 (Herb. Boldin). AUTRES RECOLTES Cote d'Ivoire : LY 7053, sur support indéterminé, Abidjan, 5 novembre 1972, leg. G. Gilles. Gabon : Makokou (leg. J. Boidin), sur support indéterminé LY 7734, 4emairel 9/63) LY 27 /SO8eteL Ye i/o on wDemat who 70 atic Poel anthophysata ei tears rtinie ere OTE — Ry SE ae \ " ee — = = = i Y y i: 2) = nee 5 CA aie, . iat ; Ree eee SEE EE eee | pS we a L., y iS = £2) LA Ce Figure 5: Hymenochaete ac ies Klainedoxa gabonensis : LY 7759, 7 mai 1976; sur Anctstro- VNU CLUES eee Vey OS Osan Sema tenho / O- Gabon "Libreville (leg! G. Crlles)jj "sur supporteinacter= mines LY 9114.32 janvier 19/93eLY 915 eaetevrtermi7o oY 919854 mars 1979s LYe92 155 .12imars 1979s LYe925SezOenoven— bre 1978s" bYe92612 tl Omnovembrem | 97 Oslo 4/0) oma vet ee oe LY 9298. 22eavril NOD LY 9310. 2S ravines! Oo ee Limo ae. Amey une OO DISCUSSION Ces nombreuses récoltes permettent de mieux cerner les caractéres de H. acanthophysata, 4 travers les variations de la couleur de l"hyménium et surtout de la stratification ou non des é@échantillons Couleur de 1l'hyménium: l'abondance des hyphes acantho- physoides, lorsque 1"hyménium n'est que trés peu ou pas fer- tile, provoque un net assombrissement de la couleur. Ainsi, LY 7053, LY 7830 et LY 9417 par exemple qui sont a peu prés stériles ont une couleur brun rougeatre foncé (5YR 4/3 a 3/3 et méme 2,5YR 2/4 pour LY 9298); cette couleur est renforcée par un aspect velouté souvent trés marqué. Lorsque 1'hyménium est bien développé par contre, la couleur est plus pale: soit plus ¢risatres (25 5YRe 5/2. ae4)/ 2senava Pap rowneR sepOU TE YEO Ol par exemple) soit plus jaunatre (7,5YR 5 a 6/4, wood brown R. pour LY 7755). Une méme récolte montre assez souvent une telle variation de couleur: par exemple, LY 7750 est tantot chocolat (5YR-3/4)cet=veloute,stancolsenissmosatrer (oYRmo 25 benzo brown R.) et aride. De facgon similaire, LY 7755 (ainsi que 7/759) offre un trés bel exemple: le centre est en effet gris légérement rosé (5YR 6/2) alors que le reste de la sur- face est bai ferrugineux (5YR 4/6). Stratitication:ssurniunatocal deml/,recoltes siscilesm> présentent une stratificatwon: LY /734,eLY9258 ely LY. 7/741) (type) sont bistratifiés. LY 9273 a quelques strates mais seul LY 7750 (dont 1'épaisseur atteint 700 pm) montre 6 4 7 hyméniums successifs. Tous les autres spécimens (épais de 30 4 100 pm) n'offrent aucune stratification. Le basidiome a dans ce cas une structure plus simple: 4 la base, un mince contexte assez sombre (15-20 jim) formé d'hyphes enchevétrées en tous sens donne naissance 4 un contexte vertical dont les hyphes se terminent presque toutes en éléments acanthophy- soides; ceux-ci, comme les spinules, sont répartis dans tou- te l'épaisseur du basidiome. Qu'il s'agisse de spécimens stratifiés ou non, la plupart des récoltes sont presque ou totalement stériles: plusieurs sporées n'ont pu étre obte- nues et les autres n'ont que peu de spores. En conclusion, H. acanthophysata nov.sp. est une espéce Zoo stratifiée ou non, dont le basidiome étalé-adhérent est de couleur brun foncé, parfois velouté, a4 1'état substérile et plus grisatre et aride lorsque 1l'hyménium est normalement constitué (les basides étant de toute facon trés dispersées). La présence de nombreuses hyphes acanthophysoides est sans conteste le point le plus remarquable avec les spinules tra- pues et massives. HYMENOCHAETE HAUERSLEVII Léger nov.sp. Basidioma resupinatum, primum tenutssimun, e Luteo ochtaceo, dein paulo cnassius, cinnamomeum, srustulosum, Laxe ad subs- cukum e badio ferruginosum adnatum; ambitu indeterminato, concolone; trama Laxe interamixta, ex hyphis braunnets, tunica Ancnassata, x 4-5-(6) pum, septatis ramosisque constante, at- que ex hyphis hyalinis, tunica tenut, x 2-3 pm, septatis namosisque; cortice nuklo; crystaklis permultis ad et inter hyphas; spinulis braunnets, acutis, Longis angustisque, 90- 170-(190) x (4)-6-8 pum, usque ad 100 pm emergentibus; hyphis paraphysoidets gracilibus, x 2 um, spinulas scandentibus ;s basidtis saepius paulum constrictis, 12-17-(19) x 4-5 pm, 4 asterigmatis 5 pm Longis; pseudoacanthophysibus tunica te- nut vek Leviter incerassata, non vel vix emergentibus, 10- 13-(20) x 3-5 pum, paucts apicalibus diverticulis 2-3 pm Lon- gis praeditis; sports cylindratis (6)-6,5-7,5-(8) x 2,8-3- (3,2) pum, apieulo saepe manifesto, hyalinis, haud amyloiders. HOCOAUMUAM: aCopenhague’(C)e leo. K. Bjdrnekaer nn, Afr.22, Mine wito Lene lat. 60°C before being stained. TAXONOMY CYRENELLA Gochenaur, forma-gen. nov., nomen anamorphosis, Deuteromycotina, hyphomycetes pertains. Mycelium ex. hy pDhiLs Ryalinice ramocic, =levyi ver tunicatae compositum zygodesmatibus praeditae. Contdtophona hyalana .macronemsa ta, solitvariac. vel GCongregatae, erecta, irresulariter ramnosa. Cellulae eOnsdiogpense monodlasticae, apiecalia jet) lateral iar globosae, ovoideae vel cylindricae vel mixtae, vacuae’ post dehiscentem conidiorum. Conidiea hyalina, obovate. oe cum Drachiist apicealipus) Tredianti bus, 2-5 ou Ula ppend iver Dasa iaei Chlamydosporae et blastosporae sint productae. Species typica: Cyrenella elegans Gochenapr ETYMOLOGY: Prom, the Greek, Cyurenes. a-mythicalwwa ter sprite, an reference’ to the probable habitat or thes funeus and. the-resemblance of ts) conidia to theses of #thesanuatiuc hyphomycetes. CYRENELLA ELEGANS Gochenaur, forma-sp. nov. Figs. 1-2. Coloniae in agaro maltoso post dies decem ad 20°C 4-5mm diam. attingentes, humectae, calendulinae, 209 Sdinpactae, wn Cenurum acervulatum, margins planis, hyphis aereis raris. Hyphae immersae leves, zygodesmatibus praeditac, ad 2ilm crassae: Conidiophors hyalina, macronemata, solitaria vel congregata, erecta, mrregulariver ramoca, ad) 100uUm lonea, 3-4ium bateae. Cellulae conldiogenae monoblasticae, apicalia et lateralia, ovoideae mel cylindrvcae vel mixtac, vacuae post dehiscentem conidiorum, 5-12um longae, 3-hum latae. UGH waahivee Isa nal, mniceliularia,wornuecleolata., 10 x Vimy cum’ brachiis S0tea lupus Pradiaget bus 3-4. 8L3-20m Joneis. Lum Latis ad basen ct sO. >UmM la tis ad eap2cem, teu Uniappendici- eccentrica Dac arike wy, =i. OO. Olims Chlamydosporae abundae, solitariae et catenatae, intercalares et terminales, frequenter gregarae, hyalinae vel melleae pallidulae, binucleolatae, globosae vel subgiobosae leves et crassitunicatae, 5.5-6.5um Latae. pUuaLUuSs Siexualie Lenotis: In arena, subs lacceaqrta trucilisatd,, Hempetead jake bia ve: Park, Lone Leland. New fork, October, 1976s HOLOtyDUSs “CULULUra (pura Siecata, (numero (05), NYBG Herb. ETYMOLOGY: From the Latin elegans=graceful, describing the appendages of the conidia. DEVELOPMENT Germination. of the conidiarot 0, felegans occurs within 24 hr on MEA at 20°C. One to 4 germ tubes arise apically between the appendages and/or from the base, but rarely from the lateral walls. Clamped septa and branches develop where the germ tubes emerge from the conidium. As the colony ages, conidia may germinate in place producing apically, laterally or from the base, one or more conidiogenous cells that develop conidia (Fig. 2C). Cytoplasm flows into the conidiogenous cell leaving the Original conidium empty. This process may continue through several cycles, so that short chains of empty conidia are common. AS a result of this proliferation, the color of the central area of the colony is diluted and much lighter than the periphery (Fig. 2A). Growth is slow. Colontes attain a diameter of 3 mm in 7 da and approximately 24 mm in 35 da. They are Persian to chrome orange (6A 7-8) with a raised, rugulose central region consisting of pulvinate masses of conidiophores and conidia and a narrow plane Margin. With age, a broad, radiately wrinkled periphery composed of mycelium and chlamydospores develops (Fig. 2A). It is somewhat fiberous in appearance with small mounds of sporulating cells scattered over the surface or coalesced to form larger irregular or wedge-shaped patches. The surface is moist. Aerial hyphae are rare or absent; when present they are usually aggregated into tufts. The myceltum is hyaline and consists of sinuous and straight hyphae of uniform width, 2.0+0.3um diam, with smooth, Slightly thickened walls. Septa with clamp connections dome b eb Lrot OCCUL approximately Joa llim Prom theacip and then’ at_intervals of approximately 507m. The cells are binucleate. A dolipore apparatus was not revealed in preparations stained with ammoniacal Congo-Red (11). 21-0 Us ieet ee Cyrene lia, elegans’. Cr Con dilact (RB) Stages wp tinetgcve lopment aor GonidiLogenous celts. and clustered chlamydospores. of cytoplasm. (C) Mature conidiophore. (D) Single Unstippled areas are devoid RACAL Adventitious simple septa are common in older hyphae and coralloid or haustorial branches similar to those shown by Otaeve (3) forvlt lobastdzum may occur. Contdtophore development begins within 2-3 da. The conidiophores arise as erect, branched hyphae indistinguishable at first from the vegetative cells. They are hyaline, of variable length and produce thumb-shaped terminal and lateral branches that stain intensely with basic dyes (Fig. 1B). These branches are separated from the conidiophore early in development by clamp connections. They also may be produced individually along the vegetative hyphae (Fig. 2B) or less commonly from a clamp. They may differentiate directly into conidiogenous cells or may branch once or twice, each branch becoming a conidiogenous cell separated from its parent cell by a clamped septum. The irregularly branched conidiophore at maturity (Fig. 1C) is generally less than 100um tall, 3-4um wide at the base and is best observed using phase contrast optics since its upper half consists of cells devoid of cytoplasm and somewhat collapsed. Conidiophores are often aggregated into sporodochial-like clusters that arise from a basal mass of intertwined hyphae and chlamydospores. The conidiophores produce single or clusters of 2-3 conidiogenous cells laterally and terminally along their branches. The contdtogenous cells are phialide-like, binucleate, have a thick-walled body and a thinner-walled neck that is approximately 1-4um long and 1-2um wide. They vary in form from globose to ampulliform to cylindrical and measure 2-4um wide, 5-12um long including the neck. Each produces a single conidium. Mature conida appear within 4-5 da. Contdta (Fig. 1A) are blastic, orange in mass but hyaline by transmitted light, binucleate (Fig. 2C), obovate, thin-walled, 10 x 4um (eit. 2 x 4.24 5m) and bear 3-4 long,, narrow, divergent, flexible, apical appendages, 16+2.4um long, lum wide at the base and tapering to 0.5um wide at the apex, and an Sccentric paca lmappendagersor suntftormewiath, 96 9F2 alex. Ooum. At 20°C, approximately 33% of the conidia examined had 3 apical appendages; 66% had 4. At temperatures above 25°C, aDOULeLSeeOf the conidiashad one, 27 and)5@arms:. Chlamydospores (Figs. 1D, 2D) appear after 3 da. They are Single and catenulate and arise terminally, intercalary, or less commonly from clamps. They typically form in large compact clusters behind the margin of the colony and just above or below the agar surface. They are subglobose to globose, 6+0.4um diameter, binucleate, hyaline to pale vamber, thick-walled and filled with guttulae. Single ones are often oval and/or unsymmetrical and borne between several adventitious septa that form as the cytoplasm retracts into the developing chlamydospore. Chlamydospores germinate within 3 da at 20°C (Fig. 2E). A single germ tube that branches and develops clamps where it exits from the spore is produced and a typical colony with conidia develops within 5 da. Chlamydospores remain viable after storage for up to 5 mo at room temperature in distilled water. Those held at 7°C exhibited a progressive FU. 2 Cyrenella elegans. (Ny) Colony on melt extract agar after 35 da. at ish deus (B) Vegetative hypha with a clamp connection. Arrow designates a young conidiogenous cell. (C) Binucleate Arrow designates a young conidiogenous cell and conidia. situ germination. secondary conidium produced during. in Three empty conidia appear on the Left. Ps decrease in the percent germinating after 45 da storage and showed no germination after 5 mos. Karyogamy, probasidia and soredia were not induced by these conditions, nor do these spores resemble, in either coloration or mode of formation, the teleospores produced by members of the genus Rhodosportdium (2, 3, and comparison in culture with Peemcacryovaum Kells, Hunters, Tallman ATCC 24502. & ATCC 24503). In view of the above, it is considered likely that these structures are true chlamydospores and not the teleomorphic state of ¢. elegane. Dedikaryotization occurs spontaneously, but infrequently, as the culture ages; a uninucleate yeast phase results. It can be detected readily by streaking material onto an agar medium containing raffinose, a sugar utilized less well by the filamentous stage. Origin of the blastospores is unknown. Subjecting the dikaryon to flooding with water or anaerobic or microaerobic conditions while growing it on media with various C/N ratios and low or abundant in nutrients failed to consistently induce the yeast phase. The yeast colontes, like the dikaryon, grow slowly, reaching a diameter of 8-10 mm on MEA at 20°C after 30 da. They are slightly elevated when viewed in cross section and show faint radial lines. Their surface is semi-glossy, deep orange to Persian orange (6A8-7A8) in color with a Slightly deeper carrot red reverse (7B7). The yeast cells (Fig. 2F-G) are unicellular, monokaryotic, non-encapsulated, elliptical to cylindrical with rounded ends, less commonly epclava leo .0+) , 86x, 3..2+0. 81m, andsreproduce, by terminal and subterminal budding. Fermentation is absent. In ME broth, most cells are single and only rarely are short chains of cells observed. No pellicle is produced. Evidence suggests that the nuclei in the dikaryon are identical. A dikaryophase is not reestablished when yeast cells from randomly selected colonies are mixed on cornmeal agar. However, it develops spontaneously in every cloned yeast colony that is over a few days old. Cells in the center of a colony at the agar surface become binucleate. It is uncertain whether these occur following conjugation between two cells or by failure to bud during mitosis. Observation of empty single yeast cells with an attached binucleate conidium suggests that the latter is the case. A sparse mycelium always develops under every yeast colony but its further expansion is limited unless the overlaying cells are scraped away. All cells on the surface of a colony remain monokaryotic so that the yeast phase can be maintained indefinitely in culture but always contaminated eventually with the dikaryon. (D) Binucleate chlamydospores. (E) Germinating chlamydospore. (F) Budding blastospores. (G) Uninucleate blastospores. Material in photographs B-G was stained with Coluidine, blue. Cells in photograph G were pretreated with Pench tore Se minemastu60 2C, 274 NUTRITION C. elegans is an acidophilic, mesophilic, obligate aerobe. Growth occurs over a wide range of H-ion concentrations. It is best between pH 3.5-6.0 and absent below pH 2.5 and above pH 7.4. Conidia do not germinate at 5°C and 37°C. Pigmentation iS more intense and colonial and cellular development best between 18-23°C. Growth is fastest at 30°C; absent above 34°C. Table I. Growth of Cyrxenella elegans on various nitrogen sources® Compound Growth Compound Growth 1., Potassium, nitrate + 6.9 Glutamic acid + Cot POLAS SLUM Ww iat ra te ~ [. Asparagine + 3. Ammonium sulfate + G7 ) Casamino tacids ++ 4, - Urea + O*e G eutart in. ++ Sp araspearct cea Cand as 20. ‘Sodium Caseinate ++ a) Cultures: were Aancubated fore5 da. ate co. Cc. +t Growth equal to growth ‘on caseamino acids + Growth weak (but, better “thanyecrowth on’ a mediumplacking a nitrogen source - No growth C. elegans utilized all nitrogen sources tested with the exception of potassium nitrite (Table I). Growth and development are best on a mixture of amino acids. The proteins, gelatin and sodium caseinate, are utilized but diffusible hydrolases were not detected when colonies on these substrates were flooded with acidified HgCl.,. A variety of compounds can serve as the carbon-energy source (Table II). Growth and development are best on glucose or fructose. Krebs Cycle acids (Nos. 37-38) are assimilated but not compounds with ester linkages (Nos. 23-24). Among the polysaccharides (Nos. 17-22), only the fructosan inulin and the starches (Nos. 18-19) were weakly degraded. Diffusible exoenzymes were not detected when colonies growing on the latter compounds were flooded with an IKI solution. Patterns of carbon and nitrogen assimilation by the blastosporic and dikaryotic stages are identical. The exogenous vitamins, p-aminobenzoate (PABA) and thiamine, are required for growth of both the mycelial and yeast phases. High concentrations of folic acid can be substituted for PABA but not a mixture of amino acids and purines. DISCUSSION Cyrenella elegans is a member of the Basidiomycotina because it produces mycelium with clamp connections but the absence of a telemorphic state makes its phylogenetic position in the subdivision uncertain and any attempt at ordinal placement a purely speculative venture. 2D Peple Il. .Growth of Cynzenella pedal on various carbon-energy sources® Compound Growth Compound Growth ie. Ay Lose plat ian.) ae Gh ~ Oe. L-Arabinose ak ane DNA - 3. Ribose - 24, Triglyceride - Wa Glucose ++ 25. Fucose = 5. Fructose ++ 26... L-Rhamnosce ++ 6. Mannose ++ 27 Glucosamine = Galactose - Oo ein thane | - 8. L-Sorbose ~ 29. Glycerol ++ 9. Trehalose + 5 Oren Gye ILE? 166.0 - A) Sucrose aap Biles DOr Ost OL ++ Thane Maltose tacts Bier Mannitol Par ieee Melibiose - Se Lnositol - ieee Cellobiose + Si eeeGalactitol - mo i Lactose a 35. Acetate = 15. Melezitose ++ 26... DL-Lactate = fore Raftinoce + 237. Succinate ++ igs snl + 3.0 eC Lo GaLe + 18. Amylose + 30 ee Dar Gen “= 19. Amylopectin + WOe 4oal Loan + 20. Pectin - i Vandel n - mits Celiulose z. re. Casamino acids = ayaa compounds were D-isomers except where noted. Cultures onwcompounds 17-24 were incubated for ll «da at 126°C van. Gunersmcor)> sda at oo 7 ek tin.rOow Oredila lato Om bevteratian ~2rowth "on —slucose + Growth weak but better than growth on a medium lacking a-carbon source ea bOwtd meCduad le tLOuOr Less (than growth on .a medium Lacking a carbon .source The blastosporic stage could be readily accommodated in the genus Rhodotorula (14) if the dikaryophase were lost. It resembles R. aurantiaca (Saito) Lodder and R#. lactosa Hasegawa in producing cylindrical cells, in colony color, in utilizing nitrate, cellobiose and maltose and in requiring PABA and thiamine. It differs from the former in being galactose negative and L-rhamnose and raffinose positive and from the latter in being unable to utilize melibiose and lactose. The requirement for PABA is confined to a very few wild-type fungi, certain species of Rhodotorula, Blastoecladta pringshetmit Reinsch and single strains of Saccharomyces cerevtstae Hansen (4). Because this requirement is so uncommon, there appears to be strong selection pressure against such fungi in nature. Its Simultaneous occurrence in Rhodotorula-and Cyrenella may Signify a much closer relationship between these genera then is evident simply on the basis of similarities in morphology and nutritional patterns. C. elegans was obtained from sand along the shore of 276 a fresh water lake in an area that is regularly submerged each spring for several months. Even though it came from a terrestrial habitat, C. elegans has several characteristics in common with organisms isolated from aquatic environments. Its thin-walled, tetraradiate-like conidia bear appendages in a pattern reminiscent of that evolved by many fresh and a few salt-water fungi. They are remarkably similar to the basidiospores of the marine fungus Wia in size and shape but not development (9) and superficially resemble the conidia of aquatic hyphomycetes belonging to the genera Acaulopage Drechsler, Clavatospora S. Nilsson and Clavartana Nawawi. Diffusible exoenzymes are a characteristic feature of terrestrial fungi (1, 8) but would be a distinct disadvantage for an aquatic organism since such proteins and their digestion products would be lost into the water. C. elegans produces no diffusible hydrolases when utilizing starch, gelatin and casein. Since its hydrolases must be retained in protective association with the cell wall, it would be well adapted for life in dilute aqueous environments. Finally, C. elegans' requirement for PABA occurs less frequently among terrestrial fungi than among aquatic ones. In R. auranttaca, for example, only strains of marine origin demonstrate an absolute need for PABA (14). These features suggest that C. elegans is not a terrestrial fungus but is an aquatic or possibly an amphibious one, whose propagules were trapped in the sand as the lake water receded, persisting there until isolated several months later. If Cyrenella is an aquatic genus, it is an extremely uncommon One., Dr. R.«Bandonivof the Univers1rty.of British Columbia (personal communication) reports that C. elegans appears to be identical to a fungus he collected from a driftwood chip on the shore of the Iowa River about 25 years ago. Other workers who have examined the conidia report they do not remember seeing spores of exactly this type in foam samples nor have illustrations of these spores appeared in the many works describing propagules from this environment (6, 7). With its slow growth rate in culture, superficial resemblance to Rhodotorula and tiny conidia, it is understandable why a Cyrenella could be overlooked. ACKNOWLEDGMENTS LaWisen to tTheank tHestollowings individuals For their aids Dre sR. Bandoni forsstudying cultures, or CUsenc Cede ond. offering many shelpPrul suggestions as welbtlas kindly YPEVIieWwing the umanuscripts Drs. W. Bridge Gooke, J. Leland Crame, C. Tt -dngold, Rs H. Petersen; and yshun-—1ehi Udacawa hor Pe ranmining slides eop thestuneue:.0r. 4c. el . hogereconet or assistance with they taxonomy; Dr. A. H. Bréenowitz ior takpnesthe photomicrographs: Prot. Lh. Ascher, language Dept. Tonecorrectine the Latam diagnosis: Ms .Julie Anderson, a former undergraduate in my laboratory who studied Cyrzenella with me as part of her honors thesis; and Mrs. W. Jaworski for typing the camera-ready copy. Dhal LITERATURE CITED meee Domsch, K., and Ws Gams. 1969. Variability -and Pocemtial of a souid fungus: population to decompose pectin, xylan and carboxymethyl cellulose. SOid Biol. Biochem. me (29-36. Pree eed a ce Phath and S.aNewell. BS yy Rhodosporidium Banno. pp. 803-814. In: The Yeasts. Ed., ae LOUder. end ed. North-Holland Publishing Co., Amsterdam. Cree ce Ll Sais else pHUR Ger, and. A. Valiman’. eae, ar Pie basidiomycetous.yeasts (Rhodosporidium spp.n.) with setrapolar and multiple allelic bipolar mating systems. Pied | Microbiol) )19: 9643-657. Lig Fries, N. 1965. The chemical environment for fungal growth. 3. Vitamins and other organic growth-ftactors. Pee t91-523- In: The Fungi. _Eds., G.” Ainsworth ‘and MmEicocman. Vol. I. Academic Press, N. Y.. N.Y: i. ia Sh Bee Me Me ees Pa iar 1OTO.. Atconol@ te LOolvidine vplue: A rapid Memo LOr staining nuclei an upiixed mycevozoa and fungi . Mycologia 62: 406-07. 6 Mascot ds Cx. NEO TG Advances in the study of so-called aquatic hyphomycetes. Amer. J. Botany 66: 218-226. foeericOld, ©. bO1>. ~ Guide to aquatic hyphonycetes: Freshwater Biol. Assn., Ambleside, England. ‘eae Cyol Per A, and=p .) -aStruwe. LO OF. Microtung i 16 f decomposing red alder leaves and their substrate Peabication.) SoriyBiol. Biochem. 12:7" 425-431. OF Kohimeyer, J., and E. Kohlmeyer. LOO. Marine mycology the higher fungi. feademiouPpress . Nia ey. seth ede 690 p. Move kornerup, A.;, and J. Wanscher. POOTE Methuen pemovook Of colour. 2nd ed.) Methuen and Cos, )Ltd., london. Uns = p's Jil. Nawawi, A., J. Webster and R. Davey. 1977. Dendrosporomycees prokifer gen. et sp. nov., a basidiomycete Birhsbranched conidia. Trans. Brit. Mycol.’ Soc. 68: 59-63. mer Newell. S., and J. Fell. LO FOR The perfect form of a marine-occurring yeast of the genus Rhodotorulka Myeotogia 62: 272-261. Ss Oa Wei, 5 ste 1968. An unusual new heterobasidiomycete with Tilletia-like basidia. pp. 261-266. In: Mycological moumtes honoring John N. Couch. Ed... We J: Kock. The Univ. Peecarolina Press, Chapel Hill. Beeeenaff, H., and D. Ahearn. 1971. Rhodotorzufla Harrison. Peeelilof-1223) “In: The Yeasts. |Ed. J. Lodder. 2nd ed. North-Holland Pub. Co., Amsterdan. MYCOTAXON Vol rex Vile N Ota. ap Demet OF 776.0 Apr un eeeho & 1 REVUE DES LIVRES par G.L. HENNEBERT Book Review Edttor, Crotx du Sud 3, B-13848 Louvatn-La-Neuve Belgtque GENERA OF HYPHOMYCETES, par J.W. CARMICHAEL, W.B. KENDRICK, I.L. CONNERS and Lynne SIGLER, 390 p., 129 pl., 8°, reliure spirale, 1980, The University of Alberta Press, Edmonton, Alberta, Canada. iD aalp ae jeuc all Frosh Cet ouvrage est une seconde édition entiérement revue et complétée de la premiére psésentation par les premiers auteurs d'une compilation illustrée et critique de la littérature originale des genres de Hyphomy- cétes dans The Fungt de Ainsworth, Sparrow and Sussman, Tome IV A, en 1973. Les auteurs ont recensé cette fois plus de 2000 noms de genre, contre 1541 en 1973. Les 61 planches de figures d'alors sont devenues 129 aujourd'hui. Aprés une introduction sur la méthode et la classifi- cation des types conidiogénétiques, on y trouvera une liste documen- tée des noms de genres, avec date, référence, type morphologique, hdte, espéce type, statut nomenclatural et synonymie, avec références. Cette liste distingue les genres acceptés par les auteurs des noms rejetés ou douteux. Viennent ensuite les illustrations, soit reproduites des des- criptions originales, soit dessinées a partir de publications récentes ou encore du type. Les 853 figures représentent pour la plupart les espéces types des genres et sont identifiées a l'espéce. Le grossisse- ment n'est pas donné mais n'est pas indispensable a ce niveau. Suivent encore 5 index: un index des genres par type conidiogénétique, un index des connections entre téléomorphes (stades sexués) et anamorphes (stades conidiens), une liste bibliographique, une liste des abréviations et une liste des figures. Les auteurs ont suivi la nomenclature illégale basée sur Persoon, Synopsts Fungorum, de 1801 comme point de départ, contrairement a l'art. 13, mais combien souhaitée par beaucoup. Le livre remplace les guides des genres publiés jusqu'a présent. Il est fort bien imprimé. Sa reliure en spirale s'inspire de celle du Barnett, Illustrated genera of Imperfect Fungt, mais on pourra le regretter.Vu son évidente utilité et son prix assez modique, le livre aura sans doute une large diffusion. Les auteurs l'ont dédié a la mémoire de Luella K. Weresub, mycolo- gue du Biosystematics Research Institute d'Ottawa, tant appréciée de OUGT CHAMPIGNONS D'EUROPE, par M. SVRCEK et J. KUBICKA, adapté en fran- Gais avec coll. de G.L. HENNEBERT et al...296 p., 448 ph. col-, 8°, cartonné, 1980. Ed. Elsevier-Sequoia, B-1940 St Stevens-Woluwe, Belgique. Ce guide des champignons est remarquable par le grand nombre d' espéces décrites - 448 - chacune convenablement illustrée d'une photo en couleurs dans le milieu naturel. Les descriptions sont précises 209 et donnent les dimensions sporales. Une introduction illustrée retrace la structure des champignons et leur classification, donne les conseils au récolteur et le documente abondamment sur la toxicité des champignons. Offrant un éventail exceptionnellement large des champignons d'Europe, tant de montagne que de plaine, ce guide surpasse un grand nombre d'autres moins complets. ECOLOGY AND DISTRIBUTION OF FUNGI, Scripta Mycologica n° 9, 144 p., 12°, broché, 1980, Academie des Sciences d'’Estonie S.S.R., Tartu MstonLe.errix., RD il, LO. Ce fascicule est intitulé en fonction des deux premiers articles quil comprend : The composttton and seasonal dynamtes of the fungal cover on mtnera sotls and Trophte groups of Estontan Agartcs, par K. Kalamees. Dans le premier travail, l'auteur reléve les flores d'agari- cales dans divers types de foréts et de prairies jusqu'a la tourbiére et le marais et note sa composition et sa richesse spécifique 4a chaque écotype végétal. Dans un second article, il regroupe les Agaricales en 13 groupes trophiques, les mycorrhizogénes, les saprophytes de divers types de substrat et les parasites étant les principaux d'entre eux. Le fascicule se termine par une revision du genre Lastobelontum par A. Raitviir, ot 1l'auteur combine diverses espéces et en décrit cing nou- velles. Texte en anglais. THE CHEMOSYSTEMATICS OF THE LICHEN GENUS PERTUSARIA IN NORTH AME- RICA NORTH OF MEXICO. par Martyn J. DIBBEN, Milwaukee Public Mu- seum, Publications in Biology and Geology N). 5, iv + 162 p., 136 fig., 4°, broché, 1980. Milwaukee Public Museum, Milwaukee, Wi. DoZo Sun UOA Prix fy 220 te postage. L'examen de pas moins de 6800 specimens, incluant entre autres des matériaux historiques de quelques 40 herbiers reconnus, a été né- cessaire 4a cette revue des 66 espéces North Américaines des lichens du genre Pertusarta. 14 espéces sont nouvelles. 33 espéces sont endémi- ques a 1'Amérique du Nord. Du point de vue de leur distribution écologi- que, 60 2 spnt tempérées, 35 % arctoboréales et 5 4 tropicales. 40 acides organiques sont recensés chez les Pertusarta, L'auteur a appliqué 1' analyse chromatographie sur silicagel a tous les échantillons, selon la méthode de Culberson. Sa taxonomie des espéces est avant tout basée sur la morphologie du thalle et de la fructification apothéciale. Sur cette base elle distingue deux sous-genres, P, sbg. Pertusaria et P.sbg. Ptonospora, l'un a fructification amplicariale ou pertusariale, l'autre a fructification lécanorale ou sorédiale, L'analyse chimique s'avére une confirmation précieuse dans la distinction des espéces. Confrontées entre elles, les deux méthodes permettent de réduire le poids donné aux variations et de ne plus subdiviser l'espéce en une série de variétés ou de formes, La différenciation chimique jointe 4a la ségrégation géo- graphique a permis cependant de confirmer des espéces proches comme dis- tinctes. L'espéce type Pertusarta pertusa, eurasiatique, n'est pas recon- nue en Amérique du Nord, celle qui lui a été confondue est décrite comme P. consoctans. L'auteur a le souci d‘une nomenclature correcte. Elle indique le nom du genre comme nomen conservandum, notant qu'il est illé- gitime (Art, 63), et référe sans plus de détail - A regret - 4 un article futur. Ses synonymies précisent lestatut nomenclatural et sont justifiées par des commentaires, Les descriptions des espéces d'une liste détaillée des spécimens examinés et d'une distribution géographique illustrée. La 280 technique macrophotographique appliquée laisse a désirer. Cette monogra- phie 4 la fois nouvelle, précise et didactique mérite 1!'attention: A PRELIMINARY POLYPORE FLORA OF EAST AFRICA, par Leif RYVARDEN & Inger JOHANSEN, 64° °p., 212 fig. , 8°) couverture papier, 1980, Funghtlora: Ed’ -P50.Box 95") Blindern, Oslou3, orway.s (PP) ix, Nka 200; Durant les 10 ans passés, alors qu'il publiaita intervale régulier les volumes des flores des Polyporaceae et des Corticiaceae d'Europe du Nord (revus dans Mycotaxon), le premier auteur a élaboré cette flore des Polypores' d'Afrique orientale par diverses explorations de récol- te et 1'étude des grands herbiers européens. La flore couvre 5 familles, les Corticiaceae, Ganodermataceae, Hymenochaetaceae, Polyporaceae et Tremellaceae, 75 genres dont 4 nouveaux (Antrodtella, Echtnoporta, Navi- sporus et Pseudoptptoporus) et 337 espéces dont 3 nouvelles. Des clés réellement dichotomiques des genres et espéces sont poposées et, évidem- ment (et pourquoi?) limitées aux seuls genres et espéces traitées. Les espéces sont décrites avec une attention particuliére aux détails anato- miques et microscopiques, lesquels sont dessinés dans la plupart des espéces. L'habitat et la distribution géographique sont indiquées, mais sans liste des spécimens étudiés (le livre se veut @6tre une flore et non une monographie). Imprimé en caractéres dactylographiques, le texte est sans relief, peu attirant. Les nombreux espaces blancs donnent au livre une épaisseur de 35 mm qui fera rapidement céder la faible couverture de papier. Le livre est dédié 4 la mémoire de M.A. DONK. SAPROPHYTIC MICROFUNGI FROM TAIWAN. PART 1 HYPHOMYCETES, by Takashi MATSUSHIMA, Matsushima Mycological Memoirs No 1, 82 p., 46 fig., 8°, paperback, May 1980. Publication Matsushima Fungus Collection, 23-19-6001 Mikageyamate-2-Chome, Higashinada-ku, Kobe, Japan 658. Price: free of charge trough official request. This is the first of a series of papers enumerating the microfungi collected in Taiwan during 1976 to 1978, by the author when he was guest as visiting senior mycologist at the Plant Pathology Division, Plant Protection Center, Taiwan. As in previous publications, the authors reports his profuse collections and cultures of the Hyphomycetes with the same wonderful ability and efficiency. His descriptions or comments on species are just what is needed, but his ful page drawings speak more than words. Not less than 250 Hyphomycetes including 2] new species and 2 new genera(Acumtspora and Chetropolyschema) are recorded, documented and 46tillustrated on the 82 pages of this issue. This is the start of a new mycological series entitled Matsushtma Mycologtcal Memoirs. We are sure of the succes which this series will receive through the mycological world. This series is distributed by the author's newly established institution, a very well standing fungus herbarium and cul- ture collection, the Matsushtma Fungus Collectton. BIBLIOGRAPHIA BOTANICA CECHOSLOVACA 1973-1974, par A. Neuhaduslova- Novotna et D. Guthova-Jarkovska, 564 p., broché, 14x20 cm,Botanicky Ustav GSAV, Prithonice u Prahy, 1978. Ce répertoire bibliographique donne toute la littérature botanique publiée en Tchéchoslovaquie durant la période citée, classée par matiére (Sle ieee EHONEEUNE ef ———e 281 UNTERSUCHUNGEN ZUR KONSERVIERUNG DE FRUCHTKORPER DES SPEISEPILZES PLEUROTUS OSTREATUS UND DER PARTIELLEN AUTOLYSE VON PILZSELLWANDEN, par Helga SCHMITZ, Bibliotheca Mycologica vol. 77, 98 p. Se yidos papier, 1980, J. Cramer ed. Dans 1'étude du mécanisme enzymatique de 1l'attendrissement observé lors de la conservation de carpophores dans des solutions acidulées ac€tique, citrique et lactique (pH 4) l'auteur met en évidence 1'action de R-glucanase et chitinase spécifiqueSdu pied et du chapeau du fruit, tandis que les S-glucanase et chitobiase du mycelium sont sans action. Thése réalisée sous la direction du Pr. G. Eger-Hummel, Marburg. MUSHROOMS OF IDAHO AND THE PACIFIC NORTHWEST. DISCOMYCETES, par Edmundtine TYLUTRIs 133° pe9027) £10.45 80 phot. =) Loxzlem, 1979; University Press of Idaho, University Station Box 3367, Moscow, Idaho 83843, USA. Ce livre est écrit pour les amateurs "enthousiastes" qui veulent dépasser les fontiéres des basidiomycétes et s'aventurer a récolter les morilles, helvelles, truffes et autres grandes Pezizales, Tubérales et Geoglossacées. Chaque espéce décrite est illustrée d'une photo blanc- noir. Une clé "de terrain, purement macroscopique" est proposée: mais elle est dichotomique et couvre 5 pages bien pleines... qu'il faut lire a chaque récolte? Une clé synoptique serait plus didactique et plus rapide. Cette remarque dépasse le cadre de ce livre qui est bien écrit et agréable. Pas moins de 122 espéces y sont classées et reconnaissables par les clés, CBS COURSE OF MYCOLOGY, par W. GAMS, H.A. VEN DER AA, A.J. VANDER PLAATS-NITERINK, R.A.SAMSON, J.A. STALPERS, 2e édition, 110 pe, JOR tL OM einaG sedosepaprer, —ChS\, baarn, Nederland. 19802 Le texte de ce cours (le édition revue dans Mycotaxon 3:558, 1976) est revu et complété de la littérature récente, sous a peu prés le méme volume. Les auteurs y développent les traits généraux de la morphologie des champignons, dans chaque groupe taxonomique, étayé de nombreuses figures et références. Le fascicule ne se veut pas un guide d'identifica tion (voir von Arx, Champignons sporulant en culture pure, 3e éd. en prép.) mais une bonne introduction au régne des champignons. A REVISION OF CHRYSOSPORIUM AND ALLIED GENERA, par C.A.N VAN OORS- CHOTeGStudiesiin Mycology sin -@20,289Fp. 86 fiptes oi ple.eines sados papier, 1980, "CBS. eBaarn, Nederland.) Prixghti25.-% Cette monographie donne les descriptions illustrées et les clés de Chrysosportum (22 espéces), Myceltophthora (8 espéces), Emmonsta (2 var.) Zymonema (1 esp.) Trtchosportella (2 esp.) et Geomyces (3 var.). La coni- diogénése blastique et thallique de ces champignons est décrite, ainsi que les relationsavec les stades téléomorphiques appartenant aux Gymno- ascaceae, Onygenaceae, Ascosphaeraceae et Sordariaceae. Cette revision est la premiére depuis le travail de Charmichael de 1962. EESTI SEENTE KOONDNIMESTIK ( LIST OF ESTONIAN FUNGI, WITH HOST INDEX AND BIBLIOGRAPHY) par J. JARVA et E. PARMASTO, 331 p., Din, dos papier, 1980, Institute of Zoology and Botany, Academy of Sciences of thesestonianis. so. Rolantu. pEstouiaraulrixcear Dl vee sha Boe Cette liste des champignons de 1'Estonie est le résultat de 15 ans de travail des mycologues de 1'Académie de Tartu pour le dépouillement de 1200 publications citées en annexe. Chaque espéce est citées avec hé- tes et références. BIOLOGY IN THR BLUE RIDGE, Fifty Years of the Highlands Biological Station, 1927-1977, by Ral piisMs,SARGENL. #l>6up ae Liao a epapeien back, 1977, Highlanas Biological Station, P.O.Drawer 850, High- lands N.C. 28741. The then named Highlands Museum and Biological Laboratory, initia- ted in 1927 by a few determined people eager to preserve local human and natural history»has been, from the early years, associated with mycology. The Highlands station, in 1934, received the Mycological Society of America for its summer foray. Dr. William Chambers Coker, who presided the institution from 1933 to 1943,and subsequently as Honorary President, has been, with Dr L-R. Hesler, Vice-President, the promotors (of ‘the’ stu- dy of the Appalachian fungi. Coker published on basidiomycetes, parti- cularly Hydnum and the Boletaceae (1939-1951). Hesler continued his Wotes on Southern Appalachtan Fungt in Mycologia from 1936 to 1955, and colla- borated with Alexander H. Smith on Hygrophorus and Lactarius (1932-1962). Dr. L.S. Olive (1943-1953) published on Tremellales and in 1965 his book Potsonous, Edtble and Halluctnogente Mushrooms. Dr. Ronald H. Petersen, more recently investigated the clavaroid fungi and the aquatic hypho- mycetes (1962-1963). Surely, the Blue Ridge Laboratory contributed much in 50 years to the American mycology. The Sargent's book, retracing, years after years, the multiple ways and reviving the unique spirit of the Highlands Biological Station will please the present and the future scientific people. PHYTOPHTHORA CINNAMOMI AND THE DISEASE IT CAUSES, by G. A. ZENT- MYER, Monograph No 10,-96 p., 39) figs’,°4°, paperpack,1980, The American Phytopathological Society, 3340 Pilot Knob Road, ‘St’ Paul MNS D2 errr cetUses: Oar How is it possible to cover such widespread and largely investi- gated fungus pathogen like Phytophthora cinnamomt in 95 pages. The author did, and on the base of 600 literature references. The pathogen is of importance for the decay of many tropical and temperate crops and trees. 900 hosts have been listed. The monograph details the many aspects of the biology, ecology, physiology and genetics of the pathogen, all data necessary in the search to. an effective control. The text is dense and illustrated of many original documents. George A. Zentmyer of the University of California, Department of Plant Pathology is devoting himself on the study of that pathogen for already 20 years and published with his collaborators more than 100 papers onthe subject. His master- ship in that fidd will help many people engaged in research through the world. A MANUAL OF ASSESSMENT KEYS FOR PLANT DISEASE, by Clive JAMES, 44 fiches illustrées, 10x18 cm., 1980, The American Phytopathologi- ealisociety 7, *StePatl< eprixrUsens 1020. Ces fiches illustrent les clés d'évaluation des dégats par Rhyncho- sportum, Puecetnta, Septorta, Drechslera sur céréales,Phoma, Leptotrochila Stemphyltum sur luzerne et tréfle, Phytophthora, Streptomyces, Rhtzocto- 283 nta sur pomme de terre et Xanthomonas sur haricot. Elles aideront les phytopathologistes a effectuer des évaluations comparables. LABORATORY HANDBOOK OF MEDICAL MYCOLOGY by Michael R. McGINNIS, 662Ep ssuilin, 68" ,ahard cover, 91980. AcademicePress a N.Y- "Owing to the increased importance of fungi in medecine, there is a pressing need to discuss important topics such as laboratory safety and emergency procedures, quality control and modern concepts,... to assist microbiologists in safety isolating and acurately identifying fungi of medical importance.... The key to understanding any field of science ne- cessitates a thorought understanding of its language..." (the author). Thus, the author devotes its first chapter to the morphological terms to be use in modern medical mycology for the description and the classi- fiecarionpoltetve pLuncie while chapter 40(2331p.) ~and.chapter 9 (63 1p..) describe the systems of classification, detail the methods of study and comment the diagnoses of genera of, respectively the Ascomycetes, Coelo- mycetes, Hyphomycetes and Zygomycetes (sic) in the former and the yeasts in the later. Chapter 9 (50 p.) adds a'synopsis of the mycoses, descri- bing and illustrating symptoms, etiology and therapeutics of each disease. The other chapters of the book are descriptive of technics: laboratory safety, handling clinical specimens, bioessay procedures, susceptibility testing, quality control, equipment maintenance, culture collection and culture media. Appendices provides with glossary, list of synonyms and list of fungal records on man. The book is no doubt interesting and useful, not only for the tech- niques described, but as an effort of providing medical mycologists with the developments in modern descriptive mycology, especially in the field of the Hyphomycetes, after the First Kananaskis Conference of 1969. Non obstant such evident value, the book presents several major defects that myself at least, I do regret. In the general disposition of the contents, the book might have been divided advantageously into two parts, one part, taxonomic, including chapters 1, 4 and 5 (the techniques excluded), chapter 9 and the appendices, the other part, technical, grou- ping chapters 2 and 3 with chapters 6 to 10, and the culture methods from chapters 4 and 5. That grouping might have avoided regrettable duplications between chapter 1 and chapter 4 in text and illustration. The illustration is, beside a few line drawings, essentialy made of black and white microphotographs of fungi, 11 x 8.5 cm (half page) in size, many of them being of good quality. Unfortunately, the illustration isdefective in several ways. The magnification is not given. The numbering of the photographs, composed of the chapter number and the figure number within the chapter is complex and unconvenient to the reader. Furthermore from a count of 219 numbered figures, 52 are composed of 2 or 3 separate full-size photographs, not mounted in plate, but printed on distinct pages most often and provided with a full legend under fig. -A which is said "continued" under fig. -B and fig. -C. This constitutes another unconve-~ nience. The procedure is in most case unjustified. Either the photographs composing a figure show the same fungus at low and high magnification, the one at low magnification being generally of no use or no value; or the photographs represent different fungi and could have been numbered and legended separately. At the same, the duplication of the same or similar pictures of the same fungus, illustrating here a term and further a gene- ric concept might have been avoided. For instance, a picture of Drechs- lera sp. fig. 1.1 is the inverted photograph fig. 4.46 of the same. Why Trichothecium roseum needs to be shown at different focusing in figures 4.105A and 4.105B on p. 303 and 304? Why Syncephalastrum ramosum must 284 appear ‘in 5 figures (1.9A, 1.9B, 4.120A, 4.120B, 4.121) and on distinct pages? Unacceptable is the fact that Fxophiala spinifera is not only figured three times in figs.1.27, 4.13 and 4.50 but also duplicated in fig. 4.49A, identical to; fig. 4.13, under the name Exophiala jeanselmei. It will be also mentioned that amongst the 161 figs. illustrating chap- ters 1 and 4, 50 figs. are of unidentified species of filamentous fungi. Why, in such an handbook designed to be informative as much as exampla- tive, the author choose to illustrate unidentified isolates rather than properly identified fungal species of medical importance which are avai- lable in pure culture in official culture collections? Why finally the photographs are supplemented of such poor and inacurate line drawings? Other defects are in the text. The author decided not to cite the authors of the names of the fungi in the text, but did cite them, in Appendic C, only for the correct names of which he listed a synonym but not for many other names used in the text. The reader will have to consult other sources to distinguish for instance Blastomyces dermatiti- dis (conidial Ajellomyces dermatitidis) from Wangiella (Hormiscium, Phialophora, Exophiala) dermatitidis. At the exception of the references inserted in the comments fol- lowing the generic descriptions of the fungi, references to the litera- ture are totaly absent from the general text, like are the references to published sources for the 77 culture media cited. The reader need to be informed on the source of every statement, term, definition, classi- fication, key, technique or formila other than the author's personal ones. : : ; ; i Dealing with terminology in chapter 1, the author restrict the application of the term chlamydospore to those terminal inflated thick- walled cells which are able to germinate and be reproductive and pro- poses the use of the term vesicle for the not germinating terminal chlamydospores as well for the intercallary ones. This proposition should be disrecommanded, the term vesicle being currently designating the ampulliform tip of the Aspergillus conidiophore stipe. Further the author illustratesthe term favic chandelier, in use in medical mycology, by a photograph of the appressorial hyphae in Botrytis cinerea. AS soon as 1870, J. KLEIN introduced the term appressorium to designate that type of hypha differentiated for the attachment of the fungus. There is no reason for abandoning that term. A-final) remark as néededvto infimum the, author s statement ti la— gellate cells are produced only by the chytrids" on page 1 to 2. Published by Academic Press, this handbook takes benefit of a high quality typographic printing and binding, as it will surely be in the hands of many mycologists in medical centers and laboratories. FUNGAL, BIOTECHNOLOGY, by J.&. SMITH, D:R. BERRY & BS KRIST IANSEN, x #308 ips, Ul. .68 ff hard cover, 419808 The British*Mycolozical Society, Symposium series n° 3. Academic Press, 111 Fifth Avenue, NewYork, NY10003..°) Price $326.50: This book is the proceedings of a joint symposium of the British Mycological Society and the Society of Chemical Insdustry, held in Glas- cow in September 1978. The recent developments of the fungal biotechno- logy has incited more research on the uses of fungi in the industrial processes. The book is a selection of topics related to the inoculation the growth and the exploitation of the fungi through the operation of liquid and solid state fermenters. The parameters of fungal growth, from the preparation of the inoculum, the genetical hazards, the rheological conditions to the reactions to the type of fermenter and the nature of the developed particles, are largely investigated and discussed for their af BOD interference in the design of the fermenters. The production of edible fungi is also considered. The potentialities of fungal biotechnology are reviewed and the achievements in different regions of the world also described. Although the repeated orthographic errors in the names of the fungi, this book remains a very interesting account to applied mycology. FUNGAL SAPROPHYTISM by Harry J. HUDSON, Studies in Biology n° 32, 2d ed., 76 p., ill., paper back, 1980. Edward Arnold Publ. see Bedford Square, London WC1B 3DQ. Price £ 2.25. It is good to realize the author's ability to reach so many goals through these modest 70 pages: enhancing curiosity and interest for the fungi, considering many aspects of the fungal ecology, guiding the stu- dent on the field and in the laboratory and demonstrating the practical interest of the fungi in industry and human life. Some titles: wood de- cay, blue stain, basidiomycetes and microfungi in litter, chitinolytic fungi, lignine decomposition, pyrophilous fungi, coprophilous fungi, moisture requirements, osmophilous fungi, storage fungi, xerophytes, aw quatic fungi, water pollution, thermophilous fungi, composts, fermented foods, antibiotics, vitamins. An exciting introduction to mycology, through ecology. FUNGI, MAN AND HIS ENVIRONMENT, by R.C. COOKE, 144 p., 111., paperaback ./1L9/7/7. Longman. Group, Ltd, London. Price £3.45. "T have attempted to write a book about fungi rather than a text- book on fungi''. The author indeed realizes a similar approach to that of Houston, but with many more information and considering also the parasitic fungi, appealing the awareness of people on the danger and value of the fungi in the present and future world. Fror® the pests and mycoses to their control, from the degadation of organic matter to the synthesis of protein and the cultivation of mushrooms for foods and drugs, the author clearly demonstrates the multiple both destructive and elabo- rative potentialities of the fungi, in which, finally, man is concerned. With science and philosophy, he suscites the desire of reading more on fungi. DIE BLATTERPILZE (AGARICACEAE) DEUTSCHLANDS UND DER ANGRENZENDEN LANDER BESONDERS OESTERREICHS UND DER SCHWEIZ, par Adalbert RICKEN ShUy +4462 p.,el22 pl. cols, im 8 ¢ relies heipzi¢gyl 9152 Reimprime par Dr. Massimo CANDUSSO, Viale Europa 5, 21047 Saronno (Varese) italia. Disponible. dans lL"allemand orzreinal: (Lit. 98,000:— net) om traduLt envitalven par Dr. Lazzari (Lit., 82,000)— net): La réimpression de 1l'oeuvre mycologique originale et unique de Adabert RICKEN (1851-1921) est un événement heureux. Publiée et distri- buée par livraisons successives de 32 pages et 8 planches colorées de 1910 4 1915, l'oeuvre de RICKEN fut trés bien accueillie par le monde mycologique de 1'époque d'aprés guerre . En effet 1l'oeuvre est le bilan de quarante année de récoltes mycologiques (1870-1910) en Allemagne et régions limitrophes et portant sur plus de 1500 espéces. L'ouvrage pré- sente dans un ordre systématique la description de 14£2 espéces d'acarics sur la base d'observations précises tant microscopiques que macroscopi- ques, avec mention de l'habitat et, pour 662 espéces, une illustration 286 en couleurs reproduisant les peintures a l'huile que réalisa 1l'auteur sur le frais. La nomenclature est traditionelle mais étayée des dates de publication. La derniére livraison comporta outre 1l'introduction et les index alphabétiques trois clés dichotomiques des genres, fort bien construites et basées sur les caractéres les plus évidents. La réimpression entreprise par 1'Imprimeur mycologue, Dr. Massimo CANDUSSO est particuliérement réussie. Le texte est reproduit de maniére réguliére sur du papier de qualité. Les planches en couleurs, éditées sur un carton plus fort comme les originales, reproduisent les tons les plus stables des différents exemplaires originaux comparés. La re- liure de type ancien est préservée dans une gaine. La qualité de cette réimpression mérite les encouragements du plus grands nombre de myco- logues. ICONOGRAPHIA MYCOLOGICA par Giacomo BRESADOLA, avec J.B. Traverso, L. Fenaroli. CG. Catoni® & J.B. Trenervediteurs, 26evols. 256.'pe., 1250 pl. col., Milan 1927-1933. Réimpression par Dr. Massimo CANDUSSO, Gruppo Micologico G. Bresadola, Viale Europa 5, 210477 Saronnes (Varese) Ltalia, in 5.volse,)l2b0rph.. col were lie, Sept. 1981-Déc. 1982. Prix souscription Lit. 100.000 before September 19895 Lit-100 S000 vamsrecumdese vols elie? sandy ale solde Lit. 60-85.000 au recu du 4e vol. L'examen de la planche proposée par 1'imprimeur CANDUSSO au Ssous- cripteur semble présenter toutes les garanties de qualité que l'on peut espérer, voire exiger, d'une telle réimpression. L'ouvrage de Bresadola est une oeuvre majeure dans la connaissance des champignons. La réimpres-— sion ne sera réalisée que si la souscription est suffisante. On ne peut que recommander aux mycologues et aux institutions de science mycologique l'acquisition d'un ouvrage que les générations futures de mycologues a travers le monde ne pourront ne pas connaitre ni ne pas consulter. Also nrecertved: LABORATORY GUIDE FOR IDENTIFICATION OF PLANT PATHOGENIC BACTERIA par N.W. SCHAAD , 72 p. 4 pl. col., mimeographed,4° in folder, 1980. 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(When and if it ever becomes necessary to sell your set of MY- POTAXON, (you should’ be aware that a complete run is far more valuable than a broken run.) This publication is available in microform. University Microfilms International 300 North Zeeb Road Dept. PR. Ann Arbor, Mi. 48106 USA. 30-32 Mortimer Street DéepealR: London WIN 7RA England - ? “ i rf i \ 2 ' j te 4 " q on , 7 Pk . ¥, ae ae | : y , t ; ‘ , 1 4 nly » Pern i ~ t ro MYCOTAXON Roiemxitl, NO. 2, pp. 209-350 July-September 1981 THE GENUS PILOPHORUS H. MARTIN JAHNS Botantsches Institut der Universitat Frankfurt, Germany Summary All species of Ptlophorus are examined and four species are ex- cluded from the genus. The remaining 10 species are described and their geographical distribution is demonstrated on maps. Character- istics used for the separation of species are discussed and an artificial key is given. Introduction Formerly the genus Ptlophorus was placed in the Clado- niaceae (Zahlbruckner, 1926), while today the genus is thought to be related to the genus Stereocaulon (Jahns, 1970a; Henssen and Jahns, 1973). In an earlier paper I attempted to delimit some species within the genus (Jahns, 1970b), but the choice of the species investigated at that time was arbitrary, resulting from the context of my onto- genetic work, so that no claim for completeness could be made. Furthermore, the investigations of most species were based on relatively limited material, so that conclusions about variation of characteristics and geographical dis- tribution were not reliable. For these reasons it seemed desirable to complete the earlier observations and to ex- tend the work to all species described by other authors and at the same time to include more specimens. This is the aim of the present work. Most of the specimens avail- able from the major herbaria were used in this investi- gation (more than 1000 specimens). A complete list of their geographical distribution is not included as it would ask to much space. For all practical purposes it seemed to be sufficient to characterize the occurring species, to mention their most important synonyms, to distinguish them from each other and to demonstrate their Gistribution on maps. The exact places of origin are given for specimens found outside the main area of dis- tribution. For a complete list of literature and synonyms see Zahnlbruckner (1927b, 1939). ; Material and methods The material investigated was received on loan from the following herbaria: Herbarium University of Michigan, Ann Arbor (MICH); Herba- rium of the University of Georgia, Athens (GA); University of Texas Herbarium, Austin (TEX); Universitetets Botaniske Museum, Bergen (BG); Herbarium University of California, Berkeley (UC); Botanisches 290 Museum, Berlin (B); Botanisches Institut, Bern (BERN); Museum of Natu- ral History, Budapest (BP); Farlow Herbarium of Cryptogamic Botany, Harvard University, Cambridge (FH); Commonwealth Scientific & Indu- Sstrial Research Organization, Division of Plant Industry, Canberra (CANB) ; Chicago Natural History Museum, Chicago (F); University of Alaska, College (ALA); Botanical Museum and Herbarium, Copenhagen (C); National Botanic Gardens, Dublin (DUB); Forschungsinstitut und Natur- museum Senckenberg, Frankfurt (FR); Conservatoire et Jardin botani- ques, Geneva (G); Botaniska Tradgard, Gdteborg (GB); Department of Botany, Dalhousie University, Halifax (DAL); Botanical Museum, Helsin- ki (H); Botanical Institute of Hiroshima University, Hiroshima (HIRI) ; Botanisches Institut der Universitat, Kiel (KIEL); Rijksherbarium, Leiden (L); Herbarium of the Department of Systematics and Plant Geo- graphy of the Botanical Institute of the Academy of Sciences of the USSR, Leningrad (LE); Institute of Botany, Faculty of Science, Lisboa (LISU) ; British Museum, Natural History, London (BM); Botanical Museum and Herbarium, Lund (LD); Herbarium of the University of Wis- consin, Madison (WIS); Botanisches Institut, Marburg (MB); Herbarium A. Henssen, Marburg (Hsn); Botanische Staatssammlung, Mtinchen (M); Herbarium of Yale University, Osborn Botanical Laboratory, New Haven (YU) ; New York Botanical Garden, New York (NY); Hattori Botanical La- boratory, Nichinan (NICH); Botanisk Museum, Oslo (0); National Museum of Canada, Natural History Branch, Ottawa (CAN); Oulun Museo, Oulu (OULU); Oxto Botanico, Padova (PAD); Muséum National d'istoire Natu- relle, Laboratoire de Cryptogamie, Paris (PC); Botanical Department of the National Museum, Praha (PR); Division of Botany, National Her- barium, Pretoria (PRE); University of Washington, Herbarium, Botany Department, Seattle (WZU); Botanical Department, Naturhistoriska Riksmuseum, Stockholm (S); Botanical Department of Tromsoe Museum, Tromsoe (TROM); Botanical Department, Museum of the Royal Norwegian Society for Science and Letters, Trondheim (TRH); Botanical Institute of the University, Turku (TUR); Institute of Systematic Botany, Uni- versity of Uppsala, Uppsala (UPS); US National Museum, Department of Botany, Washington (US); Naturhistorisches Museum, Wien (W); Botani- scher Garten und Museum der Universitat, Ztirich (Z). The specimens were mainly investigated with a stereo-microscope and some typical material was cut with a freeze-microtome. The genus Pilophorus and its subdivision In my earlier Work I explained in detail why in my Opinion the correct name of the genus and the correct epithet are “Pilophorue .Th.Fr.". Further discussion of this question followed from Culberson (1970), who favored the name " Ptlophoron (Tuck.)Th.Fr." and from Hawksworth, James and Laundon (1972), who agreed with my interpreta- tion. Closely related to this problem is the question of the type species, which in my opinion should remain P. robustus Tn. Fr.4 (Jahns? 1970p). In the last hundred years not only has the number of species subsummized under this genus fluctuated conside- rably, but attempts have also been made to subdivide the genus. An extreme proposal was the classification of Tuckerman (1882) who considered P. cereolus to be the only species of the genus - designating all other species earlier described as subspecies of P. cereolus. Stitzenberger (1862) included the genus Helocarpon Th.Fr. Pasi as a subgenus in the genus Ptlophorus. He referred implicit- ly to the description of Th. Fries (1861), who mentions for Helocarpon crasstpes short black, stalked fruitbodies. Fries himself obviously did not consider a relation bet- ween the two genera described - Ptlophorus and dHelocarpon - but later declared Helocarpon to be a synonym for Leetdea fits Fries,) 10/74). Having examined the type material of Lectdea crasstpes, I can confirm that there is no anatomical or morphological similarity with any species of Ptlophorus. A different subdivision of the genus was proposed by Satd6 (1940). When giving the description of the new spe- cies P. ntgrtcaule he believed there were principal diffe- rences between this species and all other representatives of the genus. He therefore suggested a division of the genus into a section Mgrtcaule, including P. ntgricaule, and a section E£uptlophorus, including all remaining species. It was explained earlier (Jahns, 1970a) that this subdi- vision cannot be accepted, as Sat6 based his description Simones new Section on a, far too rigid partition of, tissues pHowoOn pagmentation of the stalk tissue which, with varia- ble intensity, can be found in other species too. On the whole there seems to be no justification for any subdivision of the genus. The importance of different characteristics for the separation of species For the separation of the species the same criteria were used as described in the first paper dealing with part of the genus (Jahns, 1970b).. However, it is now pos- Sible to give a more accurate definition of the range of variation and to judge the value of the different charac- berigstics more rigidly. Thallus horizontalis The thallus horizontalis is always crustose and is ge- nerally grey or grey-green in color when dry. It consists Sreipawutve granules (fig. 1-4) or of flat or inflated scales (fig. 8). Small granules or scales may be pressed together, forming distinct areolae (fig. 7). In some spe- cies, especially in older material, the thallus horizon- talis may be almost absent. The shape of the horizontal thallus is clearly specific and relatively constant for each species. Therefore it may be used as a distinguishing eaeracveristic, but on its own it is not sufficient for the separation of the species. Principally the external morphology may be characteristic for each species but a certain degree of variation in shape is induced by factors like substrate, etc. Thus intermediate forms and transi- tions between the habits of different species may occur. Furthermore, variations in shape of the horizontal thallus among different species are difficult to explain, so that this characteristic is of restricted value for use in ertificial keys. Zoe Thallus verticalis and growth form The thallus verticalis of Ptlophorus is a pseudopodetium similar to the one found in the genus Stereocauton. The po- sition of the apothecia is always terminal. The pseudopo- detia may be simple or branched. Their size varies within ~ certain limits but offers a useful “aid for the description of the species. In nearly all species a few fertile speci- mens occur where pseudopodetia are absent and the apothe- cia are sessile on the thallus horizontalis. The habit of the pseudopodetia offers one of the best means of distinc- tion for taxonomic use. Most common are simple, unbranched and pin-shaped pseudopodetia, which occur in P. cereolus, P. strumaticus, P. nigricaule, P. fibula and P. ecurtulum (fig. 3,6, 7,8,10). If the pseudopodetia of these species bear more than one fruitbody, an indistinct terminal branching may be simulated. In contrast to these pin-shaped species, P. robustus and P. actcularts have high, branched pseudopode- tia which grow in dense clusters (fig. 1,9). An interme- diate growth form between these two types is represented by &. vegae, P. awasthianum and Py. clavatue.” Their pseudopode— tia are of middle height and somewhat branched and, at least in P. vegae and P. awasthtanum, may grow in more or less distinct clusters Gligce 2,11). Quite tyoica nm toceau species is their manner of branching, which is umbellate in P. robustus (fig .9), and Torked=in Ps acreular1s, =) = veqac and P. awasthtanum. The pseudopodetia are usually covered with an algal layer which is not continuous and often resembles the gra- nules and areolae of the thallus horizontalis. Older stalks may be partially bare. In P. ntgrtcaule the stalk is covered with conspicuous peltate scales, which are closely pressed together (fig. 8,17). Soredia In all species of Ptlophorus the pseudopodetia are cover- ed with an algal layer which is never continuous but com- posed of more or less dispersed areolae. A true cortex is only poorly developed and usually completely absent, so that the pseudopodetia in some species have a granular appearance. True soredia only occur in P. cereolus, where they give a distanet powdery Took to the stalk™ (1727) 5). Cephalodia Cephalodia are characteristic for all species of the genus Ptlophorus. However, two species which have been in- cluded in this genus (P. conglomeratum and P. stauffert ) lack cephalodia. This, among other reasons, justifies in my opinion the exclusion of these species from the genus Ptlophorus. The cephalodia of Ptlophorus are sacculate and exhibit no special wall structure. In the three tallest species, P. robustus, P. actcularts and P. vegae, some cephalodia occur on the pseudopodetia (fig. 1,9,11), but in all other species they are only to be found: on the thallus horizontalis. Their shape is irregular, the surface smooth or granular, and they are light brown to deep brown-black in color. nS The cephalodia either contain algae of the Nostoe or of the Sttgonema type. In the species investigated I found Stt- Goneia An Pe strumaticus, P. fibula, P: curtulunm -and P.-ntigrtcaule, while in P. robustus and P. awasthtanum I found only Wostoe. In P. aeteularts I observed only Wostoe, but Kurokawa and phibuichi (1970). found Stigorema an this’ species. This con- Gradi ction may not result from a misobservation, because in P. cereolus, P. clavatus and P. vegae I found some cephalo- dia with Sttgonema while others contained Wostoc . However, in general, all cephalodia of one specimen contain the same alga. Exceptions are rare. Cephalodia of P. vegae con- tain Wostoe and Sttgonema and possibly algae of the Gloeo- capsa type as well. Nostoe bearing cephalodia often have a habit distinct from those with Sttgonema. They are always characterized by a smooth surface, while cephalodia with Sttgonema as phyco- bionts show a granular surface, at least in their younger stages of development. This phenomenon is explained by the development of the cephalodium which was described else- where (Jahns, 1972). It is not impossible that from those species which seem to contain only one algal species in their cephalodia, specimens with other phycobionts will be found in the fu- ture. The occurrence of specific blue-green algae in the cephalodia is therefore not a useful taxonomic characte- sou en wal one Pycnidia, ascospores, fruitbody development Pyenidia occur abundantly in all species of the genus Ptlophorus. They are either located on sterile pseudopodetia Oreonnine thallus norizontealis.= The conidiophores: are un- branched and the conidia sickle-shaped. The size of the ecoriidia does not differ significantly in the different Species, so that this characteristic is unimportant for the taxonomy of the genus. The same holds for the ellip- Soid ascospores. The development of the fruitbodies is very uniform too (Jahns, 1970a). Some specimens of P. clavatus form an ex- ception. Numerous young apothecial primordia are located maebundles®on short stalks) (fie. 4)0 In the further course of development the primordia fuse and, with the growth of the generative tissue, form the typical elongat- ed fruitbody of the species (fig. 5,16). Anatomy and morphology of the apothecium The outer shape and inner structure of the apothecium belong to the best characteristics for the separation of the species. In most species the mature apothecia are al- most globose and quite often several apothecia are late- mallyitused) (fies 19510). An exception as “the, fruitbody of P. elavatus, where the hymenium extends a considerable distance down the pseudopodetial stalk so that the apothe- cium assumes a triangular outline in longitudinal section rine 5 :16)e. SmallerGvariations. of themspherical structure are found in P. aqwasthtanwn and P. ntgrteaule, which'both tend towards slightly triangular-shaped fruitbodies (fig. 2,8). 294 However, it is possible that the samples of P. awasthtanum that I. examined were not fully developed and mature. A longitudinal section reveals that the spherical shape of thesavpothecia 1s, noteunitrormly Lormed.wApariet yona., cue globose apothecia, fruitbodies may be flattened with a margin of the hymenium that may hang down for a considera- ble distance before curving upwards again to fuse with the Sia Tie es Po Anatomically, three major characteristics are of im- portance: the boundary-texture, the pigment-boundary and the columella. The boundary between the vegetative tissue and-ithe generative tissue of the fruitbody,) which is always easily recognized, may follow different patterns. It forms a thin horizontal zone in the lower periphery of thé. fruitbody animost species! (rigk 1) whereas int ee: robustus and P. struwnmaticus the vegetative tissue extends upwards into the generative tissue of the apothecium (fig. 12) forming a cone-like columella. The boundary between the vegetative and generative tis- sues may be emphasized by two conspicuous structures. In the transition zone the generative tissue may form a boun- dary-texture built from closely interwoven, short-celled hyphae (fig. 13). Often the junction of vegetative and generative tissue is characterized by a deposit of black- brown pigment, (fic 15), which is »referred to,aseche pigment-boundary. The pigment-boundary bears little rela- tionship to the boundary texture andmay or may not be superimposed. Black pigment is often concentrated in the subhymenial layers (fig. 14,15) as well as in the pseudo- podetial tissue (for example in P. ntgricaule, P. vegae and. P. strumaticus). The columella, the boundary-texture and the pigment- boundary are characteristics which help to separate the species very accurately. These characteristics may occur SiIngely or combamed. Secondary products Small amounts of several specimens were investigated by TLC. The results obtained agree in general with those pub- lished by Krog (1968) and Kurokawa and Shibuichi (1970). Alls species contain atrancrineand zeorin. Stictic acidiwas found in P. ntgrtcaule and P. vegae. Furthermore, Kurokawa and shibttachi found, caperatic acid: in 2. elavatus, sand scon- SsticticVacid’ in’ P. ntgrieaule cas’ well as* six other non- identified substances in different species of the genus. These observations could not be substantiated in detail. In this study theschemical characteristics will now be considered for taxonomy, as obviously only those substan- eés that are common to all species occur in sufficient amounts. In order to avoid seriously damaging the usually limited material of the herbarial specimens it is some- times impossible to demonstrate the presence even of atra- norin and zeorin. Therefore, it is almost impossible to obtain an exact and reliable picture of the distribution of these substances with the methods described. 295 Substrate The species of this genus grow exclusively on silicate stone. As an exception single colonies may extend to rig- idly pressed soil covering the stone. One specimen of P. acteularits from USA, Washington (near Lake Cushman, Mason Co., S.East Olympias (F)) was found growing on dead wood. The nu- merous other specimens from the same region suggest ex- tremely favorable climatic conditions which enable the lichen to colonize the unfamiliar substrate. Distribution The species of the genus Ptlophorus occur exclusively in the northern hemisphere. P. conglomeratum, P. colensot and P. stauffert described from Australia, New Zealand and New Caledonia are no longer included in the genus Ptlophorus. In the literature it is still maintained that P. actcularts eccurs invAustraliavand Africa, (Cape of Good) Hope). These statements refer exclusively to a note from Nylander (1860) who claims to have seen the material in the Hooker herbarium, but these specimens disappeared later. In the year 1860 the separation of the species was not well de- fined, so that it may well have been P. conglomeratum. In any Case consider i this: information’ too doubtful to an- elude in- the distribution maps. The distribution maps of the individual species are based almost exclusively on specimens which I have person- ally investigated. Only ina few cases have I included notes from other workers (Krog, 1968; Ammann and Ammann, O69; Kurokawarsand Shibuichd,,. 1970), as Ll am certain that the definition of species by these authors agrees with my own views. fer indavidual points’ inv the’ distribucion maps often include material from a relatively large area. The number of specimens examined is given for each point. However, in many cases these numbers are almost certainly slightly higher than the number of findings proper, as it was im- possible always to recognize identical specimens from the same gathering which had been distributed and preserved in different herbaria. In those cases where a single finding is represented by numerous specimens distributed as an exsiccate it is only recorded as a single point. whe exaeveplaces of origioneare indiecated@in détaLny only when either very few such places are known or when those places are outside the main area of distribution. Places which are listed in the description of the species are marked by a point, others not individually listed are marked by a square on the maps. Findings taken from other authors are marked by a triangle. In general the following large-scale distribution was established: One group of species ( P. actcularts, P. ntgrt- caule, P. elavatus) occurs around the northern Pacific Ocean. In the same area two more species are found which, how- ever, are restricted to only part of this area (P. curtulum in China and Japan, P. vegae in Alaska and the Bering Strait area). One species (P. awasthitanum ) is restricted to the Himalayas and another (P. ftbula ) to the eastern 26 parts of the USA. One species ( P.. strumattcus) occurs only in those parts of Europe which have an oceanic climate. Very typical is the distribution of P. robustus, which is found circumpolarly in the most northern areas and in high mountains. P. cereolus shows the most irregular di- stribuuion. [hie, species as most abundante ire ocandinavia but occurs in single specimens in middle and central Kurope, if Siberia and an the eastern part of North Ame— rVCa. Artificial key to the species la 1b 2a 2b 3a 3b 4a Ab 5a 5b 6a 6b 7a 7b 8a 8b 9a 9b 10a Lichen with pseudopodetia and developed apothecia 2 Lichen. without pseudopodetia or pseudopodetia without apo- thecia, sometimes with pycnidia or black primordia of apo- thecia. 15 Thallus horizontalis and pseudopodetia clearly sorediate. Pseudopodetia pin-shaped, unbranched, at most 1 cm high. Apothecia without columella (fig. 3). P. cereolus Surface not sorediate, at the most slightly granulose. 3 Pseudopodetia with continuous white cortex. Ecorticate parts of the pseudopodetia and their central part always black. Pseudopodetia 0.5=-1.5 cm high, usually dichoto-— mously branched. Large cephalodia on short branches. Apo- thecia unknown. Alaska, Bering Strait. (fig. 11) P. vegae Lichen with other characteristics. 4 Pseudopodetia smaller than 5 mm. 5 Pseudopodetia higher than 5 mm. it Apothecia cylindrical or triangular in longitudinal section, at least twice as high as broad (fig. 16). Pseudopodetia pin-shaped, unbranched. Apothecia often slightly bifurcate atethe apices (higs.5) . P. elavatus Apothecia globose, sometimes several globose apothecia are fused. 6 Pseudopodetia very thin (0.2-1.0 mm broad, 5 mm high), more or less branched, partly ecorticate and black (fig. 2). Himalayas. P. awasthtanum Pseudopodetia broader, pin-shaped, unbranched. q; Scales of the thallus horizontalis and of the pseudopo- detial cortex peltate (fig. 17), white, pressed together, with an inflated appearance. Inner parts of the pseudo- podetium with dark pigment. Apothecia with boundary-texture, globose or sometimes pointed (fig. 8). P. ntgrtcaule Lichen with other characteristics. 8 Thallus horizontalis formed by granules or minute scales. Granules or scales fused into broader, closely arranged areolae (fig. 6,7). £ Granules or scales of the thallus horizontalis scattered. 10 Apothecia always with boundary-texture and pigment-—boun- dary. Pseudopodetia usually not higher than 1 mm (at most 2 mm) (fig. 7). Eastern North America. P. ftbula Apothecia always with boundary-texture, only very old specimens with pigment-boundary. Pseudopodetia 1-3 mm high (fig. 6). Japan and China. P. curtulum Apothecia without columella, with boundary-texture and in old specimens with pigment-boundary (fig. 6). Japan and China. P. curtulumn 10b lia i aliss 12a 12b 13a 13b 14a 14b 15a i Se 16a 16b 17a 17b 18a 18b 19a 19b 20a 20b 2la 21b 22a 22b Apothecia with columella, without boundary-texture and pigment-boundary (fig. 10,12). Europe. P. strumattcus Apothecia cylindrical or triangular in longitudinal section, at least twice as high as broad (fig. 16). Pseu- dopodetia pin-shaped, unbranched. Apothecia often slight- ly bifurcate at the apices (fig. 5). Peeelavacus Lichen with other characteristics. Scales of the thallus horizontalis and of the pseudopo- .detial cortex peltate (fig. 17), white, pressed together, with an inflated appearance. Inner parts of the pseudopo- detium with dark pigment. Apothecia with boundary- texture, globose or sometimes pointed (fig. 8). P. ntgrtcaule Lichen with other characteristics. Pseudopodetia very thin (0.2-1.0 mm broad, 5-15 mm high), more or less branched, partly ecorticate and black (fi¢. 2). Himalayas. P. awasthtanum Lichen with other characteristics. Pseudopodetia umbellately branched towards the apices. Apothecia numerous, often fused at the margins, with large columella (fig. 9,12). P. robustus Pseudopodetia dichotomously branched, in combed tufts, smaller plants unbranched. Apothecia without columella ‘Getkeps Hays P. acteularts Pseudopodetia with pycnidia or with primordia of apothecia present. Only thallus horizontalis present. Thallus horizontalis and pseudopodetia sorediate. Pseu- dopodetia pin-like, often broadest in the middle Gale ENS . P. cereolus Lichen not sorediate. Pseudopodetia very thin (0.2-1.0 mm broad, 5-15 mm high), more or less branched, partly ecorticate and black (fig. 2). Himalayas. P. awasthtanum Pseudopodetia broader. Pseudopodetia with continuous white cortex. Ecorticate parts of the pseudopodetia and their central part always black. Pseudopodetia 0O.5-1.5 cm high, usually dichotomous- ly branched. Large cephalodia on short branches. Apothecia unknown (fig. 11). Alaska, Bering Strait. P. vegae Lichen with other characteristics. Thallus horizontalis persistent, formed by closely connected areolae. Sterile pseudopodetia 1-2 mm high, terminated by a pointed pycnidia. P. clavatus Thallus horizontalis formed by scattered granules, evanescent in older specimens. Pseudopodetia at least indistinctly umbellately branched in their upper part. With numerous pycnidia or primordia of apothecia. P. robustus Pseudopodetia slender, unbranched or dichotomously branched. P. actcularts Thallus horizontalis not sorediate, sometimes granular. Thallus horizontalis sorediate. P. cereolus Thallus horizontalis formed by white, peltate, inflated scales or granules. Japan, west coast of North America. P. ntgricaule Lichen with other characteristics. 12 155 14 16 4) iy 18 ue) 20 23 Zod. Boo 23a Thallus horizontalis persistent, formed by closely connected areolae. 24 23b Thallus horizontalis persistent or evanescent, formed by more or less scattered granules or scales. Determi- nation only from distribution (see maps). P. robustus, P. strumaticus, P. acteularts, P. vegae, P. awasthtanum, P. curtulum 24a Areolae thin, not distinctly formed by individual granules. P. clavatus 246 Areolae thick, composed of small granules. ao 25a Distribution in Japan and China. P. curtulum 25b Distribution in-the eastern North America. P. ftbula Description of the species Pilophorus acicularis (Ach.) Th.Fr., De Stereocaultis et Ptlophoris Comm. + “41° 41857); Baeomyces actcularts Ach., Meth. Lich.: 328 (1803). Type collection: North America, Hall (UPS, isotype). Cenomyce acteularts Ach., Lich. Univ.: 567 (1810). Thallus horizontalis small granular, evanescent in older thalli, grey when dry. Pseudopodetia (0.5-) 3 cm high, 1 mm diameter. Mostly Simple or dichotomously branched, stalks curved so as to appear as if combed; rarely erect, pin-like and 1 cm high (fig. 1). In some speci- mens richly branched in the upper part of the pseudopodetia, thus Ne ‘ os 70 ix ALB. , ‘a wva'? WbED: wpe cu) fig. 1 P. aeteularis (ce - cephalodium) Z99 faintly resembling P. robustus. Internally the pseudopodetia are solid when young, becoming hollow with age, composed of long, thin, strongly gelatinized hyphae with narrow lumina (0.5 um). The lower part of older pseudopodetia becomes blackened internally. Algal layer not continuous, in more or less distinct granules, entirely absent from some parts of the surface. Pycnidia occur in the tips of small sterile pseudopodetia or in the tips of small lateral branches of older pseudopodetia. Conidio- phores 30 pm long, unbranched, with terminal sickle-shaped conidia, 6exoi> pm. Apothecia abundant, one or several on the apices of the pseudopo- detia, black, hemispherical or indistinctly triangular, up to 1.5 mm diameter. There is no columella. Hymenium up to 240 pm high, two-thirds pigmented, lower part ste- rile, consisting only of paraphyses. Excipulum absent. Asci eight- spored. Spores rounded when young, becoming spindle-shaped when ma- pure, )21.0-29.5 x 4°75-5.5 pine Generative tissue formed¥of closely interwoven hyphae with short, broad cells with large lumina. The ge- nerative tissue is pigmented black-brown, the color being most in- tense below the paraphyses, becoming less so towards the stalk region. Phycobiont green, Pleurococcus type, cells c. 9 pam diameter. Cephalodia on the thallus horizontalis are brown and hemisphe- rical and contain Wostoc. Kurokawa and Shibuichi (1970) found cepha- lodia containing Sttgonema. Small cephalodia also occur on the pseu- dopodetia (fig. 1). Growing on silicate stone, seldom on decaying wood (one specimen). Remarks The correct citation for the species is P. actcularts Phcony yin. Fre (105 /p. 4ooe ands not P..aetcularis (Ach...) Ny 105) ep. 90) sae explained e€arlier (Jahns, 1970b); P. acteularts can be separated from the smaller species by its tall pseudopodetia. It may be confused with P. ro- bustus, especially in material from Alaska where both spe- cies occur together. But usually the type of branching is different (umbellate in P. robustus, dichotomous in P. acteularts) and the lack of a columella in longitudinal sections of the fruitbodies of P. acteularts always makes distinction easy. Th. Fries (1889, p.765) described P. actcularts var. econjugens Th.Fr. from Vancouver Island, which is said to resemble P. robustus. I have seen the original material and can see no difference from numerous other specimens which are not worthy of varietal status. Distribution (map 1) P. acteularts is probably the most abundant species of the genus. Most specimens were found on the west coast Of North: America, with a focal point in British ,Columbia and Washington. The species is widely spread in Japan too. In general this species seems to prefer an oceanic climate without extremely low temperatures, at least in comparison with other species of the genus. This assump- tion is supported by the fact that this species is found more southerly (44 Tindings in California) than all ‘other listed M in the description of the species) species and only two specimens were found in northern Alaska where, for example, P. robustus and P. vegae occur more often. East of the Rocky Mountains P. actecularts is rare too. It is very difficult .to give any estimate on the occurrence of the species in China. Although there are on- ly three places known to me where the species has been found, at least one gathering was large enough for the distribution of numerous specimens as an exsiccate (Krypt. Vind. Exs. 2842). Therefore, it may be suggested that P. acteularts is not rare in China, but simply has not been collected. Places outside the main distribution area (map 1, marked by aucirele): Canada, Ontario, Belleville (CAN). Pilophorus awasthianum Rads., Arch. Soc. Zool. Bot. Fenn. Vanamo 5:28-29 (1950). Type collection: India orientalis, Himalaya orientalis, Darjeeling, 3500 m (H, holotype). Thallus horizontalis persistent or evanescent, consisting of loosely scattered granules, c. O.1 mm diameter, grey when dry. Pseu- dopodetia simple or branched, small and very thin, O.5-1.5 cm high and 0.2-1.0 mm broad. Pseudopodetia partly covered by an algal layer consisting of small granules, but for the most part the stalks 301 fig. 2 P. awasthtanum (ce - cephalodium, p - pycnidium) are bare. The ecorticated parts are blackened, the algal layer grey. Internal part of pseudopodetia solid, formed by strongly gelatinized hyphae. Pycnidia apical on short pseudopodetia. Apothecia apical on mature pseudopodetia, sometimes simple and globose but mostly aggregated, consisting of several subglobose bo- dies (fig. 2). Apothecia 1.5-2.0 mm in diameter. In mature specimens apothecium and pseudopodetium are separated by a pigment-boundary, ~ while boundary-texture and columella are absent. Hymenium c. 100 pm high, excipulum absent. Asci eight-spored, spores spindle-shaped, c. 20 x 7 pm. Phycobiont green, Pleurococcus type, cells c. 10 pam in diameter. Cephalodia contain Nostoc, black, globose, sessile on the thallus horizontalis, c. O.1 mm diameter. Growing on silicate rock and earth. Remarks P. awasthtanum differs from all other species by the small, branched and very thin pseudopodetia and by the strongly aggregated apothecia. As there are only two S02 known specimens so far, the range of Variatvoneot ithe different characteristics is unknown. Distribution (map 2) The species is only known from the Himalayas. I have seen the following specimens: India, Darjeeling district, E. Himalayas, Awasthi, 1948 (H, holo- type); India Sandakhpu, Awasthi, 1950. (UBS). map 2 P. awasthtanum - distribution Pilophorus cereolus (Ach.) TH.Fr., Lich. Scand. 1, 1: 55 (1871). Ltehen cereolus Ach., Lich. Suec. Prodr.: 89 (1798); Meth. Lich.: 316 (1803). Type collection: Sweden, Facklelaf, E. Acharius (H, holotype). Thallus horizontalis persistent, consisting of loosely aggre- gated, sorediate granules, c. O.1 mm diameter, grey-green when dry. Pseudopodetia pin-like, usually up to 5 mm (occasionally up to 1 cm) high, 1.5 mm broad, sorediate. Sterile stalks or those 503 fig. 3 P. cereolus (ce - cephalodium, p - pycnidium) bearing only pycnidia are usually broadest in the middle and irregu- larly curved (fig. 3). In some specimens, especially in the few Ame- rican samples, the upper part of the pseudopodetium is slightly branched, each branch bearing a pycnidium or a primordium of a fruit- body. Internally the stalks are compact, composed of strongly gela- tinized hyphae with narrow lumina. The phycobiont layer is granular sorediate. Pycnidia bottle-shaped, abundant, several often produced at the apex of a pseudopodetium. Conidiophores c. 25 pm long, unbranched, with terminal conidia. Conidia sickle-shaped, 5 x 1 pm. Apothecia scarce, solitary or seldom two on one stalk, spherical, black and terminal, up to 1.5 mm diameter. In longitudinal sections the apothecium is spherical and without a columella. The boundary between the generative and vegetative tissue remains horizontal with the lower margin of the apothecium. In old apothecia a strongly co- lored pigment-boundary is situated at this juncture. An indistinct boundary-texture may sometimes be developed. Hymenium c. 80 pm high, almost unpigmented. Excipulum absent. Asci eight-spored. Spores rounded when young, becoming spindle-shaped when mature, 14.5-21 x 5.5-6.5 pm. The regions just below the paraphyses and in the pigment- boundary are deep black-brown, towards the center of the apothecium the color grades to pale brown. Phycobiont green, Pleurococcus type, cells c. 9 pm diameter. 304 Cephalodia brown, clustered,-sessile on the thallus horizontalis, c. 1-2 mm in diameter. Their surface is usually wrinkled. The cepha- lodia contain Nostoe or sometimes Sttgonema. Growing on silicate stone. Remarks P. cereolus is distinguished from all the other species by the sorediate thallus horizontalis and pseudopodetia. Without this characteristic fruiteng specimens Onn wcere— olus could besmistaken for PP, strwunaticus, Pay tbula,specureu— lum or P. ntgricaule, as all these species are of about the same height and pin-like in appearance. The pseudopodetia of P. vegae sometimes have a granular cortex, which could be considered as sorediate, but the habit of the species distanculy disiers. fromeP. cereolus (ing — jeond) 1.) An other important characteristic for the distinction of the small, pin-like species is the longitudinal section of the fruitboodiese( tre. ei 5 en Tuckerman, in his later works (e.g. Tuckerman, 1882, p.235) considered all other species to be varieties of P. cereolus. As a result, many subsequent authors have attributed all species of the genus to this species. As pointed out by James (1965, p.239)°"P. cereolus" as used map 3 P. cereolus - world distribution (places listed @ and not listed M in the description of the species) 305 by British authors refers mostly to P. strwmatreus and not to the present species. In the same way "P. cereolus" as used by many American authors refers to P. actcularts. Rasdnen (1946, p.2) described P. cereolus var. ecephalodt— ferve has, which he saidvas to be distinguished by the smaller pseudopodetia and the abundant, markedly tuvercu- late cephalodia. The syntype material of this variety was distributed in Lich. Fenn. Exs. as no. 960. The samples Pre an no Way diiferént from a multitude of other poorly- developed specimens and should be considered merely as ju- venile specimens which are not worthy of varietal status. map 4 BP. cereolus - distribution in Europe (places listed @ and not listed MM in the description of the species) 306 Distribution maemo) P aeneolus US thesmost abundant Species sor sence eciie aya Europe. Most samples were collected in Scandinavia where sts occurrence overlaps with the distribution of P. robustus i and P.tetmeatieus. However, P. .strumaricus: Usyrestricved) Ge | the oceanic climate of the Norwegian and Swedish west | coasts, and P. robustus occurs in the Norwegian highlands and in the Arctic only, whereas P. cereolus is spread over | on 15 20 reo) 7 bey) Ss CGT sth ROT map 5 P. cereolus - distribution in Scandinavia (places listed @ and not listed # in the description of the species) DOr, Scandinavia, including Finland. The absence of P. cereolus in Great Britain suggests that this species, unlike P. strumatteus, dislikes an oceanic climate. This agrees with the observations made by Ammann and Ammann (1969). Outside Scandinavia P. cereolus shows an interesting distribution (maps 3-4). These rare but widely-spread findings are listed further on. While a specimen from northern Poland links up reasonably well with Scandinavia, the occurrence in Siberia appears somewhat isolated. How- ever, the material from Siberia is well developed and was most certainly very abundant as samples of this material can be found in many herbaria. A somewhat bigger group of findings originates from the Carpates and the Tatra (Cze- choslovakia), partly distributed as an exsiccate (Lojka, rc. Huet Oo!) In the Alps this lichen is apparently far less abun- dant than might be expected from the older literature. Although I have not seen all material mentioned in older publications it seems that most samples belong to Stereo- caulon, especially Stereocaulon ptleatum, which is small and sorediate and looks somewhat similar to P. cereolus. Nume- rous specimens originate from the Oberpfalz in southern Germany. These and material from South Tyrol have been distributed as exsiccatae (Arnold, Lichenes exs. 823, 2008), but apparently it has been found only in a few places. The lichen seems to be absent from the rest of Germany and the more western countries. The literature does not give any clue to the distri- bution of P. cereolus in North America as P. cereolus, P. acteularts and P. ftbula were completely confused until re- Cenoly. In particular, P. actcularis was designated as, \P. cereolus in many cases. Krog (1968) remarks, that she has not seen any authentic specimen of P. cereolus from wes- tern North America. She mentions one single specimen which she has not seen herself, but which from the de- Scription could have been P. cereolus. I had the chance to see this sample and it proved to be P. aectcularts too. We May, therefore, conclude that P. cereolus does not occur in the western parts of North America. In the eastern part of the continent there are three places where true P. cereolus has been found. One of which (Grand Marais) was described by Fink (1910) who, however, thought it to be identical with the P. cereolus described by Tuckerman and which in fact was P. acteularis. The American material is not fully developed and slightly different from the Kuropean material. Nevertheless, it seems to be the same species and its distribution in North America is linked bo, the Mam vdistributionm area in Europe by findanes from Newfoundland, Greenland and the Azores. Places outside the main distribution area (map 3-5, marked by a circle): Canada, Newfoundland, upper Humber River (H); Canada, Newfoundland, New Harbour, Trinity Bay (M); Canada, Minnesota, Grand Marais, along the northern shore of Lake Superior (4 specimens MICH, 1 speci- men US); Canada, Minnesota, Cook County, 8 miles SW of Grand Marais 308 on US Route 61, Cascade River State Park, along Cascade River (CAN); USA, New York State, Adirondack Region, Chapel Pond, near St. Hu- berts (MICH); Greenland, Diskofjord, Kuanersuit suvdlat (C); Azores, Santa Maria, Pico Alto (James and Henssen unpublished) (MB); Germa- ny, Oberpfalz, above Spielberg near Nabeck west of Schwandorf (11 specimens BM, C, H, KIEL, M, O, PR); Germany, Oberpfalz, Schutzen- gelsteinbruch, Eldensteiner Forst (M); Austria, Tyrol, Silz, Intal (2 specimens M); Italy, South Tyrol, Margola near Predazzo (25 spe- cimens:B, BM, ‘BP, FH; H; LD, M, 0, S, W)? sitaly, South Tyrol, Set— ween Bellamonte and Paneveggio (4 specimens BM, BP, M, W); CSSR, Presov, 0. Szatala (BP; CSSR, alpes Dzurowa prope pagum Teplicska, com. Lipto (W); CSSR, Slovakia, Nizke Tatry, in valle rivi Svidovy (PR); CSSR, Slovakia, Nizke Tatry, in valle rivi Svarinsky (PR); CSSR, in sylva Stefkofka prope Bresztova, com Arva (16 specimens B, BG, “BPS iFH,. G) H,;) LD, 3M; 0, PR iS; Us (oN) eo Poland, sproperhabiau (H); USSR, Dudinka, Yenisey, lat. bor. 69°35! (7 specimens FH, H, OS US re | Pilophorus clavatus Th.Fr.,Bot. Notiser, 1888: 214 (1888). Type collection: Canada, British Columbia, Vancouver Island, Mt. Mark, 1887, Macoun (UPS, holotype). P. acteularts £. hallit Tack./Preoe,; Amt Acad ArtstSci. sl2-sl77 | (1877) . Type collection: USA, Oregon, Hall (UPS, isotype). | P. cereolus var. hallit (Tuck.) Tuck., Syn. N. American Lich.: 235 (1882). Py hatltit (Tack.) -Vain.;. BotzeMag., Pokyoe35=. 59n (1021 | P. japontecum Zahlbr. (nomen nudum), Catal. Lich. 4: 432 (1926); Bou, Mage, “Lokyoc41: 387i, G1927 je. fig. 4 P. elavatus, with young fruitbodies (ba - bush-like pri- mordia of apothecia, ce - cephalodium) af SOS Thallus horizontalis thin, granular persistent. Granules often aggregated, forming scales. Granules minute, O.1 mm diameter, grey- green when dry. Pseudopodetia up to 2 cm high, c. O.3 mm thick (fig. - 5). Internally the central area of the stalk is massive and consists of thin, strongly gelatinized hyphae with thin lumina (0.5 pm). Al- gal layer not continuous, formed by more or less indiscrete granules, the stalk usually being naked just below the apothecium. Central lower part of the stalks sometimes blackened. Pycnidia bottle-shaped, apical on short pseudopodetia, about 1 mm high, often bearing thalline scales which form a collar halfway up the stalk. Conidiophores unbranched, 25 pm long, with terminal coni- tay OO ox. O 5 At. Apothecia black, occurring Singly at the apex of each pseudopodetium, often spread- ing some distance down the stalk. The apothecia are tri- angular in longitudinal sec- eLOM eid. 16), about. 2 W=5) mm high, 1 mm broad at the upper end, tapering to O.4 mm at the base, often they are more or less bifurcated at the apices (fig. 5). Apothe- cium sometimes formed by fu- Sion of several primordia growing on bush-like branched pseudopodetia (fig. 4). No columella, boundary-texture, or pigment-boundary is formed between the vegetative and generative tissue. Hymenium c. 200 pm high. Asci eight- spored. Spores rounded when young, becoming spindle- Shaped when old, 23.5-26.5 x 5.0-6.0 pm. Hymenium green- ish, and the generative tis- sue uniformly pigmented, black- brown. Phycobiont green, Pleurococcus type, cells c. 8 pam diameter. Cephalodia brown-black, irregular, lumpy, about O.5 mm diameter with a velvety surface. Growing on the thallus horizontalis. They contain Sttgonema and sometimes Nostoc. Plants growing on silicate rock. figs 59h. elavacus Remarks In most specimens the triangular outline of the apo- Cheevalissurficienteto dietinguish Pa clavatussirom all other species. P. eclavatus could only possibly be confused with small specimens of P. aeteularts bearing slightly tri- angular apothecia. In this case the two species may be separated by their persistent or evanescent thallus hori- zontalis respectively. Tf was *explained, earlier (Jahns, #970b) thatr2. hall is a synonym for P. elavatus and that the name P. clavatus Den Th.Pr. (1608) has pricrity at the species Jever over.) halit4 . (Tuck.) Vain. (1921). PF. japontewn Zanibraelseaanomen nudum. Zahlbruckner knew it to be a synonym of P. halltt but nevertheless preferred his new name. Distribution (map 6) The distribution of -P.~clavatus 13) essentially 1denivical with the occurrence of P. acteularts. Both species are found on the coasts lot the northern Pactra Ocean, a sme ardcue apparently being relatively more abundant than P. aeteula- rts in Japan. i surprisingly Pwelavatus, nas siiou yeu bec found in Northern Alaska but was present on the Asiatic side of the Bering Strait. Kurokawa and Shibuichi (1970) have examined material from Formosa and their observation is included in map 6 (triangle), they also mention the species in Korea but don't seem to have examined the ma- terial themselves. fry eye ech MAA map 6 P. clavatus - distribution ( M® places not listed in the description of the species, aA specimens not examined) Pilophorus curtulum Kurok. and Shib., Journ. Jap Bot. 45, 3: 78 (1970). Type collection: Mt. Ontake, Prov. Hida, Japan (TNS, holotype; FH, O, isotypes). Thallus horizontalis persistent, consisting of minute granules, O.1 mm diameter, usually aggregated forming small squamules, 1.5 mm oe ————E—EeEE—————— ea. oe 4 S fig. 6 P. curtulum (ce - cephalodium) diameter (fig. 6). Sometimes granules scattered and hardly aggegat- ed. Thallus grey-green when dry. Pseudopodetia pin-like, 1-3 mm high, 0.5-1.0 mm broad. Mostly covered by a continuous algal layer consisting of small granules. Only sometimes ecorticate. Internally the stalks are compact, composed of strongly gelatinized hyphae. Central part of older pseudopodetia colored by deposition of dark pigment. Pycnidia sessile on the thallus horizontalis. Apothecia apical on the pseudopodetia, nearly always single, but sometimes two apothecia developed from two primordia have partly merged. Sometimes apothecia sessile on the thallus horizontalis. Apothecia about 1 mm in diameter. In longitudinal sections the apo- thecium is spherical and without columella. Pseudopodetium and apo- thecium are separated by a boundary-texture which is formed in the young primordium. In older specimens the boundary-texture is masked by a pigment-boundary. Hymenium c. 150 pm high, almost unpigmented. Excipulum absent, asci eight-spored. Spores rounded when young, becoming spindle-shaped when mature, c. 7 x 22 pm. Phycobiont green, Pleurococcus type. Cephalodia contain Sttgonema, they form dark-brown flattened discs between the squamules of the thallus horizontalis. Growing on silicate rock. lee Remarks , This species resembles P. ftbula. The thallus horizonta- lis of P. curtulum seems to be a bit thinner and less areo- lated, and the pseudopodetium is somewhat higher and more granulated than ain P.. frbula; but probably theyonLy ‘cood reason for keeping the two species separated is their dis- JUNCteGLStrrpution. Pe curtuium is only Known vomeapane ane China, while P. ftbula is endemic to north-east America. Distribution (map 7) This lichen has been found in Japan several times. I have seen samples from 10 places. From China I have seen two samples: Cleblieel, Vathewete, lens, creennel, (@)) 9 Claalinel, wAGheNeia, leenraevceulGl. 6 Ike imines Waha prope pagum Yungning regione frigide temperata infra casulam Maoniubi, alt. 3800-4030, Handel-Mazetti (Diar.Nr.1286) (W). OS BAC: map 7 P. curtulum - distribution (places listed e and not listed ™ in the description of the species) Pilophorus fibula (Tuck.) Th.Fr., De Stereocaults et Ptlophoris Comm.: 42 (1857). Stereocauton ftbula Tuck., Proc. Am. Acad. Arts Sci. 1: 238 (1847). Type collection: USA, New Hampshire, White Mts., 1844, Tucker- man (FH-Tuckerman, holotype). Thallus horizontalis squamulose, squamules large, up to 2 mm dia- of el ee meter, composed of small granules, persistent, grey-green. Pseudopo- detia seldom exceeding 1 mm hight, sometimes totally absent so that the apothecia are sessile (fig. 7). Stalk massive, composed of gela- tinized thin hyphae with small lumina (0.5 pm). Algal layer more or less areolate, not continuous. Pycnidia single, in small diminutive pseudopodetia. Conidiophores ce 30 pm long, unbranched, with terminal sickle-shaped conidia, 6 x 1 pm. Apothecia black, hemispherical, 1 mm in diameter. A pigment-—boun- dary and boundary-texture lies between the generative and vegetative tissues. There is no columella. Hymenium up to 120 pm high, lower- third pigmented. Lower edge of apothecium sterile, consisting only of paraphyses. Excipulum absent. Asci eight-spored. Spores rounded when young, becoming spindle-shaped when old, 17.0-23.0 x 5.5-6.5 pm. A dark-brown pigmented layer occurs under the paraphyses, which be- comes paler towards the center of the apothecium. Phycobiont green, Pleurococeus type, cells c. 6 pm diameter. Cephalodia contain Sttgonema, they form brown, Be Ee granular discs, c. 1 mm diameter between the scales of the thallus horizon- ee S ta ea ee Growing on silicate stone. 7 fig. 7 P. fibula (ce - cephalodium) Remarks Since the first description of Stereocaulon ftbula by Tuckerman in 1847 this specific epithet has tended to be applied to widely differing small or depauperate speci- mens of the genus. But P. strumattcus and P. cereolus differ distinctly from P. ftbula by the occurrence of a columella 314 in the apothecium or of soredia respectively. Most diffi- cult: to" distineulseh trom Py tbulatisw?. curren sea one differences between the two species are only gradual. The thallus norizontalis of 2, curtulio 2svusually bul noe always less squamulose and less persistent. Moreover, only very old apothecia of P. ecurtulum are separated from the stalk by a pigment-boundary in addition to the boundary-texture which is always present. Distribution (nao fo P. ftbula occurs only in north-east America and appears to be restricted mainly to the area of the White Mountains. Only two samples originate from the coast. I have seen ma- terial from *the following places: USA, New Hampshire, Mont. Alb. (BM, FH, M, MICH, W); USA, New Hamp- shire, Roche entrance to Ilume, Franconia (F, US); USA, New Hamp- shire, montibus Nord Anglio (PC); USA, New Hampshire, Warren (US); USA, New York, rock on trail up Mt. MacDetyse near Lake Placid (MICH) ; USA, New York, Adirondacks, Mt. Marcy (S); USA, New York, Adiron- dacks, Indian Falls, near Mt. Marcy, near Lake Placid (FH); USA, Maine, Mt. Desert Island (M, US). map 8 P. ftbula - distribution ( e places listed in the description) se epke Pilophorusenigricaule Sato, Journ. Jap. Bot. 16: 173)(1940) £1g.7. Type collection: Japan, Honsyt, Mt. Tydkai (Tokyo, holotype). Pr lopnorom cereolus (non Th.Fr.) Sat6, Journ... Jap; Bot. 9: 214° (1933). Thallus horizontalis persistent, white or light grey, granular. Granules c. 2 mm broad and c. 1 mm high, subglobose, aggregated or scattered on the substrate. Most granules slightly peltate. Pseudopo- detia pin-like, 1-6 mm high, 1 mm diameter. Internally the stalks are compact, composed of strongly gelatinized hyphae, colored black by the deposition of dark pigment-granules. The pseudopodetia are covered by subglobose granules, showing the same color, morphology endestructure as the granules of the thallus horizontalis (fig, 17) . In some specimens a few stalks are branched. fig. 8 P. ntgrtcaule (ce - cephalodium) Pycnidia apical on short pseudopodetia or sessile on the thallus ho- rizontalis. Conidiophores long, slightly branched with terminal Sickle-shaped conidia, c. 10 x 1 pm. Apothecia terminal on mature pseudopodetia or sometimes sessile on the thallus horizontalis, 1.0-2.5 mm diameter. Apothecial margin downturned as far as the point of attachment to the stalk. Apothe- cia subglobose or slightly conical (fig. 8). No columella is present. Apothecium and pseudopodetium separated by a broad boundary-texture. No pigment-boundary is present. Hymenium c. 180 pm high, subhymenium 120 pm. Excipulum absent. Asci eight-spored. Spores rounded when young, becoming spindle-shaped when mature, c. 18 x 7 pm. Phycobiont green, Pleurococcus type, cells c. 10 pm diameter. Cephalodia on the thallus horizontalis, thick, brown to black, 316 with wrinkled surface, c. O.5 mm diameter, containing Sttgonema. Growing on Silicate rock. Remarks P. ntgrtcaule is distinguished from other species by the white color and the thick, subglobose to peltate structure of the thallus horizontalis and of the algal layer of the pseudopodetium. With these characteristics the species is easily dstinguishéd: from Pi veurtubims whieh as aiecorshor, stalked and occurs in the same area. On the other hand, the dark color of the central part of the pSeudopodetium, from which the species takes its name, is not unique to the species. The stalks of P. vegae, P. awasthtanum, P. curtulum, P. strumatteus and P. eclavatus are also more or less blackened within. As Kurokawa and Shibuicha (1970) pointed out, an P. curtulum the color of the axis seems to vary with envi- ronmental factors. The same applies to the other species With blackened axes, where colorless and hyaline specimens can always be found. The blackened axis appears to have no consistent taxonomic value and therefore this criterion should not be used to divide the genus Ptlophorus into sec- tions as done by Sat6 (1940). The taxa Ptlophoron section EupiLophoren "Sato (Journ. Jap.) Bou. 10: 175.040) eandeee— lophoron section Wrorieaulia Sato. (hournmes Jap. boy 10% ae 1940) therefore should be abolished. Oe CESK map. 9 P. ntgrteaule - distribution of oi 7 In figure 8 of his paper Sat6 (1940) gives a scheme of the transverse section of pseudopodetium and apothecium of his two sections. This rigure is misleading in several respects. The species of his section Euptlophoron are shown corbe hollow, Which 1s not true for most species of Prlo= Pioris. Only 4ane Pr. aetcularte, and P. robustus do -old.pseudo-— podetia sometimes become hollow. The section of Mgrtcaulta shows a black central column of the pseudopodetium, which passes directly into the central part of the apothecium. In reality the apothecium and the pseudopodetium are not epaly ot different ontogenetic origin, but are also dis- tinctly separated by the boundary-texture. Furthermore, the pigment is not restricted to the axis of the pseudo- podetium, but the phyllocladia and the thallus horizonta- lis are also pigmented. Distribution (map 9) The distribution of P. ntgrtcaule corresponds with the occurrence of P. acteularte and P. clavatus, but the spe- cies is much rarer. Until now P. ntgricaule has been found in two areas only: Japan and the west coast of America. Pilophorus robustus Th.Fr., De Stereocaults et Ptlophorts Comm.: 41 (1857). Type collection: Norway, Hedmark, Osterdal, Amot, Austa-elv, 1837, Blytt (UPS, holotype). Ptlophoron polycarpum Tuck., Am. Journ. Arts Sci. 2, 15: 427 (1858). Type collection: Bering Straits (FH). fig. 9 P. robustus (ce - cephalodium, p - pycnidium) vy. 3 Thallus horizontalis granular or pulverulent, or indistinctly squamulose, evanescent in older specimens, grey-green when dry. Gra- nules about 1 mm diameter. Pseudopodetia stout, massive, seldom be- coming hollow when old, irregularly or umbellately branched in the upper part, normally 1.0-2.5 cm high, but sometimes up to 5 cm high and 3 mm broad (fig. 9). Hyphae of stalk strongly gelatinized with very small lumina (c. 0.5 pm), ecellishaty least. LO in length, ori- entated more or less parallel to the longitudinal axis of the pseu- dopodetium. Algal layer not homogenous, separated in loose granules. Old pseudopodetia becoming more or less decorticate. Cortex-granules c. O.2 mm diameter. Pycnidia bottle-shaped, apical on the tips of short lateral bran- ches or at the tip of smaller young pseudopodetia (fig. 9). Conidio- phores c. 10 pm long, unbranched with elongate cells. Conidia ter- minal, sickle-shaped, 5 x 1 pm. Yoo rR ete e ate e ee 2 ? A nN Both map 10 P. robustus - world distribution (places listed @ and not listed M® in the description. a specimens not examined) 319 Apothecia numerous, crowded at the apices of the pseudopodetial branches, ustially’c. 2.5 mm (from 0.5 to 4 mm) broad, 1.5 mm high, globose. Margins of crowded apothecia sometimes grown together se- condarily. Margin of the hymenium extending downwards below the point of attachment to the stalk, curving inward and upward before reaching the pseudopodetial surface. Vegetative tissue forming a large columella within the apothecium (fig. 12). In young specimens only a small columella is present. Boundary-texture and pigment- boundary absent. Hymenium c. 200 pm high, lower-half pigmented. CHS 4 map Tl P. robustus - distribution in Scandinavia 320 Asci eight-spored. Spores simple, rounded when young, spindle-shaped when mature, 18.0-24.0 x 4.0-6.5 pm. In one specimen some apothecia without pseudopodetia sessile on the thallus horizontalis. Phycobiont green, Pleurococcus type, cells c. 9 jam diameter. Cephalodia large, brown to orange, lumpy, 1.0-1.5 mm diameter, Situated on the thallus horizontalis and on lower parts of older pseudopodetia (fig. 9), with Wostoc. Growing on silicate rock and small stones. Remarks Ptlophorus robustus can be separated from all the other species by the relatively tall, umbellately branched pseu- dopodetia, the evanescent thallus horizontalis, and the large and well-developed columella in the apothecium. In some specimens the very young and small pseudopodetia re- semble P. cereolus, but the latter is always sorediate. In some cases certain specimens of P. robustus and P. acteu- laris might be confused, especially in material collected in Alaska, where both species occur together. This error is possible if the pseudopodetia of P. acteularts are rich- ly branched in their upper part and thereby resemble the umbellately branched thalli of P. robustus. But the two species can always be distinguished from the occurrence of a columella in the apothecium of P. robustus. Unusually small or very large specimens are quite fre- quently encountered in this species. Some specimens from different parts of the world are barely 1 ecm high and have pseudopodetia of less than 1 mm in diameter. Others may have pseudopodetia up to 5 em high and 6 mm broad at the base. Among these*large specimens is P. robustus f. magnus Savicz (1941),' but the type-material is in-no way unique | among the specimens examined. It is very interesting that unstalked apothecia which grew directly from the thallus 3 ® ie SNe. Aker Oy a s ee b sry me 8 Sos, ee Os) ovae SO PAL MESS Pe SE Re Mig Se SRT. a SS ZA =| a Soe ‘ge be —— —— = aS Imm fig. 10 P. strumatteus (ap - primordium of apothecium, ce — cephalodium) horizontalis were found. This is quite common in species with small pseudopodetia such as P. ftbula, but is rather unusual in the larger species. Distribution (map 10-11) WAAL Pe ropustus «snows -ardastinct arctic, circumpolar distri- bution. In Scandinavia where most samples were found the Species is almost exclusively restricted to the high moun- tains as was already established by Ammann and Ammann (1969); but here too, its clear dominance in northern re- Plonseis, Obvious. Alsecond centeryor distribution aseAlas— ka and the Tschuktschen Peninsula. These two main areas are connected by specimens collected in Siberia, Spits- bergen and Greenland. I have not seen the material from Spitsbergen myself, but the information was given by Krog Specimens found outside Scandinavia and Alaska: Spitsbergen, Svenskegya, Kong Karlsland, E.Dahl, 1936; Greenland, Laksefjord (C); Greenland, Laksefjord, Proven District (C); Green- land, Diskofjord, Kuanersuit (C); Greenland, Disko, Nordfjord, Kugssinerssuaq (C, 2 specimens); USA, Alaska, Aleutian Islands, Amchitka Island (WIS); USSR, Tschuktschen Peninsula (2 specimens LE, M); USSR, Arakamchechen Is., Tschuktschen Peninsula (BM, FH, PC, US); USSR, Wrangel Island (3 specimens LE); USSR, Taimyr Peninsula (3 specimens LE; GB, 2 specimens S); USSR, Laja River, Bolchese- melskaja (LE); USSR, Tobolsk (LE); USSR, Beresow, near Tobolsk (LE, Oa Pilophorus strumaticus Ny]. ex Cromb., Monogr. Lich. Brit.: 115 (1894). Type collection: Great Britain, S. Aberdeenshire, Brae- mar, Morrone, 1869, Crombie (BM, lectotype). P. robustus f£. dtstans Hult., Bih. K. svenska Vetensk. Akad. Handl. 26: 17 (1900). Type collection: Sweden, Dalsland, Hattefjdll, prope Vagsdter, 1870, Hulting (GB, lectotype; UPS, lectopara- type (1895)). P. dtstans (Hult.) Magn., Bot. Notiser 108: 298 (1955). Thallus persistent, minutely granular or nearly squamulous; gra- nules O.1 mm diameter, often becoming confluent to form aggregations c. 1 mm in size, grey when dry. Pseudopodetia pin-like, up to 2 mm high, O.5 mm diameter (fig. 10). Internal hyphae strongly gelati- nized, cell lumina O.5 wide, hyphae of upper part usually grey- brown. Pseudopodetia mostly covered by a layer of algae in con- tinuous granules or squamules. -Pycnidia apical on short pseudopodetia, up to O.5 mm high, often surrounded by a collar of granules or cortical scales. Conidiopho- res c. 45 pm long, unbranched with terminal sickle-shaped conidia Dex il pmein’ size. Each mature pseudopodetium nearly always bears a single hemi- spherical apothecium, c. 1 mm diameter. In some cases two separate apothecia develop at the tip of the pseudopodetium but later merge together. Apothecial margin down turned as far as the point of attachment to the stalk. The boundary between the generative and vegetative tissue arches upwards to form a small columella at the center of the apothecium (fig. 12). The boundary-texture and pig- ment-boundary between the generative and vegetative tissue are ab- sent in mature apothecia. The development of apothecial primordia, which have a pigment-boundary is very characteristic for the spe- Sa7 cies (Jahns, 1970a). Hymenium c. 230 pam high, lower part pigmented, margin sterile, consisting only of paraphyses. Excipulum absent. Asci eight-spored. Spores rounded when young, becoming spindle-shaped when mature, TATOS19, 08X46. SS 25 pm. The brown-black pigment is densest directly below the paraphyses and tends to become less intense towards the columella. Phycobiont green, Pleurococeus type, cells c. 8 pm diameter. Cephalodia brown, granular, surface irregular and lumpy, contain— ing Sttgonema. The development of the cephalodia was described else- where (Jahns, 1972). Growing on silicate stone. map 12 P. strwmaticus - distribution in Great Britain ae EAE: Remarks The distinct columella in the apothecium is character- bstic Of Mature specimens of Chis species,. distinguishing them from the other small species. P. robustus the only other species with a columella, has a distinctly different habit. map 13 P. strumatteus - distribution in Scandinavia ( mm places not listed in the description, A specimens not examined) 324 The name P. strumattcus was first used by Crombie (1875, p. 140), but as no description was appended it was a nomen Udine Cine re. Distribution (nap 12-13) P. strumatitcus has been found in western Scandinavia and in Great Britain. In contrast to P. robustus and P. cereolus, this species obviously prefers an oceanic climate and ex- tends far less to the east and north of Scandinavia than the other two species. Similar observations were made by Ammann “and, Ammann (19090)\eeTwo of their locations have been included in map 13 (marked. by a triangle), without having been examined by me. Pilophorus vegae Krog, Norsk Polarinstitutt Skrifter, 144: 56-57, Oslo (1968), fig. 14. Type collection: Alaska, Bering Sea district ,sNunivak ishand,. Roberts Mt. vl7O0.ft, Kroqt(0;, holotype). Thallus horizontalis crustaceous, consisting of loosely aggre- gated or nearly evanescent granules, c. O.5 mm diameter, white when dry. Pseudopodetia simple or branched in the upper part. Branches either dichotomous or irregularly bush-like. Pseudopodetia 1.0-1.5. cm high and c. 3 mm broad. Younger parts covered with a continuous algal layer, consisting of minute granules or scales. Color of scales white, later becoming blackened at the base of the pseudopo- detia. Basal part of old pseudopodetia often ecorticate. Central column of the young stalks hyaline, while the interior of older stalks is completely black. fig. 11 P. vegae (ap - primordium of apothecium, ce - cephalodium, s - broken stalk with pigmented center) RES Pycnidia not observed. Mature apothecia are absent in all specimens examined, but pri- mordia of apothecia are found terminal at the tip of the stalks. Phycobiont green, Pleurococcus type. Cephalodia stipitate on the pseudopodetia (fig. 11), containing Nostoe in the type-specimen and Sttgonema in other specimens. Small cephalodia-like structures on the substrate contained algae of the Gloeocapsa type, but it seems doubtful, whether they belong to this lichen. Growing on silicate rock. Remarks P. vegae can be confused with P. robustus and P. actcula- rts occurring in the same region. The clearest character- istics of the species, which are lacking in P. robustus and P. acteularts, are the white color of the young pseudo- podetia and the black pigment of the central column. P. ntgrtcaule, which also has a white cortex and a black cen- tral column is smaller and easily recognized by the glob- ular or peltate shape of its cortex granules. map 14 P. vegae - distribution ( e@ places listed in the des- cription of the species) 52:0 Distribution (map 14) P. vegae is known from Alaska and the Bering Strait area only. I have seen the following specimens: USSR, Tschuktschorum, Bovidential, 64°45'N, 174°w (2 specimens LE, O); USA, Alaska, Bering Sea district, Nunivak Island, Roberts Mt., 1700 £t(O); USA, Alaska, Central Pacific Coast, Talkeetna Mts. (0); Canada, British Columbia, Queen Charlotte Islands, Moresby Island, Laing Point Mountain, SE of road into Peel inlet, c. i mile of Laing Renliae,; LOO) ae (GAN) ~ Lichens to be excluded from the genus Pilophorus Four species have been described as belonging to the genus Ptlophorus, which must be placed elsewhere: Pilophorus colensoi (Bab.) Knight, Trans. Proc. New Zealand 16: 400 (1884). Stereocauton colensot Bab., in Hooker, Flora N.Z. 2: 294 (1867). Type collection: New Zealand, Colenso 2746 (BM, holotype). Corynophoron colensot (Bab.) Nyl., Lich. Nov. Zeland.: 15 (1888). Thallus horizontalis thin, granular and evanescent. Pseudopode- tia up to 5 mm high, branched. Cortex of irregular scales. Cepha- lodia sacculate, on the pseudopodetia, containing Sttgonema. De- velopment of apothecia following the typical ontogeny of a Stere- ocaulon species (Jahns, 1970a). Remarks The helical ascogones are especially typical and quite different from the straight ones found in the genus Prlo- phorus. The species therefore belongs to the genus Stereo- caulons @ View salso taken by Th.Fries (les7) and aby mam, Lamb in a note attached to the type-specimen. Raésdnen (1932, p. 24) described P. colensot var.reagens Ras. from Tierra del Fuego. I have seen material from the following places: New Zealand, Colenso 2746 (BM, holotype); Fuegia occid. supra Ru- pium (B); N-East of New Zealand (G). Pilophorus conglomeratum F.Wils., Linn. Soc. Journ. Bot. 28: 372 (1891). Type collection: Hab. supra truncum arboris smortuum muscosum in Black spur. Victoria, Australia (BM, holotype). Ptlophorus cartosum Hue, Nouv. Arch. Mus. 3, 10: 280 (1898). Thallus horizgontalis thin, grey-green, consisting of minute gra- nules up to O.1 mm in diameter. Thallus horizontalis persistent or evanescent. Thallus verticalis up to 2 cm high, pin-like, un- branched. In young specimens more or less smooth, covered with a thin layer of minute granules similar to those forming the thallus horizontalis. Thallus verticalis quickly becoming bare and divided by lengthwise-oriented furrows. In old specimens the stalk consists of numerous nearly-separated columns. fig. 12-16 anatomy of apothecia (bt - boundary-texture, c - colu- mella, pb - pigment-boundary) Tig. P. ntgrtcaule with peltate scales, longitudinal section of the stalk QL Sat LED Sey ny ay ay ba Sw Say a Seng ~ =; fy S — = = Ss = = — —— Ss — Fs — ZS ¢ Ss S “a i( 3 Noy 16 1%, DLO Several small apothecia crowded at the tip of the stalks, gradual- ly merge into one large fruitbody. Spores 14 x 3.5 pm, spindle- shaped. Apothecia dark-brown. Cephalodia absent. Remarks The species somewhat resembles P. robustus, but the ab- sence of cephalodia and the brown pigment of the apothe- cia are striking differences. The whole habit is slightly different from a Ptlophorus and therefore the species should be excluded from the genus. Perhaps the lichen is related to Baeomyces or to one of the genera of the Clado- niaceae (e.g. Heteromyces, Thysanothectum) but the question can not be settled before more is known of its ontogeny. I have seen material from the following places: Tasmania, Hartz Mountain plateau, on rotten wood in forest, extreme- ly damp and well shaded, alt. 3700 ft (BM); Central Western Tasma- nia, forest at western side of Mt. Arrowsmith near Franklin River, (BM); North Western Tasmania, Ball Room Forest Track, Dove Lake, near Cradle Mt. (BM); New Zealand, South Island, Fjordland, Eglinton Valley, Cascade Creek, alt. 1600 ft (BM); Victoria, Australia (BM, holotype) . Pilophorus staufferi Frey (nomen nudum), Bot. Jahrb. 86, 1-4: 242 (1967). Type collection: Neukaledonien, Monts des Koghis, Mt. Bono, Gipfelgrat, 1070 m (BERN). One's first impression of this species is that 16 as very much like P. robustus. The thallus verticalis is um- bellately branched in the upper part, each branch bearing one to several apothecia. But on close inspection the lichen seems to be related to P. conglomeratum. The color of the apothecium is more brown than black, and the stalks Show the same furrows as in P. conglomeratum. Cephalodia are also-absent. Thelthallus horizontalis of —Peterausrerr, .un= like the granulated thallus of P. conglomeratum, consists of small phylocladia about 1 mm in diameter. I think that P. stauffert and P. conglomeratum belong to the same genus; their exact taxonomic positions must be decided later. Pilophorus pileatum (Mont.) Zahlbr. (nomen nudum), Catal. Lich. 1023S) Se (1939)e Cladonta ptleata Mont., Annal. Scienc. Nat. Bot. 3, 18: 310 (1852) Type collection: Coquimbo in Chile. I have seen no samples from this species, but the descriptions of Montagne (1852) and Vainio (1887) mention brown-black apothecia. Therefore it certainly does not belong to Ptlophorus. References Acharius, E. (1798) Lichenographiae Sueciae Prodromus. Lincopiae. Acharius, E. (1803) Methodus qua omnes detectos Lichenes. Stockholm. Acharius, E. (1810) Lichenographia universalis. Gdédttingen. Ammann, K. & B. Ammann (1969) Die fennoskandische Verbreitung von Pilophorus (Tuck.) Th.Fr., Stereocaulaceae. Herzogia 1:87-94. Crombie, J.M. (1875) Recent Additions to the British Lichen-flora. J.Bot.Lond. 13:140-142. SENS) Crombie, J.M. (1894) A Monograph of Lichens found in Britain. British Museum, London. Culberson, W.L. (1970) The Typification of the Lichen Genus Pilo- phoron. Bryologist 73:630-632. Fink, B. (1910) The Lichens of Minnesota. Contr. U.S. nat. Herb. 14, 1:1-269. Frey, E. (1967) Die lichenologischen Ergebnisse der Forschungsreisen des Dr. Hans Ulrich Stauffer. Bot. Jahrb: 86, f-4:3209-—255. Fries, Th. (1857) De Stereocaulis et Pilophoris Commentatio. Uppsala. Fries, Th. (1861) Lichenes Arctoi Europaeae Grénlandiaeque hactenus COgniti.. Nova, Acta Reg ..soc. scient.,Uppsal.. 3,°322/8; Fries, Th. (1871) Lichenographia Scandinavica I, Uppsala. Fries, Th. (1874) Lichenographia Scandinavica II, Uppsala. Fries, Th. (1888) Nagra anmarkningar om Sladgtet Pilophorus. Bot. Notiser 1888: 212-214. Fries, Th. (1889) Einige Bemerkungen Uber die Gattung Pilophorus. Bot. Centralblatt 38:764-766. Hawksworth, D.L., P.W. James & J.R. Laundon (1972) The Nomenclature OrePilopnorusc., Taxon 21 ,52/ 5532/2920. Henssen, A. & H.M. Jahns (1973) Lichenes. Stuttgart. Hooker, J. (1867) Handbook of the New Zealand Flora II. London. Hue, A.M. (1898) Lichenes extra-europaei. Nouv. Arch. du Muséum 3, 10:280 Hulting, J. (1900) Dalslands Lafvar. Bih. K. svenska Vetensk. Akad. Handl.1.626,00/3:17. Jahns, H.M. (1970a) Untersuchungen zur Entwicklungsgeschichte der Cladoniaceen unter besonderer Berticksichtigung des Podetium- Problems. Nova Hedwigia 20:1-179. Jahns, H.M. (1970b) Remarks on the taxonomy of the European and North American Species of Pilophorus Th.Fr. Lichenologist 4:199- 213. Jahns, H.M. (1972) Die Entwicklung von Flechten-Cephalodien aus Sti- gonema-Algen. Ber. Dtsch. Bot. Ges. 85, 10-12:615-622. James, P.W. (1965) A new Check-list of British Lichens. Lichenolo- GSC ess Eo=i 53. Knight, C. (1884) On the Lichenographia of New Zealand. Trans Proc. New Zealand 16:400. Krog, H. (1968) The Macrolichens of Alaska. Norsk Polarinst. Skrifter 144:1-180. Kurokawa, S. & H. Shibuichi (1970) Notes on Japanese Species of Pa OpnOron. WOurn. sap. Sot. 45, 2 /5=<04. Magnusson, H. (1937) Férteckning 6ver Scandinaviens vaxter, 4 Lavar. Magnusson, H. (1955) New or otherwise interesting Swedish Lichens XV. Bot. Notiser 108:298-299. Montagne, D.M. (1852) Diagnoses phycologicae. Ann. Sci. Nat. 3, 18: 302-319. Nylander, W. (1857) Enumération générale des lichens. Mém. Soc. Sci. nat. Cherbourg 5:85-146. Nylander, W. (1860) Synopsis Methodica Lichenum I. Paris. Nylander, W. (1888) Lichenes Novae Zelandiae, Paris. Rasanen, V. (1932) Zur Kenntnis der Flechtenflora Feuerlands. Ann. Socs Zool. Bot. Fenn. Vanamo 2, 1:24. Rasdanen? V. (1946) Lichenes novi II. Ann. Soc. Zool. Bot. Fenn. Vanamo,, Notil. ©Bot. 2isi=7. Rasanen, V. (1950) Lichenes novi VI. Ann. Soc. Zool. Bot. Fenn. Vanamo 5:28-29. 330 Sat6, M.M. (1933) Notes on some Japanese Lichens determined by Dr. Bav. A.) Wainio Lil. Journ. Jap. Bot. 9221i-214. Sato, M.M. (1940) East; Asiatic) Lichens ITi< \ Journ. Jap. Bot. 16: 173-147: Savicz, Vee. (1941) De nonnullis lichenibus URSS. Notul. syst. inst. GLY DECG .sHOrLILSDOLtEreLLOpol. =o 4/—9". Stitzenberger, E. (1862) Beitrag zur Flechtensystematik. Ber. Utiber die Tatigkeit St. Gallisch. naturwis. Gesellsch.:124. Tuckerman, E. 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(1939) Catalogus Lichenum Universalis 10. Leipzig. af MYCOTAXON Otel NO) DD. wos 558 July-September 1981 NEW DEMATIACEOUS HYPHOMYCETES FROM) TROPICAL, RAIN FOREST LITTER S. ONOFRI, D. LUNGHINI, A. RAMBELLI and L. LUSTRATI Cattedra di Micologia, Istituto dell'Orto Botanico, Universita di Roma, Largo Cristina di Svezia 24, 00165 Roma, Italy. Summary In this paper, anew genus, Chryseidea gen.nov. with the new species C.africana, and two new species, Beltrania onirica sp.nov. and Phaeo- trichoconis aurata sp.nov., from Tai National Park (Ivory Coast) fo- rest litter, are described. Among mycological investigations carried out by this Chair on the Tai project (M.A.B., Project n.1) to study the influence of human ac- tivities on the south-west forests in the Ivory Coast and with the pur- pose to improve the environmental knowledge and quality, three new species and a new genus of Dematiaceous Hyphomycetes are described. They were found on dead leaves collected on the soil surface in the Tai National Park forest. Every attempt at isolation in pure culture was unsuccessful. The following descriptions concern observations on na- tural media, Exsiccata are available in the Mycological herbarium of the Rome Botanical Garden, (RO). Chryseidea Onofri gen.nov. Conidiophora macronematosa, synnematosa, erecta, simplicia, pig- mentata, in apice semper sterilia, Cellulae conidiogenae a latere po- sitae, determinatae, discretae, simplices, ampulliformes, apice den- ticulato, circiter in medio synnemati circumpositae, Conidia hyalina, falcata, Sp.typ.: C.africana Onofri sp.nov. Conidiophores macronematous, synnematous, straight, simple, pig mented, with apices always sterile. Conidiogenous cells lateral, de- terminate, discrete, simple, ampulliform, with a denticulate apex, surrounding the conidiogenous apparatus approximately in the mid part. Conidia hyaline and falcate, 40 um i rs —— —Y == =—— ee ee —— SSS Nk h 555 Chryseidea africana Onofri sp.nov. Conidiophora macronematosa, synnematosa, solitaria, recta, simpli- cia, adtenuato et sterili apice, ex constantibus et rectis hyphis (2 um crassis) composita, pallide brunnea, usque ad 560 wm longa et 42 um ad basim crassa, Cellulae conidiogenae a latere positae, discretae, simplices, ampulliformes, cum apice saepe deflectens conidiorum forsitan pondere vel magna copia et denticulato ob subsequentes loco- rum qui conidia generant proliferationes; orientes ex hyphis laterali- bus, adherentibus et parallelis ab hyphis synnematis et orientibus pro xime sub saeptis; ab circiter tertio altitudinis a basi usque ad dimidium synnema circumpositae, sufflavae; 7,8-11,2x2,3-2,7 wm. Conidia fal- cata, asaeptata, extremitatibus acuta, hyalina, omnia agglutinata; cir cum conidiophora permanent in formam similia Chaetopsina fulva; 14,5-18,5x1,4-1,9 wm. In foliis emortuis, Tai, Ora Eboris, holotypus: H.B.R. 116A, (RO). Conidiophores macronematous, synnematous, solitary, straight, simple, narrowing to the sterile apex, composed of straight and regu- lar hyphae, Ochraceous-Tawny in colour (Ridgway, 1912), up to 560 um long and 42 wm thick at the base, Conidiogenous cells lateral, discrete, simple, ampulliform and denticulate from successive proli- ferations, with the apex frequently repent, presumably under the weight of conidial production; they originate from lateral hyphae that grow just under the septa of conidiophorous apparatus at about one third of the height from the base and extend up to half of the height of the coni- diophorous apparatus, along which they grow side by side, Light-Buff in colour (Ridgway, 1912); 7,8-11,2x2,3-2,7 wm. Conidia 0-septate, falcate, with very sharp apices, hyaline, 14,5-18,5x1,4-1,9 wm; they remain coiled at the apex of each conidiogenous cell in a compact mass surrounding the conidiophorous apparatus. The microorganism described presents morphological characters different from others belonging to genera with synnematous conidio- phorous apparatus, It differs in the position of the conidiogenous cells along the conidiophorous apparatus, in the resulting general structure (Chaetopsina like) and in spore shape. For these considerations we think necessary to propose the new genus, Chryseidea, and the spe- cies C,africana for our strain, Beltrania onirica Lunghini sp.nov. Coloniae effusae, Mycelium immersum, Setae erectae, simplices, le- ves, atrobrunneae, usque ad 506 um longae et 4,6 wm ad basim crassae, Conidiophora macronematosa, mononematosa, simplicia aut rarius ra mosa, saeptata, flexuosa, levia, non constanti crassitudine, pallide brunnea, 184-236x4,6 um; ab eadem basi setarum oriuntur, Cellulae conidiogenae integratae, terminales, polyblasticae, denticulatae, sym 334 20 wm 40 um a. conidiophores; b, conidiophores Fig. 2 - Beltrania onirica sp.nov.: and seta; c. conidia. sf SO podiales, locis qui conidia generant magno intervallo positis, pallide brunneae. Conidia levia, acrogena, protoplasmate granuloso, solitaria, conica-campanulata, zona supraaequatoriali subhyalina et tenui orna- ta, ad basim denticulata, ad apicem breviter appendiculata, pallide brunnea; 19-21x11,5-12,5 wm; appendix circiter 2 wm longa. in foliis emortuis, Tai, Ora Eboris, holotypus: H.B.R..117A, (RO). Colonies effuse. Mycelium immersed. Setae straight, simple, with thick and smooth walls; Argus Brown in colour (Ridgway, 1912); length up to 506 wm and 4,6 wm thick, Conidiophores macronematous, mono- nematous, simple or rarely branched, septate, slightly curved and irregulary thick, Deep Olive-Buff in colour (Ridgway, 1912), 184-236 x4,6 wm; originating from the base of the seta. Conidiogenous cells integrated, terminal, polyblastic, denticulate, elongating by sympo- dial proliferation, Deep Olive-Buff in colour (Ridgway, 1912). Conidia smooth, acrogenous, with granular protoplasm, solitary, co-- nic-campanulate, with a thin, not easily observed transverse band, with a very short appendage at the apex and denticulate at the base, Deep Olive-Buff in colour (Ridgway, 1912); 19-21x11,5-12,5 wm; appen dage about 2 wm long. The microorganism described presents characters different from those of related species, In particular it differs from Beltrania querna Harkness (Pirozynski, 1963) with conidia with a different shape and longer appendage; the conidial shape of our strain is very close to those of B, mangiferae Munjal et Kapoor (Pirozynski and Patil, 1970) and B, muelleri Rao et Mani Varghese, 1978, but these species are characterized by different conidiogenous cells, For these considera- tions we propose the new species B.onirica for our strain. Phaeotrichoconis aurata Rambelli sp.nov. Coloniae effusae, Mycelium immersum, Conidiophora macronematosa, mononematosa, solitaria, erecta, simplicia, saeptata, levia, brunnea- sufflava, usque ad 126 wm longa et 3,6-5 wm crassa (ad basim 8-10 wm), Cellulae conidiogenae polytreticae, integratae, sympodiales, cicatri- cibus conidialibus parum manifestis preditae, Conidia solitaria, sicca, acrogena, obclavata, 5-saeptata, ad basim trunca, ad apicem longo rostro praedita, levia, pallide brunnea-flava, protoplasmate omogeneo; 34,5-43,7x4,6-6,9 um; rostrum rectum, asaeptatum, hyalinum, usque ad 25 um longum et usque ad 2 wm crassum, In foliis emortuis, Tai, Ora Eboris, holotypus H.B.R. 118A, (RO). Colonies effuse, Mycelium immersed, Conidiophores macronematous, mononematous, solitary, straight, simple, septate, with smooth walls, Buffy Brown in colour (Ridgway, 1912), up to 126 wm long and 3,6-5 wm thick (8-10 um at the base), Conidiogenous cells polytretic, integrated, sympodial, Conidia solitary, dry, acrogenous, obclavate, transversely ———_—— Oe 356 Fig. 3 - Phaeotrichoconis aurata sp.nov.: conidiophores with conidia, Koy ii SRO acc Sette rte = sco saieneatensnnpeeceremnenraenstitene rgesesian Fig. 4 - Chryseidea africana: a. and b. Phaeotrichoconis aurata: c. and d. Beltrania onirica: e, and f, 338 5-septate, rostrate, with a large scar at the base, smooth, Tilleul- Buff in colour (Ridgway, 1912), with homogeneous protoplasm, 34,5- 43,7x4,6-6,9 um; scar about 2,5-3 wm wide, beak straight, aseptate, hyaline, up to 25 wm long and 2 um thick, Ellis (1971) modified the original diagnosis of the type species P, crotalariae (Salam et Rao) Subramanian to include also species characterized by polytretic, integrated, terminal and sympodial co- nidiogenous cells, Our microorganism seems includable in the genus Phaeotrichoconis as modified by Ellis (1971), but differs from the two described species, P,crotalariae and P,urariae Bharadway, in seve- ral characters and mainly in the dimensions; for these reasons we propose for our strain the name P, aurata as a new species. ACKNOWLEDGEMENTS The Authors wish to thank the Minister of Scientific Research of the Ivory Coast, for the assistance he has given, Dr. P.M. Kirk of the C.M.I. for a slide of P.crotalariae kindly sent us and our collea- gues, in particular Dr. A.M. Persiani, of the Laboratory of Mycology for their criticisms. We also thank Prof. O. Verona, Istituto di Micro biologia Agraria eTecnica, Universita di Pisa, Italy, for kindly revie- wing the manuscript. REFERENCES Bharadway, S.D. (1969). A new species of Phaeotrichoconis on Ura- ria picta from Varanasi, India. Indian Forester 95(1): 24-26. Carmichael?) [,W:, W.B.° Kendrick, “Isl. Connerswande py. o1glerg( 1980). Genera of Hyphomycetes. The University of Alberta Press, Edmon- ton, Alberta, Canada. Ellis, M.B. (1971). Dematiaceous Hyphomycetes. Commonw. Mycolog. Inst., Kew, England. Ellis, M.B. (1976). More Dematiaceous Hyphomycetes. Commonw. My- colog.=Inst., Kew, England. Matsusnhimaset = .(L0/))- Microfungi of the Solomon Islands and Pa- pua-New Guinea. Publ. by the Author, Kobe, Japan. Matsushima, T. (1975). Icones microfungorum a Matsushima lectorum. Publis byethie, Author mKobDewaapan. Morris, E.F. (1963). The synnematous genera of the Fungi Imperfec- ti. Western sillinois| Univ; {Ser sbiclogicalmociencesmio ws). Pirozynski, K.A. (1963). Beltrania and related genera. Mycological Papers, 90. Pirozynski, K.A. and S.D. Patil (1970). Some setose Hyphomycetes of leaf litter in south India. Can. J. Bot. 48: 567-581. Rao, V.G. and K.I. Mani Varghese (1978). Litter-Fungi: two unde- scribed Hyphomycetes from India. Nat. Acad. Sci. Letters 1(2): 49-50. Ridgway, R. (1912). Color standards and color nomenclature. Wash- ington, 1b.CG. Subramanian, C.V. (1971). Hyphomycetes, An account of Indian spe-— cies, except Cercosporae. Indian Council of Agricultural Research, New Delhi, India. MYCOTAXON VORPENTDIC RENO S825 epp5ji559=545 July-September 1981 COCHLIOBOLUS RAVENELIT SP. NOV. AND C. TRIPOGONIS SP. NOV. J. L. ALCORN Department of Primary Industries, Indooroopilly, Queensland, 4068, Australta Abstract Cochltobolus raveneltt sp. nov. and C. trtpogonts sp. nov. are described from culture, as the teleomorphs for Btpolarts ravenelit and B. tritpogonts comb. nov. respectively. Pompe Vimo (ames LUC Vaso EEpenenCsCOnCceptsein ticacomplex of fungi assigned to Drechslera sens. lat., I have attempted to produce teleomorphs for species where no such connection was known. This has been done to test the hypothesis that conidial state characters allow prediction of teleomorph correlations, and that new associations will prove to be Drechslera sens. str. - Pyrenophora, Btpolarts - Cochltobolus, and Exserohtlum - Setosphaerta (Luttrell 1977, 1978). Three new teleomorphs were described previously, two in Cochlto- bolus (Alcorn 1978a) and one in Setosphaerta (Alcorn 1978b), and those results support the predictive aspect of anamorph classification. Subsequently Cochltobolus states have been induced to form in cultures of other species. Two of these teleomorphs are described here as new. COCHLIOBOLUS RAVENELIIL Ascal states in Cochltobolus were found by McKenzie (1968) for two 'Helminthosporium' species which infect Sporobolus inflorescences in Queensland. The work was reported in a thesis, and subsequently none of the data presented have been published. McKenzie did not assign specific epithets to the Helminthosporium collections 340 studied. I have examined specimens she deposited in BRIU and there is no doubt that one of the anamorphs connected with a teleomorph was Bipolarts ravenelit (Curt.) Shoem. Luttrel1 (19:76) was unable to produce a-per tech State rior this species, and I made many unsuccessful pairings of cultures before a teleomorph developed. The methods used in the successful attempt are described briefly below. Ten collections of Sporobolus spp. with infections of the inflorescence caused by B. ravenelit were made in south- eastern Queensland, in an area approximately 60 km north of Brisbane. Conidia from each of five panicles in each collection were streaked over water agar plates and allowed to verminate., Pour verminatecasconidiamtrom cachap aveswere transferred to PDA, making a total of 20 transfers for’ each coblection. “Cullturesaweresparred=on sachs agar (hevere 1971).) each plate being amended with» maize lear sand barley Uraim. sihrcenmethodss0t. pairing. weresecmployed, A. Bulked conidial inoculum was prepared for each collection. Conidial suspensions from all isolates from a single collection were mixed and inoculated onto the plates. B. All possible pairings: bDetweenscollectwons weresmade, using the bulked conidial inoculum from each. C.. “All possable pairings between collections, using one single-conidium isolate from each. Inoculum consisted of blocks from PDA cultures. Ascocarps were obtained in only one pairing, namely with bulked inoculum from collection 7957 (Sporobolus ferttlts (Steudel) W.D. Clayton, Glasshouse Mountains, 24.v.1979) and collection 7963 (S. elongatus R. Br., .Maleny., J4.V,19/9)4, “Pour ascocarpsuwere: formed, each iwi tit aso LOpose basal’ body and a long cylindrical neck. They were visible only because the necks projected above the dense hyphal mat investing the basal section. Ascocarps contained many ripe asci which released ascospores freely in water. Single- ascospore culturés were established. These cultures were paired in all possible combinations, but only one pairing resulted in the formation of ascocarps. None were formed in single-ascospore cultures, but all produced an anamorph on PDA indistinguishable from B. ravenelit. 34] In another test, small numbers of ascomata formed in Dalringspotucultures. fromecollections 79/6 (5.7 fertiiis, St. Lawrence, 24.vii.1979) and 7977 (Sporobolus sp., Claliviewm 4 .Vi119 79). andy /976.and 79/89(S. fertilie, between Sarina-and Mackay, 24.vii.1979). Some single- ascospore isolates from these ascocarps produced the teleo- morph when paired, but not when grown alone under the same cultural conditions. Subsequently ascomata were formed in greater numbers when inflorescences and grain of S. elongatus Were Used iasathe plant substrate ian oach"s agar plates, mnsteadeotemarze, lean or Darley agradin:. Cochltobolus ravenelit sp. nov. Figures 1 - 4 Ascocarpi atrobrunnei vel atri, basi globosa 360-515 um diam. et collo cylindrico 980-1800 um alto, ad basim 95-135 DMparall eae CON so ouiine G1 alemAS Cia cy WIndri Ca. Vestigialessbitunicatdi , 165-500)x 14-1/-5 im... /AScosporae hyalinae, filiformes, versus extrema angustatae, rectae vel torsivae, 5-14-septatae, 155-310 x 5-7.5 um. In inflorescentiis et granis Sporobolt elongatt in AC ATOM d Chile ile OSU] BRI Pals LOomenolOLY DUS em DRUP 5027), IS0ZSeparatypi . Ascocarps dark brown to black with a globose body 360- J Omid lalimeanded paler cylindrical neck 980=1800) x 95- IS oeiMeate chesbase and) 95-1 loimeat. thesapex = Occasionally with a cylindrical base up to 280 um x 250 um below the ascogenous section. Ascocarp body often covered densely by reddish brown hyphae, the beak glabrous or with a few short Nag TSeiGl oie SASCIMCYINGtICal wUSidallyetanered «si only near the base but. lacking a distinct, pedicel, straight vor eurveds 9165-4002 x 114-175 ime Often, with asblunt baturcate foots o- Om median... VeSit1 ol alebitunicate Ur lens 2)/0 mASCOs Sporesuri ii tom: Stralont) ovecoi led helvcaluiys in thepascus. slightly tapered to obtuse apex, more so to base, 5-14- Ceptate loo LUE Xo oe Time Ore) | In some asci the degree of ascospore coiling is only very slight, while in others it is more pronounced. Some- times ascospores are tightly coiled in the apical part for 20-25 um, and loosely in the remainder. Ascospore diameter at the apex is about 4 ym, and at the base 2.5 wm. Conidia from two collections which gave rise to inter-ferile cultures of B. raveneltt are illustrated in Figure 4. 342 Cochltobolus ravenelii is quite distinct from C. sporobolt Castellani, which occurs on Sporobolus leaves (Castellani 1951). C. sporobolt has smaller, short-beaked ascomata, smaller asci, and shorter 4-6-septate ascospores. COCHLIOBOLUS TRIPOGONIS Drechslera trtpogonts A.S. Patil & V.G. Rao was described from India, infecting the inflorescences of Tritpogon jacquemonttt Stapf (Patil §& Rao 1972). Subsequently L-collected a similar fungus onl. cOLt7) Ormes etl eMuel 1} C.E. Hubbard in Australia (Yetman, New South Wales, 12.v. 1977, J. LA. 9/7158) =Comparison with thestype specimen (IMI 161251) showed that the fungus on this host was con- specific with D. tripogonis. No other collections have been made, although 7. lolttformis has been examined at three other sites in Queensland (Bunya Mountains, Cooroy, Peregian Beach)® Herbarium specimens Ofpthisespecrcsen lebki mi Lon many different localities in eastern Australia, were also searched. No infections Of the, int lorescenceswerescetececd. Single-conidium isolates from the collection on T. lolitformts were paired. A teleomorph referable to the genus Cochltobolus developed in these paired cultures. Single-ascospore isolates sometimes formed ascomata when: paired, but never when grown alone. The anamorph formed by Single-ascospore cultures is indistinguishable from that produced by the original single-conidium isolates. Cochltobolus trtpogonts sp. nov. Figures 5 - 8 Ascocarpi atri, basi globosa 300-590 um diam. et collo cylindrico 880-2000 x 50-150 um. Asci cylindrici, vesti- Plales bitunicati, (17/0-410 xml sb als oem. en sScOSpOTde hyalinae, filiformes, versus extrema angustatae, torsivae, 4-9-septatae, 175-275 x 4-6 um. BivSeyet ea Cochltobolus ravenelitt (1) Ascocarp x50 (2) Asci x200 (3) Ascospores x320 (4) Conidia from collections 7957(a) and 7963(b), x320. Isolates from these collections formed the teleomorph of C. raveneltt when paired. Figs. 5 - 8 Cochltobolus tripogonis (5) Ascocarp x50 (6) Ascus x320 (7) Ascospore x320 (8) Ascosporic chlamydospores x320. 343 Sess 344 In) foli1sZeae=maydte in agarovoachil tt 80 (ee DRIP 123757 sho lLotypuss BRIE 227 3542paraty pus. Ascocarps black, with a globose basal part 300-590 um diam. bearing a long, ‘straight or slightly curved cylindrical neck 880-2000 x 50-150 ym (Fig. 5). . Conidiophores: are not formed on ascocarp body or neck. Asci cylindrical, vestigial bitunacates slightly tapéredsnear the basco /U—St0e x al 252 18.5 um (Fig. 6). Ascospores filiform, hyaline; helically coiled in the ascus and completely filling it, commonly somewhat loosely coiled in the median section and more tightly in the extremities, gradually tapered to approx- imately half the maximum width at each end, 4-7-septate, 175-275 x 4-6 pm Fie ew Single cells of some ascospores become dark brown and thick walled, while retaining their original shape (Fig. 8). These darker cells are regarded as ascosporic chlamydospores. I am unaware of any other reports of this phenomenon in Cochltobolus species. In additional testing, inflorescences of 7. lolitforms on Sach's agar were used as the substrate in an attempt to produce greater numbers of ascocarps. Ascomata were formed on the rachis and florets, but no more abundantly than previously. In some, a basal pedicel 30-150 um high and 30-58 um diam. was present below the swollen fertile section OteulCEascocanp. The teleomorph of C. trtpogonts is distinct from that of most other Cochltobolus species because of the very long ascocarp neck. Species such as C. cymbopogonis J.A. Hall & Sivanesan, C. hawattensis Alcorn, and C. raveneltt, which also have ascomata with long necks, can be distinguished by other morphological characteristics of teleomorph and anamorph. Evidence supporting recognition of Btpolaris as a genus distinct from Drechslera will be presented elsewhere. Part of that evidence relates to D. trtpogonts, for which a new combination is necessary. Btpolarts trtpogonis (A.S. Patil §& V.G. Rao) comb. nov. Drechstlera trtpogonis A.S. Patil & V.G. Rao, 1972, Transeepy.e er my.cOlceoOC moons 4. 345 Single-ascospore cultures representing opposite mating types of both Cochltobolus species have been deposited in the culture collection of the Commonwealth Mycological Pasertutes (I Mim255052mande255655,"0e ravenel77;) JIMIe 235294 Bide 259295 68C. Lr tpodcnts):. ACKNOWLEDGEMENTS I- am grateful to Dr R.F.N. Langdon (BRIU) and Prep eemoutron a IMijeeromercidingmspecimens in their Keeping, and to Mr R.G. O'Brien for assistance in obtaining some of the Btpolarts ravenelizt collections. REFERENCES Alcorn, J.L. 1978a. Two new Cochltobolus species. Trans. Demeny CO.) O Cis Ung Ol O07, Alcorn, J.L. 1978b. Setosphaerta monoceras sp. nov., ascigerous state of Hxserohtlum monoceras. Mycotaxon 7: 411-414, Castellani, E. 1951. Una nuova specie di Cochltobolus. MY.cOpDa claeMy CO law app lg hOceeo oD fas Hebert, T.T. 1971. The perfect stage of Pyrtcularia grisea. Phytopathology 61: 83-87. Luttrell, E.S. 1976. Ovarian infection of Sporobolus potrettt by Bipolarts ravenelit. Phytopathology 66: 260- 268. Luttrell, E.S. 1977. Correlations between conidial and ascigerous state characters in Pyrenophora, Cochltobolus and Setosphaerta. Rev. Mycol. 41: 271-279. Luttrell, E.S. 1978. Biosystematics of Helminthosportum: impact on agriculture. In Btosystematics in Agriculture. Beltsville Symposia in Agricultural Research 2, Bcdlm..AveRombercer, RH. bootes, LV Knutson, GP. l. Lentz, pp. 193-209. Allanheld, Osmon §& Co., Montclair, New Mersey. McKenzie, F. 1968. A study of false smut disease of Sporobolus spp. M. Sc. Thesis, University of Queensland, Brisbane. Patil, A.S. § Rao, V.G. 1972. a immature asco- the field. Spore, sou mature, CUP-MM 1566, xn OOO: MICROGLOSSUM Gillet 1879 One known Maraconesian species 1. Microglossum olivaceum (Pers. : Brioeeo letme lam pr inmenvean Wicca) MNes ali Yoo Nevis) RECENT TAXONOMIC TREATMENTS: Dennis (1978), Mains (1956), Nannfeldt (1942). PREVIOUS MACARONESIAN RECORDS: None. mY Ere LOCA hy see urope. KNOWN MACARONESIAN DISTRIBUTION CANARY ISLANDS. Tenerife. CUP-MM 1307(TFC). SUBSTRATA: On soil. Notes: Mains (1956) included Thuemenidium (= Corynetes) in his very broad concept of the genus. Ic ID Tea ay a M. olivaceum, 6 asco- SPOCGS, a Amea SClee WA + Pole wiwallechanne 16 ascospores, CUP-MM 1307, xe L000: Th. atropurpureum, mature ascus with J+ pore wall channel, immature ascus, 12 \ascospores, CUP=MM 1371; x LOO: THUEMENIDIUM Kuntze 1891 emend. Maas Geesteranus 1964 One known Macaronesian species 1. Thuemenidium atropurpureum (Batsch : Fr.) Kuntze, ReEVisS. 3 OC Dil wees) el oo e. = Corynetes atropurpureus (Batsch : Fm) Dur= and. ‘Anne Mycol. 86:4] 42161 908" = Microglossum atropurpureum (Batsch : Fr.) Karst Acta. Soc shatindgw hl. Leen. .o1n LiOe 1885. 365 RECENT TAXONOMIC TREATMENTS: Dennis (1978), Maas Geesteranus (1964), Mains (1956: Microglossum), Nann- feldt (1942: Corynetes). PREVIOUS MACARONESIAN RECORDS: *Bafiares Baudet & eee 1980), nt Beltran, Tejera (1980),**Cool (1924), ***Cool (1925). Pees OCA TRY eEurope. KNOWN MACARONESIAN DISTRIBUTION a CANARY “I1SUANDS: *Gomera. *CUP-MM 1371(TFC|[=TFC 770])). xx ***Tenerife. CUP-MM 1136(0), ** ***1652(L). SUBST RATA: On soil: Notes aaa NOuGns sthesechilerion. Of Malleamorphous,. colored pseudoepithecium has been used to distinguish Thuemeni- dium, it is often absent. Mains (1956) I think ‘correctly Synonymized 3 species Durand (1908) distinguished in has’ monograph, Corynetes atropurpureus, C. purpuras-— cens, and C. robustus, the last with an epithecium "lacking or inconspicuous," and with more robust asco- mayulayswand= “somewhat, donger » spores. The) collection Bronte, Gomera, — ceporied by iBanaress Baudet : Pr. a % Mes ann he tay RE EE ey JL GNG hs eee id Ms as SES Tee eee ore EE or MEST “ Rar pater 416 long, usually branched, gelatinous, sparsely septate, con- stricted by blackish septum. at its base. Lateral append- ages 26-52 um long, simple or variously branched, gelati- nous, extending upward, arising from triangular, subsessile cells formed at one or both upper corners of receptacle cells, these appendages sometimes producing one or more antheridia terminally. Simple rhizoidlike appendages 180- 350 um long, somewhat rigid, extending downward from more or less cubical cells in suprabasal cluster of receptacle cells. Antheridia ca. 9 um long, evanescent, gelatinous, solitary or clustered, produced directly upon somewhat tri- angular basal cell or borne on short, one-celled branch. Perithecia 60-79 x 24-36 um, ovate, usually curved outward and truncate at its apex. Holotype: TAIWAN. Nantou Hsien, Chitou, at ca. 1300 m alt., No.w)/83,) May 293) 19V7. Keeleradallec..20n elytrago. Osorius formosae Bernhauer (Coleoptera, Staphylinidae). Paratypes: TAIWAN. Nantou Hsien, Chitou, Nos. 782, 7/84, 786-790, data as for the holotype; Nantou Hsien, Tsuifeng, Nos. 791, 792, July 25% 1972) (Kew leradagles. .oOngvaktous parts of body of Osortus formosae Bernhauer. Osortus fotmosae occurs in mountainous regions of Tai- wan, where it lives in rotten wood. It is often found with Prtochtrus stlvestrit Bernhauer, host of Monotcomycés leptochtrt Thaxter (Terada, unpublished). Fichtatully mature specimens of Osortomyces and more than ten immature ones were collected from thirteen specimens of Osortus formosae. Collection of additional specimens would be necessary for a thorough study of thallus development. After spore germination, a series of horizontal divi- sions takes place that results in the formation of a uni- seriate thallus having a darkened, constricted septum near its apex, which may break off early (Fig. 1). Vertical divisions soon occur in the lower cells of the thallus, resuiting in the formation of a biseriate lower. receptacle CEL mel The number of cells in the thallus increase so that in the thallus shown in Fig. 2, there are eight super- posed cells separating the constricted, dark septum from the cluster of cells just above the basal cell. Rhizoid- like appendages (ra) extend outward just above the base of the thallus, whereas gelatinous, erect appendages (la) arise laterally at a higher level. The primary axis of the thal- lus terminates in a cluster of gelatinous appendages (Figs. 2-4) (it was not possible to determine the exact position of the spore septum, which separates the primary appendage 417 from the receptacle). Byamaturitysetive on six. simple, long, rhizoidlike appendages are present that extend down- ward from the suprabasal group of cells in the primary axis cs / sald). Triangular, subsessile cells are formed on one or both upper corners of the upper cells of the re- ceptacle. Each of these cells produces a short, gelati- nous branch. Secondary axes that arise from the upper part of the primary axis (two are indicated in Fig. 17) may deteriorate later (Fig. 18). The positions of the perithe- cia are variable; however, the lowermost one usually occurs on the suprabasal portion of the primary axis (Fig. 18). In the material examined, one specimen bore gelatinous, unicellular, elongate structures directly on triangular, subsessile cells (Fig. 5) or in clusters on a stalk cell produced by a subsessile cell (Figs. 5, 6). Although sper- matia could not be detected, these structures are regarded as simple antheridia because they are very narrow apically. Consequently, Osortomyces belongs to the Laboulbeniaceae. The antheridia seem to cease their function in a short time; possibly, they are converted into sterile appendages (Fig. Dea GLOwraleloeht shies ou) AlO)ie During perithecial ontogeny in Osortomyces rhtzophorus, a single cell (a') that arises laterally from the recepta- cle (Fig. 9) soon divides obliquely into an upper (d) and aelower celli'(c) (Fig. 10). The lower cell (c) divides PiCowanwupper cel lacG )mandhaw lower, Celle (ce) (FhTo bn), which undergo further divisions and finally form the peri- ehecial iwalls. The upper cell (d) produces the female sex- ual organ, which initially consists of three superposed cells - a lower carpogenic cell (f), a median trichophoric Pou (em) ew andeal terminals trichogyne, (tr) (Fig. 12). Only one— or two-celled trichogynes were observed. By the time there are three tiers of outer wall cells, an ascogenous cell (ac) and three supporting cells (ss, ist, is) are pres- ent below the trichophoric cell (e") within the perithecium (higt 3). This pattern of perithecial development is the Same as that described in detail by Thaxter (1896) for Stigmatomyces baert (Knoch) Peyritsch and other taxa. By the time perithecial wall consists of four tiers of cells, asci have been formed (Fig. 16). The second tier of cells in the fully mature perithecia is 3-4 times taller than the lowest tier (Fig. 14). Sometimes the apical out- er wall cell on the inner side of the perithecium (toward Ene thallus) axis) divides (Fig. 15). The stalk cells and the basal cells are strongly appressed to the receptacle so that the perithecia are sessile in appearance (Figs. 14, 15). 418 The genus Osortomyces cannot be allied with any genus of the Laboulbeniales except Scelophoromyces Thaxter (1912; in 1931, Thaxter altered the spelling of this genus to SkeLophoromyces, but the earlier spelling should be used). Osortomyces resembles Scelophoromyces in the following ways: 1) the primary axis consists of a variable number of super- posed cells; 2) rhizoidlike appendages arise at right and left from the suprabasal portion of the primary axis; 3) lateral appendages extend upward from one or both upper cor- ners of receptacle cells. On the other hand, remarkable differences exist between the two genera: 1) the perithecia of Osortomyces are normally sessile and lateral, whereas in Scelophoromyces, they are long-stalked and never arise di- rectly from any cell of the primary axis; 2) the primary axis of Osortomyces is highly divided in the suprabasal por- tion, whereas in Scelophoromyces, the suprabasal portion is less divided or undivided; 3) lateral appendages of Osorto- myces are gelatinous and the protoplasts of their short, subsessile basal cells taper almost to a point apically, whereas in Scelophoromyces, they are rigid like the rhizoid- like appendages and the protoplasts of their basal cells are broad at the apex. ACKNOWLEDGMENTS I wish to thank Dr. I. I. Tavares, University of Cali- fornia, Berkeley, for reviewing the manuscript as well as for helpful suggestions and Mr. Y. Shibata, Tokyo, for identification of the ,jhost insects. LITERATURE CITED Thaxter, (R.as1890; Contribution towards a monograph of the Laboulbeniaceae. Mem. (Amer soAcad Ss 7AntsaSc in lZ sao L298 Pls. I-XXVIL. Pe tie Pap New or critical Laboulbeniales from the Argentine. ProcsvAmern. (Acad. Arts @SCint 46 2553-22, LOS Ts. Contribution towards a monograph of the Laboulbeniaceae. Payiaas Mem. Amer. Acad. Arts Sci. 16: ,1-435% Bboy lak. af MYCOTAXON Moles WiyeNO.e25 pp. 419-427 July-September 1981 THE GENUS CODINAEA. THREE NEW SPECIES FROM THE AMERICAS ADRIANNA D. HEWINGS Department of Plant Pathology, University of Illinois, Urbana, IL 61801 J-L. CRANE Illinois Natural History Survey and the Departments of Botany and Plant Pathology, University of Illinois, Urbana, IL 61801 Three new species of Codinaea Maire are described and illustrated from the U.S.A. and Venezuela. A key modified from Hughes and Kendrick (3) to include those species described since 1968 is also included. Codinaea illinoensis Hewings & Crane SD.enOVs ieee. A and B. Coloniae effuse, pallide brunneae, demum fusco-brunnescentes. Mycelium in substrato immersum, e hyphis crasse tunicatis, ramosis, septatis, subhyalinis vel pallide brunneis compositum. Setae nullae. Conidiophori macronemati, mononemati, simplices, recti vel paulum arcuati, septati, hyalini vel pallide brunnei, tunicis crassis, levibus induti, singillatim e hyphis vel cellulis nodulosis orti. Cellulae conidiogenae monophialidicae, integratae, terminales, paulum lageniformes, apicem versus tenuiter, prope basim crasse tunicatae, 30-37 x 5.6-6.7 um. Collarula conspicua, infundibuliformia, expansa, marginem versus tenuiter tunicata, ad basim crasse tunicata, refringentia, 2.3-3.3 x 3.3-4.4 um. Phialoconidia hyalina, continua vel nonnumquam uniseptata, paulum arcuata, symmetrica, ad apicem attenuata in finem obtuse rotundatum setulam rudimentarium apiculatum exhibentem, ad basim attenuata, cicatrice obtusa notata, 14.4-18.9 x 3.3- 4.4 um. Typus: ILLS 42169. 420 Colonies effuse, pale brown becoming dark brown. Mycelium immersed see the substrate, composed oD thick-walled, branched, septate, subhyaline to pale brown hyphae. Setae absent. Conidiophores macronematous, Fig. 1. Codinaea illinoensis, A. Conidiophores; eae a ee B. Mature Conidia. of A21 mononematous, simple, straight or slightly curved, septate, hyaline to light brown, thick-walled, smooth, arising Singly from the hyphae or nodulose cells. Conidiogenous cells monophialidic, integrated, terminal, slightly lageniform, thin-walled at apex, thick-walled at base, 30-37 x 5.6-6.7 um. Collarettes conspicuous, funnel- shaped, flaring, distally thin-walled, thick-walled and refringent at base, 2.3-3.3 x 3.3-4.4 um. Phialoconidia hyaline, unicellular or occasionally 1-septate, slightly curved, symmetrical, tapering to a bluntly rounded distal end bearing a rudimentary apiculate setula, basal end tapering with a blunt scar. Holotype: On decayed wood, cypress swamp, N.W. of Vienna, Johnson County, Illinois, 9 July 1969, J.L. Crane 98-69, ILLS 42169. Codinaea lunulospora Hewings & Crane sp. nov. Fig.2, A-F Coloniae effusae, pallide demum obscure brunneae. Mycelium in substrato immersum, e hyphis crasse tunicatis, ramosis, septatis, subhyalinis vel pallide brunneis compositum. Setae nullae. Conidiophori e basibus sculptis orti, macronemati, mononemati, simplices, recti, septati, obscure brunnei, crasse tunicati, leves, ad phialidis basim paulum attenuati. Cellulae conidiogenae phialidicae, integratae, terminales, lageniformes, ad apicem tenuiter tunicatae, hyalinae, crasse tunicatae, subhyalinae vel pallido brunneae ad basim. Collarula conspicua, infudibuliformis vel aliquantulum cylindrica, apicem versus tenuiter tunicata, sympodialiter semel vel bis instaurata, 3.6-4.5 x 1.8-2.7 um, prope basim ad 1.4-1.8 um attenuata. Phialoconidia hyalina, continua, lunata vel sigmoidea, symmetrica, 8.8-12.0 x 0.8-1.0 um. Holotypus: Dumont-VE 4619 (NY). Isotypus: (VEN). Colonies effuse, light brown becoming dark brown. Mycelium immersed in the substrate, composed Of thick-walled, branched, septate, subhyaline to light brown hyphae. setae absent. Conidiophores arising from sculptured bases, macronematous, mononematous, Simple, Straight, septate, dark brown, thick-walled, smooth, tapered slightly at base of phialide. Conidiogenous cells Pphialidic, integrated, terminal, lageniform, thin-walled and hyaline at apex, thick-walled, subhyaline or light brown at base. Collarettes conspicuous, funnel-shaped to somewhat cylindrical, distally thin-walled, developing sympodially with 1-2 successive proliferations, 3.6-4.5 x A422 ’ Codinaea lunulospora, A-E. Conidiophores ue Fig. Mature Conidia. Ee af 423 1.8-2.7 pm narrowing to 1.4-1.8 um at base. Phialoconidia hyaline, unicellular, crescent-shaped or Sigmoid, symmetrical, 8.8-12.0 x 0.8-1.0 um. Holotype: On unidentified herbaceous stem. Trail between Manacal and Los Pocitos, N.W. of Irapa, Edo Sucre, Venezuela, 10 July 1972. K.P. Dumont et al., Dumont VE-4619 (NY), Isotype: (VEN). Codinaea matsushimae Hewings & Crane sp. nov. Fig. 3, A-D Coloniae effusae, pallide brunneae, demum obscure brunneae. Mycelium in substrato immersum, e hyphis crasse tunicatis, ramosis, septatis, subhyalinis vel pallide brunneis compositum. Setae singillatim vel gregatim ortae, rectae, crasse tunicatae, leves, ad vicies septatae, ad basim obscure brunneae, ad apicem obtuse rotundatum pallide brunneae vel subhyalinae, usque 340 um longae, e basi 6.7 um lata ad apicem 4.4 um lata attenuatae. Conidiophori e cellularum nodo setarum socio orti, macronemati, mononemati, simplices, recti vel paulum arcuati, septati, hyalini vel pallide brunnei, crasse tunicate, leves. Cellulae conidiogenae phialidicae, integratae, terminales, hyalinae, paulum lageniformes, ad apicem tenuiter, ad basim crasse tunicatae, 14-27 x 3.0-6.0 um. Collarula conspicua, evanescens, ad marginem tenuiter, ad basim abrupte crasse tunicata, synpodialis, semel, bis, vel ter instaurata, 2e0=3-0. x 2.0-4.0 im. Phialoconidia setulis ornata, hyalina, ter ver raro semel vel bis septata, paulum arcuata, symmetrica, ad apicem obtuse rotundatum et basim cicatrice notatum attenuata, 20-27 x 3.3-5.4 um. Setulae alia ad apicem, alia juxtam cicatricem basalem affixae, 5.5-7.8 um longa. Typus: ILLS 42170. Colonies effuse, light brown becoming dark brown. Mycelium immersed in the substrate, composed of thick-walled, branched, septate, subhyaline to light brown hyphae. setae arising singly or in groups, erect, thick-walled, smooth, up to 20-septate, dark brown at base, pale brown to subhyaline at bluntly rounded apex, up to 340 um long, tapering from 6.7 um wide at base to 4.4 um at apex. Conidiophores arising singly from a knob of cells associated with setae, macronematous, mononematous, simple, Straight to slightly curved, septate, hyaline to light brown, thick-walled, smooth. Conidiogenous cells phialidic, integrated, terminal, hyaline, slightly lageniform thin-walled at apex, thick-walled at base, 14-27 x 3.0-6.0 um. Collarettes conspicuous becoming evanescent, 424 Codinaea matsushimae, A-C. Conidiophores and Ficw 3, D. Mature Conidia. setae; 425 distally thin-walled, abruptly thick-walled at base, Ssympodial with 1-3 proliferations, 2.0-3.0 x 2.0-4.0 um. Phialoconidia with setulae, hyaline, 3-rarely 1 or e-septate, slightly curved, symmetrical, tapering toa bluntly rounded distal end and a proximal basal sear, 20-27 x 3.3-5.4 um. Setulae attached at distal end and to one side of basal sear, 5.5-7.8 um long. Type: On exocarp of Carya, Benson’s Bluff, S.E. of TELINOLS seeeteh prt les 9095) elas Other Material Examined: On Cypress Swamp (Deer Pond), Johnson islet ea isl Crane, 91-69, ILLS Gorevill, Johnson County, Crane 12-1-69, ILLS 42170. decayed wood, Elvira County Illinois, 22 May 42171. Key to the Species of Codinaea Maire* 1. Conidiophores synnematous ......... C. obesispora (3) Conti da ODNOneSHMOnenema cousin. sisrcisisiciclele sicleicis + + 01s sl eless sane Zs Piva loconid Lamwit nese UULAC mets crclescholeie ts cherelelots otslekete erecennes Pntaroconid ramwschOUGMSeCULACEI cr culstsisisiclietslc sloisiere siete sme’ B Phialoconidia with a setula at eachvend 2... ....e-+n0 Phialoconidlamwichiel api Ca larse Cu lasts cle seis lotta ice leleaee 4, Setula definite, 13-15 um long ..... C. unisetula (7) Setula apiculate eeoseeeoeeeeweeeepeeeeweeeseseeeeeeeoeeevseeee see @ 5 5. Phialoconidia 14-19 x 3.3-4.4 um. C. illinoensis Phialoconidia 20-30 x 4.0-5.5 um ... C. apiculata (6) b PHPAMOCOM MOAT ARSE DLOUEC iW elels siclstcl sisicle.ctsisie.e sie eitte oiebetetaioieioue, Rita bOCOnT di amnONn=SEDLALCmrleisse cles sic sisle cs cleiets slo oie sss uec iemeesetulae w250 Uma LON Rass alstcis com cic stele cciete rented er here mL O Setae slightly geniculate at apex ... C. fertilis (3) Setae straight, cyclindrical to gradually Caperingsaceapex tei. ole e ec views Cemascam Camus) Setulae of unequal length ........... C. aristata (4) Setulae equal in length ..cccccccccccccees Setutacv5-9 WmulOne@ae vse le eels ocr Dr Evanicam ain Setulaepwup .UOuil « Onw Mal ON it ote ols cuss ecko lelelel oie cieteietetecetels Chaetosphaeria pulchriseta anam. Codinaea (3) Setulae’ 1 ComDAnVaAmC 3) Phialoconidia 17-23 um long ......... C. vulgaris (3) Phialoconidia with acute apex, eoeceveveoeeree vee ee ee © © 6 19 Pare eooeoevevreecee oe eere eee eee ee eee ee oe oe Lad, gonytrichodes (10) espe 0016, 6 cole elec eleraie cre .o MELO eeoeeeoeeeeseftfeeeseeeee @ @ @ C. ooo oe 8 @ @ 1 23-28 um long e@eeveeeveveeve eee ee ee © © C. longispora (3) Phialoconidia 14-19 um long .... Phialoconidia Phialoconidia Phialoconidia Phialoconidia Conidiophores and ornamented ..ccccseeseeveee Co Conidiophores Phialoconidia with bluntly pointed apex, sessecsersoeceovce Cs Simplex: (35) SCPCAL!S covcccccccvervsersescsescecees OF NON=SCDtLALEL <.c.0 1c cies creretenele cit 6 clets ele seeeO B=SEptate tes cece seme map LCdl Smo T~Septateiic sc cists oles cc cistere seeleteiote aioe samc branched glauco-nigra (3) SIMPLE And “SMOOUNete sietctetetele cic cl etetereiatel aac 10s 169 Xt 225=5. 5 msec eC ena Lam) Phialoconidia 20-24 x 2.0-2.8 um ...... C. setosa (3) Conldlopnores] Drancnedigs steterct ele cletsiclelc’c cleteleteretere! erence Conidiophores simple @eeoeseeoeeeeseese@eeeeseerpeeeeeeeet eee 6 @ @ @ 28 Phialoconidia Phialoconidia 20=26 um long ......... Co hughesi ind) 10—15 um long eoeeevoeveneeeeveee eee eee 8 & Chaetosphaeria callimorpha anam. Codinaea (3) Phialoconidia Phialoconidia Phialoconidia Phialoconidia SCGONE ya CUIVECS « cicicleisie c1e.sipis sieteisiete omc straight or slighly curved ......-.. 30 Sigmoid ............ C. lunulospora botuliform’......... C. botulispora’ (3) Phialoconidia cylindrical ™..... C. ~ eylindrospora Gp) Phialoconidia tapering towards base Mtn eens) Phialoconidia abruptly tapered AL DAS] ccccecccccssccoesesceveees Co Clavulata (2) Phialoconidia gradually tapered at base .........6. Chaetosphaeria talbotii anam. Codinaea (3) 427 Literature Cited webs sao. 5. 1976. More dematiaceous Hyphomycetes. Commonwealth Mycological Institute, Kew, 507 pp. - Gams, W. and V. Holubova-Jechova. 1976. Chloridium and some other dematiaceous Hyphomycetes growing on decaying wood. Studies in Mycology No. 13, Baarn, 99 Pp. Hughes, S. J. and W. B. Kendrick. 1968. New Zealand fungi 12. Menispora, Codinaea, Menisporopsis. New Zealand J. Bot. 6: 323-375. Maire, R. 1937. Fungi Catalaunici, Series alterna. Contribution a 1’etude de la Flore Mycologique de la Catalogne. Publcions Inst. bot. Barcelona 3: 128 Pp. Matsushima, T. 1971. Microfungi of the Solomon Islands and Papua-New Guinea. Pub. by author, Kobe, the) We Matsushima, T 1975. Icones Microfungorum 9 a Matsushima Lectorum. Pub. by author, Kobe, 209 pp. Morgan-Jones, G. and E. B. Ingram. 1976. Notes on Hyphomycetes XV. Two new species of Codinaea. Mycotaxon 4: 504-509. RIPOZyYNSKis Ka tA. sand S. D. Patil. 1970. Some setose Hyphomycetes of leaf litter in south India. Ganadiamtd .meeBOU. P4056 507—50 16 Reddy, S. M. cheyel | Sr S. 1978. A new species of Codinaea Sydowia 30 (1-6): 186-188. Shearer, | Ce A. Andee ot BOL ANC mero (clea Ul e1ero fy the Chesapeake Bay and its’ tributaries. I. Patuxent River. Mycologia 63: 237-260. Sutton, Bb. .Cs and’. 6S. Hodges, 91975. Eucalyptus microfungi: Codinaea and Zanclospora species from et Brazil. Nova Hedwigia 26: 517-525. MYCOTAXON Vol. XIII, No. 2, pp. 428-430 July-September 1981 A NEW SPECIES OF DACRYOPINAX FROM BRAZIL B. Lowy Botany Department, Louisiana State University Baton Rouge, LA 70803 Dacryopinax maxidorii Lowy, sp. nov. Fructificatio in humido aurantiolutea, elastico- gelatinosa, pileata, 4.5 cm alta ad 5 cm lata; sicca cornea, ferrugineo-brunnea; caulis elongatus, firme radicatus, conspicuus, crassi-venatus; in sicco dense albido-tomentosus; apicis alte ramosus, deinde poly- cephalus, lobuli flabelliforme ad 450 um crassis cum mar- gine crenulata; hymenio glabrum, unilaterali, inferiori, pauci pilosa; probasidia cylindraceae, aseptata, 30.0-36.5 X 4.5 um, metabasidia furcata, aseptata, bisterigmata; hyphae enodosae, 2.0-3.0 um diam; basidiosporae leviter | curvulo-cylindraceae vel subovoideae, uniseptatae, in cu- mulo aurantiaco-lutea, (7.0-) 8.0-10.0 (11.5) X 4.5-5.0 um, per promycelium aut conidia germinantes; conidia sub- spherica, 1.5-2.5 um diam. Fructification when fresh rubbery gelatinous, orange- yellow, stipitate, pileate, up to 4.5 cm in height, 5 cm broad (Fie. 1). drying horny, rusty brown; marisinge: roma broad, elongated, firmly rooted, sterile stalk; thick veined, densely whitish-pilose when dry, with pilosity diminishing upward; deeply branched apically, frequently becoming polycephalic with broad, veined, flabelliform lobes + 450 um thick, often with crenate margins; hymenium unilateral, inferior, producing a dense palisade of basidia; abhymenium thinly covered with hyaline, cylin- drical to slightly inflated, unbranched hairs, unicellular to sparsely septate, arising from a layer of irregularly inflated hyphae; probasidia cylindrical, unicellular, 30.0-36.5 X 4.0-4.5 wm; occasional cylindrical, slender, unbranched dikaryophyses 20-25 X 2.5-3.0 um; metabasidia furcate, bisterigmate; hyphae without clamp connections, 429 2.0-3.0 um diam; basidiospores slightly curved-cylin- drical to subovoid with narrowed apices and prominent apiculus, with a single, thick, central septum, (7.0-) 8.0-10.0 (-11.5) X 4.5-5.0 um, producing subspherical conidia 1.5-2.5 um diam, or germinanting by germ tube. Holotype, Brazil. Lowy 190 BR (TYPE), km 405 Manaus- Porto Velho road, Amazonas, 16-I1X-1980. Leg. 8B. Lowy, D. Coelho. On unidentified log. This species is named for Maxine and Doris Lowy, enthu- siastic collectors of tremellaceous fungi. Figure 1. Basidiocarps of Dacryopinax maxidorii with scale shown in millimeters. In gross morphology the new species somewhat re- sembles D. indacocheae Lowy (1959), except for the lat- ter's stubby stalk, thinner lobes, and brownish to earth- colored pigmentation. However, the long stalk, bright carotinoid pigmentation, and flabelliform lobes of the new species places it closer to D. martinii Lowy (1971). The basidiospores of these three species are predominantly l-septate, but D. maxidorii shares with D. indacocheae somewhat curved-cylindrical spores with a single, thick septum whereas the spores of D. martinii are character-— istically thin septate. 430 The following revised key includes the seven known species of Dacryopinax occurring in tropical America. 1 Basidiocarp slightly expanded apically; pileus simple r,orvbranched (en) (Pete Woe CaN mer eeee sy ut nee 2 2. Stipe tomentose; basidiospores becoming septate, en. ea. oe I D. spathularia 2 obi pemno tu coment ose: Reet deepens: becoming S=SeP Cates eer Uh ie ere eee: ; Deedennisit Basidiocarp broadly expanded apically; pileus gross- ly lobed to ftlabelln form onpepolycephalice.. =... 93 3b. slongestipmtateseplleuse:)abel 1. Orn: ee ee 3. Short stipitate to substipitate; pileus not Flabelli form. ges ean ee ells Eh cre ode ee een aE 4, Abhymenium and stipe smooth; dried pileus thin, fraci lev wen. ; Dinas Gd nda 4. Abhymenium and Eoime miteeer Farid pileus thick, LODUS ET wi pee omen a ens eerie Detmas ident 5. Hymenium coarsely Penis yellow when bresh yt mass ee: D.- yungensis 5. Hymenium sAereiel va ec when fresh . 6 6. Pileus spreading-orbiculate, undivided, drying dark brown to blackish. . ; : D. elegans 6. Pileus foliose, SSP Toned drying light Jopeeymaaie isos (ie Pa : D. indacocheae ACKNOWLEDGEMENTS Thanks are due The New York Botanical Garden, the Instituto Nacional de Pesquisas da Amazonia (Manaus, Amazonas, Brasil), and Louisiana State University (Baton Rouge, LA), under whose sponsorship the field work was completed as part of the Projecto Flora Amazonica. LOWY sab. LITERATURE CITED 1959. New or Noteworthy Tremellales from Bolivia. Mycologia 51: 840-850. 1971. Tremellales. Flora Neotropica. Mono- graph 6, Hafner Publishing Company, Inc. New York. MYCOTAXON OMX UTS UNO meee AOL July-September 1981 NEO Sie is Car THIRD INTERNATIONAL MYCOLOGICAL CONGRESS IMC. will be held in Tokyo, Japan, from 28 August till 3 September, 1983. Several affiliated organizations have indicated they wish to be involved in the organization of the program, particularly with regard to symposia and the provision cf time for business meetings they may wish to hold. Possible titles for symposia, with names and addresses of potential organizers, and de- tails of any other suggested activities should be submit- ted as soon as_ possible to the Organizing Committee (Secretary General, Prof. K. Tubaki, Institute of Biolo- gical Sciences, University of Tsukuba, Sakura-mura, Ibaraki 300-31, Japan). A copy of any letter should be sent to the Secretary of the International Mycological Association, Dr. D. L. Hawksworth, Commonwealth Myco- logical Institute, Ferry Lane, Kew, Surrey TW9 3AF, England. Suggestions should be submitted as_ rapidly as possible. 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It is a convenience to typists, but certainly not an essential, since rectangles of the appropri- ate size can be prepared on any paper using a non-photoreproducing blue pencil. Each package of 50 sheets is sent postpaid for $2.20 US. BIOPLATE is a special sheet of transfer letters for the use of authors in the preparation of plates and graphs for publication. It is manufactured for us by Prestype, and available in either black or white. Each sheet is approx- imately 30 X 39 cm., with a wide assortment of numbers, letters (including important Greek characters), symbols, etc. These are sent postpaid for $4.95 US (black will be shipped unless white is specified). eT Les i ‘ ok at be » MYCOTAXON AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS Volume XIII October-December 1981 NO CONTENTS Studies in the lichen genus Psoroma I: Psoroma tenue and POPC CLIAINOME WI 6 6a ass oS ae od ele e's A. HENSSEN AND B. RENNER 433 An undescribed pleomorphic species of Codinaea... NORIHIRO TOYAZAKI 450 The psilopezioid fungi. VIII. Additions to the genus Pachyella. DONALD H. PFISTER AND FRANCOISE CANDOUSSAU 457 Pithomyces pavgitt, a new combination for Trtchocladtum pavgtt and Pithomyeces funtculosa...... MARY E. PALM AND ELWIN L. STEWART 465 Validation of the Herpomycetineae and Herpomycetaceae in the SPUD ERE ALCS 2505s pacn, PURSE Taree ete, wee it UG ake Si ISABELLE I. TAVARES 469 Erynta (Zygomycetes: Entomophthorales): validations and new PRES Wh et A ESP ran Bit Sie Aisa sats umes Ie Whee at alti nie foegia.y ae wie RICHARD A. HUMBER 471 A review of the nonentomogenous Entomophthorales... BRUCE E. TUCKER 481 Erynta (Zygomycetes: Entomophthorales): emendation, synonymy, Mane Transfers. 3 ih sila. aes RICHARD A. HUMBER AND ISRAEL BEN-ZE'EV 506 eS LINTOS Aro Wo iia eee ia eee ahs ee G. L. HENNEBERT 517 Notices Pere DI ce oCTence, CONPTESS., Klis is oe) oh Cite onin date whine se ea wiea An ples 525 IMC: A SA ECOS DS TBA ANAT o suuia vy novi Gerns fe elas bow Bhs pi wel ee! ay date Sova aby 525 PEE a ENG pte seed i oa og a bs oe ae Wee cp aie, She Wg AhG Gra sino a ine Bia © et bMeeW tale 526 MEE TROIS GL LCNCMOLAXAGL iui. Se ice Oo mk lh oye Mieipid ote Wide Wego: 8 528 ewen.ON publication dates 712 (2), 13(1) ;.and 13. (2). oo. elk. 534 See ee AN ee Te, SMU Ae lia tis Who Re es iehteed oe Wo ee Re ele meen 535 RE DR eae ias Becht EG tole th hy LCL ely) TUE: plate Vepa aia aA ba Sep Epo Ro edodel ic ovpnohnnc of JD [MYCOTAXON for July-September 1981 (13: > MANN was issued July 1, 1981] atBERT RM LID Ie ITHACA, N. Y. 14850 NOV 02198) ISSN 0093-4666 MYXNAB ST (3) 493-5 364( 19819 Library of Congress Catalogue Card {fumber 74-7903 Published quarterly by MYCOTAXON, Ltd., P.O. Box 264, Ithaca NY 14850 For subscription details, availability in microfilm and microfiche, and availability of articles as tear sheets, see back cover a ue - t . ¥ yi 7 pee Fs ah ; 7. Ay ry -? - —- Pd aes ee MY COTAXON eee Pete ALTE NO. 3, pp. 433-449 October-December 1981 STUDIES IN THE LICHEN GENUS PSOROMA I PSOROMA TENUE AND PSOROMA CINNAMOMEUM A. HENSSEN and B. RENNER macnberevehaBiologie ‘der Universitat Marbung D-3550 Marburg/Lahn, Germany SUMMARY The new species -Psoroma tenue Henssen belongs to the P. hypnorum-group and is most closely related to P. cinnamomeum. Two varieties are described. P. tenue var. tenue is restricted to the Southern He- misphere; the lichen has been collected in Tierra del Fuego and the South Shetland Islands. P. tenue var. borealis Henssenis a wide-spread northern ta- xon; localities are given for the Alps, Scandina- via, Iceland and North America. P. tenue is a spe- cies of snowbanks, glacial till and tundral heath. The lichen contains porphyrilic acid methyl ester and pannaric acid. The occurrence of these meta- bolites in other members of the P. hypnorum-group is mentioned. A short description of anatomical structures in P. cinnamomeum is included. INTRODUCTION The lichen genus Psoroma Michx, a member of the Pannariaceae, occurs predominantly in the Southern Hemisphere, in South America and New Zea land. The type species P. hypiorum (Vahl) S.F. Gray (Synonyms: P. bryanttt Dodge as "Bryantt" in imac aeAmMersrM1 crOoscop. Soc. 842511719055. holos type: hb. Dodge; and P.follmannit Dodge in Trans. aie wee Croscopm cOC. 164: 51.0,)a196500 1SOLY pe has SEL) only is known to be world wide distributed. 434 Our studies in Psoroma are mainly restricted to South American species, based on collections made by A. Henssen and G. Vobis in southern Chile and Argentina in 1973/74. P.W. James who was planning a monographic treatment of the genus encouraged us to collect Psoroma species intensively. The series of Psoroma papers, pub- lished in part together with P.W. James and K. Marton, may be regarded as preliminary studies LOX a world monograph ings thtseini Lid apapon we... Henssen is responsible for the taxonomic treat- ment; and B. Renner, for the identification of the lichen substances. InvLeS CuUrrentecirCcumMSeHl pELOnge se rCngm smd heterogenous genus, and affinities of certain species to Pannarta Del. in Bory have been sug- gested (James & Henssen 1976, J¢rgensen 1978). The species of the P. hypnorum-group, for example, form a distinct assemblage of terricolous lichens of cold temperature or subalpine / alpine areas. They may be more closely related to Pannarta pestzotdes (G.H. Web.) Trevis. than to the corti- colous / saxicolous, squamulose or lobate species of Psoroma with which they are at present allied (James & Henssen 1976). Thesnew) Species WP cucu De lLOnGgSme tL Omstlicomne. hypnorum-group. The outstanding character of the species is a pattern of lichen substances not previously known) to soccuGn in Vesorengus.. me ne specimens of P. tenue so far studied always con- tain pannaric acid and the new compound porphy- rilic acid methyl ester and some specimens con- tain an unknown substance (U,) (Renner et al. 1981). Specimens from the Northern Hemisphere differ slightly in morphology fromethe “southern hemispheric ones and are included in the variety borealts. In P. hypnorum, no lichen substances were found by TLC (J¢grgensen 1978). We confirmed this observation in the type specimen (C) and in the types of the synonymous species P. bryantit (hb. Dodge) and P. follmanntt (KASSEL) as well as ina number of our specimens from northern and southern 435 localities. All of these specimens were similar with regards to external and internal morphology. However, certain specimens, deviating to some extent in morphology, were found to contain one, two or all three of the compounds described from P. tenue. These specimens need further taxonomic Pavestigation. Porphyrilic acid methyl ester and mennaric acid also occur in the new P, rubro- marginatunP. James & Murray (Henssen et al. in manuscript) , and the unknown (U, ) occurs ina South American specimen of P. paleaceum (Fr.) Nyl. (2? P. hirsutulum Nyl.) (Henssen 24353g: MB). Within the P. hypnorum-group, several patterns in thallus anatomy may be recognized. P. tenue corresponds to P. ctnnamomeum in external and internal morphology of the thallus and cephalodia: The two species, therefore, are regarded to be closely related. Although the South American species P. etnnamomeum was well - described by Malme (1952), additional information obtained by modern methods is included in the presented paper. MATERIAL AND METHODS MATERIAL. The abbreviations of herbaria follows those given in Index Herbariorum; hb. Dodge means the private herbarium of C. Dodge, Burlington, Vermont, U.S.A., and hb. Henssen means the private herbarium of the senior author. Psoroma cinnamomeum: Argentina, Tierra del Fuego, Rio Grande, 1896, Dusén 115 (holotype: S); Cabo San Pablo, on plant debris in a tundra heath, overgrowing a dead liverwort, 1973, Henssen & Vobis 24448t (MB). P. hypnorum (selected specimens): Norway, Vahl (lectotype by Jgérgensen: (C). Sogn og Fjordane, Nordfjordeid, on rock overgrowing mosses, 1959, Henssen 5339 (hb. Henssen). - Finland, Lap- -ponia enontekiensis, Toskaljarvi, 1955, Henssen 6145 (hb. Henssen). - Argentina, Tierra del Fuego, Glaciar Martial, on rock overgrowing mosses, 400m s.m., 1973, Henssen & Vobis 24412a (MB); Laguna Escondida, Henssen & Vobis 24439h and 24460d (MB). - Antarctic Peninsula, Refuge Is- land, 1940, Bryant 30-2 (holotype of P. bryantii: hb. Dodge). - South Shetland Islands, Greenwich Island, Bahia Soberania, on mosses, 1963, Follmann 11843 (isotype: 436 Fig. 1. Psoroma tenue var. tenue (paratype), young and mature densely aggregated apothecia; scale = 1 mm; drawn by H. Becker. of P. follmannii: KASSEL). P. paleaceum: Argentina, Tierra del Fuego, between Ushuaia and Lapataia, on rock overgrow- ing mosses, 1973, Henssen & Vobis 24353g (MB). P. tenue: see below. ef 437 MORPHOLOGY. Samples were sectioned by freezing micro- tome and the sections mounted in lactophenol cotton - blue. Measurements of spores and anatomical structures were made from permanent preparations; measurements of gross morpho- logy on air-dried specimens. For studying the ascus structure, KI, solution was added directly to the squash preparation. CHEMISTRY. Crude lichen extracts were chromatograph- ed in solvent systems A, B, C of Culberson (1972), and D, E of Renner (1980). The methods of mass spectrometry and UV/VIS spectroscopy were used for the identification of the lichen substances; details are given in Renner et al. WEOGI)= TAXONOMIC PART Psoroma tenue Henssen, sp. nov. var. tenue Hab) eb Ole cA Ore a | MismalalOny yen lOO ATe . ast Lue— Luresor the ascocarp, Hige4A-D. DIAGNOSIS. Thallus granulatus vel subsquamulosus, pallidus vel fuscescens, e verrucis formatus semiglobosis, circiter 0.2 - 0.4 mm latis vel e squamulis plus minusve adscentibus, .0-45— 078mm) latis, crassis,) dispersis vel subcontiguis. Thallus corticatus, hyphae medullosae reti- culum formantes, alga symbiotica laete viridis, ad fami- liam Chlorococcaceae pertinens. Cephalodia globularia vel ramosa, plus minusve aggregata, atropurpurea, corticata, alga ad Nostoc pertinens. Apothecia usque ad 2 (-3)mm lata, disco plano vel sub- plano, atrofusco, margo thallinus vulgo pallidus, tenuis, subtiliter crenulatus. Hymenium 95-120 (-150)um altum, Asci subcylindrici, 80-110 x 8-12ym, 8-spori, annulo amy- loidea in apice asci incrassato, Sporae eseptatae, inco- lores, polymorphae, ovoideae vel subfusiformes, 19-24 x (5-)7-10 (-12)um, superfice verruculosa. Pycnidia minuta, circiter 0.1-0.16 mm lata, conidiophora brevicellularia, conidia terminalia et lateralia formantia. Conidia bacil- bi formia, Circiter 2-4 x 1 um. Chemistry: thallus and apothecia contain porphyrilic acid methyl ester, pannaric acid, in the holotype the un- known substance (Uj) was found. 438 af 439 Holotype: Argentina, Tierra del Fuego, Parque Nacional de la Tierra del Fuego, Laguna Escondida, on loamy soil near the lake shore overgrowing Cephaloziella and plant debris, at c. 270m s. m. 1973 Henssen & Vobis 24439b (MB); isotypes: BM, LPS. Paratype: near the type locality at Hosteria Petrel, on soil overgrowing plant debris ina stand of Nothofagus at the lake shore, 1973 Henssen & Vobis 24460w (MB). FURTHER SPECIMEN EXAMINED: South Shetland Islands, Greenwich Island, on earth and plant debris, 1963, Follmann 14065 (KASSEL) . Thallus of pale or more rarely brown colour, dull, composed of dispersed or aggregated minute granules and squamules, fastened by rhizoidal hyphae with colourless or brown thick walls. Granules up to 0.4 mm, squamules 0.4 - 0.8 mm broad, more or less ascending. Thallus sections (2559>)) 150-250 (450) um high, at least on the upper surface distinctly corticated. Upper cortex in young thalli compo- sed by thin-walled cells (Fig. 3A), later on the walls gelatinizing, and the lumina in the gelatinous matrix up to 18um broad, connected by pits (Fig. 3B). Lower cortex composed of isodiametric or elongated cells (Fig. 3C). Medullary hyphae (2.5-)4-6um thick, orientated reticula- tely and enclosing single or several algal cells. Phyco- biont a member of the Chlorococcaceae, cells c.6-10pum broad, distributed throughout the thallus but more numer- ous in the upper part. Crystals of lichen substances de- posited in clusters in the medulla. Pig. 2. Habit photographs of Psoroma species. A-B, P. tenue var. tenue (paratype); A, lichen overgrowing plant debris; B, enlarged apothecia. C, P. tenue var. tenue (holotype) ; dark colored, aggregated cephalodia between thallus squamules and apothecia. D, P. cinnamomeum (holotype). E-F, P. tenue var. borealis (holotype); E, mature apothecia; F, cephalodia (arrowed) between squamules. G, P. cinnamomeum (holotype) , crowded apothecia surrounded by a thick thal- line margin; the black apothecia belong to a Badicia species. H, P. tenue var. borealis (hb. Henssen 6130), Squamules and apothecia. A-H, scale = 1 mm. 440 af 441 Cephalodia granulary or coralloidly branched, dark purplish brown, singly or forming clusters up to 0.8 mm (Fig. 2C). Cephalodia corticated, cortex 4.5-12yum thick, formed by 1 to 2 rows of cells; medullary hyphae orientated in a network in young stages, later forming pseudoparen- chyma (Fig. 5E). Alga a Nostoc species. Apothecia dispersed or closely aggregated, up to 2(-3) mm broad, disc flat, dark brown, surrounded by a pale, finely crenulated margin (Fig. 1, 2A, B). Hymenium 95-120 (-150)um high; hypothecium - layer including the ascoge- nous hyphae - 20-25(-50)um; subhymenial layer of varying thickness, frequently extending into a stipe of adglutinat- ed hyphae with enlarged cells towards the base. Margo thal- linus corticated, basal part 100-120um broad, composed of radiating hyphae with broad, up to 25um wide cells; joint walls between the cell lumina 3.5-6um thick (Fig. 4A, B). Crystals of lichen substances in clusters in the medulla of the margo thallinus. Pycnidia inconspicuous, in sections 0.1-0.16 mm large. Conidiophores branched and anastomosing, conidiogenous cells short, the conidia terminally and laterally produced. Conidia rod-shaped, c.2-4 x lpm. Psoroma tenue var. borealts Henssen var. nov. Poot eee, (so challus anatomy, /f19. 3D-F; seruc— Lure sOL apothecium, Fig. 5C. DIAGNOSIS. Differt a var. tenuethallo majori semper distincte corticato et colore cinnamomeo vel luteofusco. Chemistry: thallus and apothecia contain porphyrilic acid methyl ester and pannaric acid, in the holotype and in the specimens from Austria, Ost-Tirol and Canada the unknown substance (U,) was found. Fig. 3. Thallus anatomy of Psoroma species (microtome Sections). A-C, P. tenue var. tenue (holotype); A, 1.s. of young thallus, crystals of lichen substances indicated by arrow; B, t.s. of upper cortex and algal zone; C, t.s. of thallus adjacent to an apothecium. D-E, P. tenue var. bore- alis (Henssen 22679g); D, 1.s. of young thallus; E, 1.s. of a cephalodium at the base of an apothecium. F, P. tenue Var. borealis (Henssen 23157x), t.s. of corticated thallus. A-F, scale = 20 um. 442 ae 443 Holotype: U.S.A., Colorado, Boulder Co., Niwot Ridge, on snowbank, between 3300 and 3750 m, 1961, Henssen 13030b (hb. Henssen). Paratype: Austria, Tirol. Samnaungruppe, Zeblasjoch, on snowbank at 2450m, 1975, Henssen 22679g (MB). FURTHER SPECIMENS EXAMINED: Austria, Ost-Tirol, Matrei, Johannishitte, in glacial till at c. 2200 - 2400 mn, overgrowing mosses, 1938 F. & Koppe (hb. Henssen 6130). - Norway, S6r-Troéndelag, Dovrefjell, Knuthd, on snowbank at 1500 m, 1959, Henssen 4974 (hb. Henssen). - Sweden. Torne Lappmark, Abisko. Nuolja, on snowbank at 1160 m, 1954, Henssen 6127 (hb. Henssen). - Iceland, Bard, Kollsvik, overgrowing mosses in a lichen heath, 1972, Henssen & Kristinsson 23157x (MB). - Canada, Alberta, Banff National Park, Peyto Lake, Dryas heath at 2150-2250 m, 1962, Henssen & Cain 14471y (CAN, MB). Thallus squamules up to 1 mm broad, brown or reddish brown, thick; thallus distinctly corticated on the upper and lower surface by isodiametric cells (Fig. 3D, F). Apothecial disc brown and margo thallinus of the same colour, or disc dark brown and margo thallinus slightly paler. Crystals of lichen substances deposited within the medulla of the thallus and apothecial margin or not. Pie emecenuesvar. Lenue the lobes are smaller Brae mevatemlOneca, is (Fig. 92) ,andvthe cortex is usually less well developed since it is restric- ted to the upper surface. Broad cortical struc- tures on both thallus surfaces have been observed in thalli closely attached to margins of apothe- Slam fg. 56)r in such lobes the thallus corres= ponds exactly to that in P. etnnamomeum (cf. meGs. 3B with’ 5A). The cephalodia occur either between the lobes Or are producted in groups below the apothecia (Puge038).) the alga in the cephalodia is a Fig. 4. Apothecial structure in P. tenue var. tenue (micro- tome sections of the holotype). A, median 1l.s. of young apothecium, note the numerous algal cells in the stipe; B, cortex structure in apothecial margin; C, upper part of apothecial margin, crystals of lichen substances indicated by arrows; D, ascus and spores. A-D, scale = 20 um. species of Wostoc as is found in most Psoroma species. The margin of the apothecia develops in the Same Way as in ‘Pannaria rubtginosa (Ach.) Bury (Henssen 1969, 1981). The cortex in the margo thallinus is differentiated in part by the thal- lus cortex and in part by the medullary hyphae growing up between the hymenium and the thallus cortex (Fig. 4A, B). The cells formed by the me- dullary hyphae are considerable smaller in size (Fig. 4C, 5C). As seen from above the finely cre- nulated margin surrounding the relatively flat disc resembles that of certain Pannarta species. In very old apothecia the margin may bear granu- les and the disc becomes folded or medianly de- pressed. Young developing apothecia are at first urceolate. The ascus wall and hymenial gelatine stain blue in iodine, and an amyloid ring structure is seen in the ascus apex of the same shape as that Pound in 2. 2ypnorum (Keuck 1977). hie emer vee var. tenue yellowish to brownish crystals of lichen substances are usually depo- sited in large amounts in the medulla of the eoellus and apothecial margin. In var. borealis Similar crystals have only been observed in the specimen collected in Canada (Henssen 14471y ). miese crystals are difficult to demonstrate in the sections by black and white photography but are rather impressive in microscopic examination of stained permanent preparations. The varying occurence of the unknown substance (U,) in both varieties of the species is interesting. Fig. 5. Anatomy in Psoroma species (microtome sections). A-B, P. cinnamomeum (holotype); A, 1.s. of thallus; B, _ cortex structure in apothecial margin; C, P. tenue var. borealis (Henssen 22670g), apothecial margin; D, P. cinna- momeum (Henssen 24448t), cortex structure in apothecial Margin; &, ~. tenue var. tenue (holotype), 1.s. of cepha- lodium; F, P. cinnamomeum (Henssen 24448t), 1.s. of cepha- lodium. A-F, scale = 20 um. 446 Fig. 6. Psoroma cinnamomeum (Henssen 24448t), young apo- thecia; scale = 1 mm; drawn by H. Becker. PSOrOMma Lene mS GnanractenuZeds Dy eLnempEesence of porphyrilic adic methyl ester and pannaric acid’ in) correlation with, small, flatvanotnecia surrounded by a thin evenly crenulated margin, by dark purplish brown cephalodia, aggregated to clusters, and by an upper thallus cortex formed by periclinal hyphae with isodiametric cells. In .comparisoni toe. venue.) P. vetnnamoneumsnas a Similar anatomy and the same type of cephalo- dia but) differs eapartetrom stiews acwromeltecnen substances, by having a very thick apothecial margin in the young, strongly urceolate apothecia 447 and by the smaller lumina in the cells forming Pies COLGCxK@ ine thesmargo thallinus (cf. Figs, 4¢ Pom Gawiltlimb, mi )ree ect nnanomeumn OCCULS in the PameeDegioOns as) Ps venue Var. tenue. By the pale So OUImMOms ies thialius tand athe abundant crystals meapothecialior thallus sections, £. tenue var. tenue is easily distinguished from P. etnnamo- meum. Reddish brown specimens of P. tenue var. borealts resemble P. etnnamomeum but the variety is only known from the Northern Hemisphere at this time. ECOLOGY. P. tenue var. tenue has been collec- ted on soil, long covered by snow, together with Pe Aypnorum and P. pateaceum. P. tenue var. bore- alts has been found, hitherto, on snowbanks in glacial till and in tundral heath, where the li- chen was growing together with Lectophysma ftn- Po ieee merbivett 96 235 ox) (OLei meri urascens, (Ny!.) Gyeln. (14471y) respectively, among other lichens. Psoroma ctnnamomeum Malme 1925 P. 11, 12 The specimen studied in comparison to the ho- lotype (Henssen 24448t) was growing on the same type of substrate. The prominent apothecial mar- gin is especially distinct in younger apothecia (Fig. 1G, 6). The cortex was well developed in all'sections studied. In the upper cortex the cell lumina are embedded in the gelatinous matrix of the cell walls. The lumina are roundish and @onnected by pits (Fig. 5A). In young thalli the cortex is formed only by periclinally arranged hyphae, later on cells are added by vertically growing medullary hyphae. In the lower thallus cortex the cell walls are less strongly gela- man Zed. tThe, cortex in the margo thallinus is formed by radial or somewhat reticulately arran- ged hyphae with thick walls and relatively small Poin amiie LC .aeoB, )D) The cephalodia are dark purplish brown and form clusters between the squamules or they are produced in groups below the apothecia. They are surrounded by a brown pigmented cortex, composed PeeOnewOL,sinupabt, two)rows Of cells (Fig. 5F). 448 In young stages the hyphae are thin and form a network and later the enlarged cells become ag- gregated into a pseudoparenchyma. ACKNOWLEDGEMENTS These studies were supported by a grant of the Deutsche Forschungsgemeinschaft, which also provided a travel grant for the field studies in South America. In Tierra del Fuego we were guests at the Marine Biological Station in Ushuaia, and the excursions were guided by Dr. S. Guarrera from the University of La Plata. We are greatly indebted to him and to Dr. Kthnemann for the profitable stay we had in this remote part of the world. We wish to thank Dr, C. Dodge, Burlington and the curators of the herbaria for the loan of the type material. Mrs. G. Traute is thanked for skilled technical assistance and Dr. Nash for reading and correct- ing the manuscript. ZUSAMMENFASSUNG Die neue Art Psoroma tenue Henssen, eine Flechte auf Schneebdéden und in Tundra-dahnlicher Vegetation gehort zur Psoroma hypnorum-Gruppe und ist am ndchsten mit P. cinna- momeum Malme verwandt. Morphologie und Anatomie der beiden Arten werden verglichen. Sie stimmen in der Wuchsform, der Struktur des Lagers sowie in der Gestalt und Anatomie der Cephalodien weitgehend tiberein. Zwei Varietdten der neuen Art P. tenue werden unterschieden. P. tenue var. tenue ist auf die stdliche Hemisphdre beschrdankt und wurde bisher in Tierra del Fuego und auf den South Shetland Islands ge- sammelt. P. tenue var. borealis ist anscheinend eine weit verbreitete Flechte der nérdlichen Hemisphare; Fundorte werden ftir die Alpen, Skandinavien, Island und Nordamerika angegeben. Beide Varietdten von P. tenue enthalten den neuen Flechtenstoff£f Porphyrilsaduremethylester und Pannarin- sadure, Sowie in einigen der untersuchten Proben zusdtzlich eine unbekannte Substanz (U,). Das Vorkommen dieser Stoffe in anderen Arten der P. hypnorum-Gruppe wird erwahnt. Psoroma bryantii Dodge und P. follmannii Dodge werden als Synonyme Zu P. hypnorum (Vahl) S. F. Gray gestellt. 449 RE RE RENGES Culberson, C. F. 1972. Improved conditions and new data for the identification of lichen products by a standardized thinglayerschromatographic method. J. «Chromatogr. 72: ie 7 Henssen, A. 1969. Die Entstehung des Thallusrandes bei den Pannariaceen (Lichenes) mit einer generellen Diskussion uber die Entwicklung lecanoriner und biatoriner Flech- tenapothecien. Ber. Dtsch. Bot. Ges. 82: 235-248, Henssen, A. et al. 1981. The Lecanoralean Centrum. In Ascomycete Systematics The Luttrellian Concept (D. R. Reynolds ed.): 138-234. New York, Heidelberg and Ber- lin: Springer-Verlag. James, P. W. and HensSen, A. 1976. The morphological and taxonomic significance of cephalodia. In Lichenology: Progress and Problems (D. H. Brown, D. L. Hawksworth and R. H. Bailey, eds): 27-77. London, New York and San Francisco: Academic Press. Jgrgensen, P. M. 1978. The lichen family Pannaricaceae in Europe. Opera Botanica 45: 1-124. NFR: Stockholm. Keuck, G. 1977. Ontogenetisch-systematische Studie tber Eridoderma im Vergleich mit anderen cyanophilen Flech- tengattungen. J. Cramer, Bibliotheca Lichenologica WO Lem own Vaduz. Malme, G.O. 1925. Die Pannariazeen des Regnellschen Herbars. AG DOC a Z0AENOgs 2 .l—2 3. Renner, B. 1980. Untersuchungen zum Einflu8 der symbio- tischen Alge auf den Stoffwechsel und die Struktur des Flechtenlagers. Dissertation, Philipps-Universitat, Marburg (microfiche). Renner, B., Henssen, A. and Gerstner, E. 1981. Pannarsaure und Porphyrilsduremethylester - Sekundarstoffe der Flechtengattung Psoroma. Z. Naturforschung (in press). MYCOTAXON Vol. XIII, Nox. 3, pps 450-456 October-December 1981 AN UNDESCRIBED PLEOMORPHIC SPECIES OF CODINAEA Norihiro TOYAZAKI Public Health Research Institute of Kobe City, Minato- jima-naka-machi 4-chome, Chuo-ku, Kobe 650, Japan AND Shun-ichi UDAGAWA National Institute of Hygienic Sciences, Kamiyoga l-chome, Setagaya-ku, Tokyo 158, Japan In the course of an investigation of airborne fungi in Kobe, Japan, a hyphomycete was obtained in pure culture which proved to represent a previously undescribed species of Codinaea Maire. When grown on a variety of media such as cornmeal agar, malt extract agar, oatmeal agar, potato- carrot agar and potato-dextrose agar, sporulation is pleo- morphic: lt produces (a) polyphialidicysdarker wa thickoas walled, and regularly septate conidiophores and allantoid, hyaline, 3-septate, and terminally setulate conidia charac- teristic of Codinaea, and (b) monophialidic, paler, and less septate conidiophores with fusiform non-septate con- idia somewhat suggestive of Chloridium. Due to the latter features, the isolate superficially resembles Chloridium codinaeoides Pirozynski (1972), which produces monophiali- dic conidiogenous cells and narrowly fusiform-obclavate, aseptate conidia. A recent account of the morphology and taxonomy of the genus Codinaea was given by Hughes and Kendrick (1968), who recognized 13 species and four anamorphic forms of Chaeto- sphaeria (Sphaeriaceae) and described in detail numerous collections from New Zealand. Subsequently the following 13 species have been added to the genus: C. maharashtrensis Pirozynski § Patil (1970), C. novae-guineensis Matsushima (1971), C. eucalypti Sutton § Hodges (1975), C. septata Sutton §& Hodges (1975), C. apiculata Matsushima (1975), Cc. Tunata Matsushima (1975), =CosbritannicasM «= Byeoll ts eUuLoyOl. C. hughesii M. B. Ellis (1976), C. cylindrospora Morgan- Jones & Ingram (1976), C. unisetula Morgan-Jones §& Ingram 451 (1976), C. clavulata Holubov4-Jechova (1976), C. parkhalen- sis S. M. & S. S. Reddy (1977), and Codinaea anamorph of Striatosphaeria codinaeophora Samuels §& E. MUller (1978) (as '"S. codinaeaphora'). Dictyochaeta fuegiana Speg., the type species of the genus Dictyochaeta Spegazzini that was con- sidered as a possible earlier name for Codinaea (Hughes and Kendrick, 1968), has been re-described by Godeas et al. (1977). However, there are too few diagnostic features available on the type to properly define the type species of that genus. Since none of the fungi of this genus are pleomorphic, the fungus from Kobe has been described here as a new species. Codinaea dimorpha Toyazaki & Udagawa, sp.nov. (Figs. 1-6) Solonvacmineagaromucormmealy slentcecrescentes wm itciuts— simae, ex mycelio vegetativo submerso compositae; hyphae aeriae sparsae; conidia abundantia, aggregata, brunneo- grisea; reversum incoloratum. Coloniae in aliis agaris communibus celerius crescentes, planae vel rugosae, tenues, penitus auratae vel aurantiacae. Mycelium immersum vel semi-immersum, ex hyphis hya- linis vel subhyalinis, ramosis, septatis, 1-3 um dian, laevibus compositum. Boudacmsteri cSeansentes: Structurae conidiogenae macronematae, dimorphae: (a) polyphialidicae et (b) monophialidicae. (a) Conidiophora ex hyphis repentibus superficialibus orientia, singula vel 2-3 aggregata, simplicia, brunnea, cylindrica, 200-280 x 6- 8 um, laevia, incrassata, omnino diametro uniformia, apicem versus pallescentia, recta vel flexa vel plerumque genicu- lata, usque 6-8 septata, superne successive elongascentia. Cellulae conidiogenae elongatae, cylindricae, vulgo usque ad 40-60 um longae, polyphialidicae, raro discretae, deter- minatae, pallide brunneae vel subhyalinae, multis collari- bus conspicuis praeditae; collaria infundibuliformia, 3-4 x 1.5-2.5 um, subhyalina vel pallide brunnea, postremo effracta. Conidia semi-endogena, acropleurogena, hyalina, allantoidea, ellipsoidea vel oblonga, parum curva, 22-28 x 7-8 um, 3-septata, laevia, ad basim interdum hilo incon- spicue et utrinque setula singula, simplici, hyalina, 5-6 um longa praedita, in massa mucida aggregata. (b) Conidio- phora ex hyphis superficialibus singula oriunda, simplicia vel interdum irregulariter ramosa, 2-3 phialides ferentia, eyiindrica; 45-100 x 2.5-4 ym, hyalina vel obscure oliva- ceo-brunnea, prope basin septata, laevia. Cellulae coni- diogenae monophialidicae, integratae, determinatae, cylind- 452 ricae, superne gradatim angustatae, collari distincto 2-2.5 x 1-2 wm praedita. Conidia semi-endogena, acrogena, hya- lina, cylindrica vel fusiformia, 10-18 x 1-2 um, parum curva, aseptata vel raro uniseptata, utrinque plus minusve truncata, in massa mucida aggregata. Chlamydosporae terminales, obscure olivaceo-brunneae, ellipsoideae vel pyxritormes, slacves,inCrassal ae, Ul on 6-10 um, aseptatae vel raro uniseptatae. Holotypus: cultura NHL 2891 ex aere urbis Kobe in Japonia,in I2.vi: 1980 (maeNw Loyazaki, 1sOlata |e inecol tece tione fungorum "National Institute of Hygienic Sciences (NHL), Tokyo, Japan." Etymology: lat. dimorphus = having two forms, referr- ing to the dimorphic conidiogenesis. Colonies on cornmeal agar (CMA) growing slowly, reaching 2.8 cm in diam after two weeks at 23 C, very thin, vegetative mycelium largely submerged; white aerial hyphae sparsely developed on the agar surface; conidia-bearing structures abundantly produced in central colony areas, "brownish grey'' (Kornerup and Wanscher, 1978; pl. 6-E-2); reverse uncolored.) Coloniesson malt extract _ycascmcxtrace agar (MYA) growing rather restrictedly, consisting of a somewhat thick basal felt, wrinkled conspicuously, velvety to floccose, producing abundant aerial hyphae, "pastel yel- low'' (Kornerup and Wanscher, 1978; pl. 2-A-4); conidia slowly developed, not affecting the colony appearance with- im three weeks: sreyerse "Orange ma hormerupeancendisciic ia. 1978; pl. 6-B-7). Colonies on oatmeal agar (OA) or potato- dextrose agar (PDA) growing more rapidly, reaching 3.0-3.6 Cm) in diam arter stwo-weekssaty25 UC, plane chil, .« cecum) ec. — low" or "deep orange’! (Kornerup and Wanscher, 1978; pls. 4- A-8, 5-A-8). Mycelium immersed or semi-immersed, composed of hya- line to subhyaline, branched, septate, 1-3 um wide, smooth- walled hyphae which often swell to knots of pale brown to brown cells up to 5.5-10 um wide. Sterilessetgemiacking. Conidia-bearing structures macronematous, of two types; (a) polyphialidic, and (b) monophialidic. Type (a) commonly developed on CMA and at later stages also on MYA, OA and PDA. Conidiophores arising singly from superficial- ly repent hyphae or in groups of two or three from hyphal knots, simple, brown, cylindrical, 200-280 x 6-8 um, smooth and thick-walled, uniform in width throughout, paler to- wards the apex, straight or flexuous, often markedly genic- ulate in the upper region, up to 6-8-septate, successively proliferating with age at short distances. Conidiogenous 453 Fig. 1. Codinaea dimorpha (Ni Dez Soe A. Conidiophores and polyphialides. B. Conidia(3-septate) . C. Monophialides and conidia. D. Chlamydospores. (All measurements are in um.) 454 Figs. 2-6. Codinaea dimorpha (NHL 2891). 2. Conidiophores. 3. Three-septate conidium. 4. Mono- phialidic conidiogenous cell and 0-l-septate conidia. 5 and 6. Polyphialidic conidiogenous cells. (Bars in 3-6 = 10 um and in 2 = 100 um.) cells elongated, cylindrical, usually up to 40-60 um in length, -polyphialidic, rarely discrete, determinate, pale brown to subhyaline, each phialide in turn becoming exter- nally scarred with the persistent remains of the collaret- tes; collarettes funnel-shaped, 3-4 x 1.5-2.5 um, subhya- line to pale brown, aged ones often broken off. Conidia semi-endogenous, acropleurogenous, hyaline, allantoid, el- lipsoid or oblong, slightly curved, 22-28 x 7-8 um, 3-sep- ae 455 tate, smooth-walled, sometimes with a basal hilum, provided at each end with a single, simple, hyaline setula 5-6 um in length, aggregated in a slimy mass. Type (b) developed at an early stage on MYA, OA and PDA. Conidiophores arising singly from superficial mycelium, simple or sometimes ir- regularly branching into two or three stalked phialides, cylindrical, 45-100 x 2.5-4 um, hyaline to dark olive- brown, septate near the base, smooth-walled. Conidiogenous cells monophialidic, integrated, determinate, cylindrical, tapering gradually into a well-defined, terminal collarette measuring 2-2.5 x 1-2 um. Conidia semi-endogenous, acro- genous, hyaline, cylindrical to fusiform, 10-18 x 1-2 um, slightly curved, aseptate or rarely uniseptate, more or less truncate at both ends, aggregated in a slimy mass. Chlamydospores often developed on MYA, OA and PDA, terminal, dark olive-brown, ellipsoid or pyriform, smooth, thick-walled, 10-15 x 6-10 um, aseptate or rarely unisep- ate’. NG Gye NOmOLrOWtl sOCCULS. Specimen examined: an isolate from air, at Kano-cho, Ikuta-ku, Kobe-shi, Japan, June 12th, 1980, N. Toyazaki, NHL 2891, holotype. Ofechesspecies, currently, assigned sto the genus, C- dimorpha somewhat resembles C. fertilis Hughes § Kendrick (1968) because of the crowded appearance of polyphialides which are produced at the portion where cylindrical coni- diophores proliferate, and because it forms prominent phialide collarettes, as well as fusiform, curved and setu- late conidia. It differs from C. fertilis in that it lacks sterile setae and has larger, 3-septate conidia and mono- phialidic conidiogenous cells. Chloridium codinaeoides (loc. cit.) may be confused with the monophialidic fruiting structures of the present Species, but in the former, conidia are smaller 5-9 x 1-1.5 um as opposed to 10-18 x 1-2 um in C. dimorpha. The another superficially similar hyphomycete, Hypho- discosia europaea Hol.-Jech. § Borowska has recently been described by Holubov4-Jechova and Borowska (1981). Both are characterized by cylindrical, setulate phragmospores produced acropleurogenously at the elongating part of sim- ple, erect, pigmented conidiophores. Hyphodiscosia euro- paea is distinct from the present fungus in that the coni- dia are sympodial-polyblastic and dark-colored at maturity, and in lacking a monophialidic conidia. ACKNOWLEDGMENT 456 We are grateful to Dr. Walter Gams, CBS, for his crit- ical review of the manuscript and for many helpful sugges- Cions. REFERENCES Ellis Ma Bl oy Ceeeeiore Demat vaccousmily DROMy Cele Samm Dr. 472-474. Commonwealth Mycological Institute, Kew, England. Gams, W., and V. Holubova-Jechova. 1976. Chloridium and some other dematiaceous Hyphomycetes growing on decay- TN WOO tame CUCTEBMY GO Lramm Ls mrp Godeas,/ A. M:.) S.G. Marchand @andsD. Gabraly 01977) lora criptogaémica de Tierra del Fuego. Tome X. Orden Hy- phomycetales. Fasciculo 1. Fundacién para la Educa- cién, la Ciencia y la Cultura, Buenos Aires, Argen- CinavemLloep. Holubova-Jechova, V., and A. Borowska. 1981. Hyphodisco- sia europaea, a new species of lignicolous Hypho- MY.CECES es GESkauMy iO lems ome 2 0 5 lee Hughes, S. J., and W. B. Kendrick. 1968. New Zealand fungi 12. Menispora, Codinaea, Menisporopsis. N. Z. ULB O Ghat Oc meg 25-150, oe Kornerup, A., and J. H. Wanscher. 1978. Methuen Handbook ote Colours 5 rdeedsm nme Met huchemLOndClhwmmE > cme Matsushima, T. 1971. Microfungi of the Solomon Islands and Papua-New Guinea (Kobe). p. 14. Matsushima, T. 1975. Icones Microfungorum a Matsushima Uectorumm (Kobe) Meme ppresOec7e Morgan-Jones, G., and E. G. Ingram. 1976. Notes on Hypho- mycetes XV. Two new species of Codinaea. Mycotaxon 4: 504-509. Pirozynski, K. A. 1972. »Microfungi of Tanzania I. Miscel- laneous fungi on oil palm. II. New Hyphomycetes. CMI My,.cO Seb ap ae ZrO LU. PirozynsSkae Kell rand > 0 ba tds Lo (Ore SOmems etose Hyphomycetes of leaf litter in South India. Can. J. BO Cai Gime0 foe Reddy, S. M., and S. S. Reddy. 1977(1978). A new species of Codinaea. Sydowia 30: 186-188. Samuels, G. J., and E. Muller. 1978. Life-history studies of Brazilian Ascomycetes 1. Two new genera of the Sphaeriaceae having, respectively, Sporoschisma-1like and Codinaea anamorphs. Sydowia 31: 126-136. Sutton, B. C., and C. S. Hodges. 1975. Eucalyptus micro- fungi: Codinaea and Zanclospora species from Brazil. Nova Hedwigia 26: 517-525. MYCOTAXON eee p Ole Di Nos 63) pp 45/7 =464 October-December 1981 fhemhsilopezvordatuneieeVvadle Additions to the genus Pachyella. Donald H. Pfister Farlow Reference Library and Herbarium of Cryptogamic Botany Harvard University, Cambridge, MA and Frangoise Candoussau 22, rue H6o-Paris, 64000 Pau, France SUMMARY A revtew of the genus Pachyella ts gtven tn vtew of recent collections from France and continued study by the sentor author. One new spectes ts proposed, P. peltata, and one new combtnatton tn made, P. pseudosuccosa. A key ts gtven to the nine spectes now tneluded in the genus. The senior author has revised the genus Pachyella (Pfister 1973) and emended its circumscription. The junior author has made significant new collections of species Of the genus in France. This paper presents one Mewespeclcsnotm acnyella trom trance, asreportsot.?. punettspora from France, and a revision of the nomencla- ture of one species. A revised key to the genus Pach- yella is given. The literary history of the genus Pachyella was re- viewed by =riister (1975). “Sance that Cimesseveral smaller contributions have been made (Pfister 1974, 1975). Pristens(1979) also added’ on species to thesgenus er. hydrophtlta (Peck ex Sacc.) Pfister, and Berthet and Dona- dini (1978) described Peztza aquatilis which was later transferred to Pachyella by Donadini (1978). This species is discussed below. A recent paper by Donadini (1980) should be men- tioned. In it Donadini reduced Pachyella to the rank of a subgenus under Pezgtzga. There is no indication that 458 specimens were studied. Unfortunately several bibliogra- phical improprieties have occurred. We feel that Pachyella should be maintained as a distinct genus. The species are not only morphologically and anatomically distinct from Pegtza but they are ecolo- gically quite different as was pointed out earlier by Pfister (1973). The following is a synopsis of the species now placed in Pachyella. Complete synonymies may be found for previously treated species in Pfister (1973). SVNODS smo les peeics Pachyella adnata (Berk. & Curt.) Pfist., Canadwnum pou. Sle 20 L0te o/s. Earlier Aleurta annamttica Pat. was added to the synonymy (Pfister 1974). Few additional collections have come to light. The distribution is wide but disjunct. Additional collections deposited in FH have been examined from Ohio (W. B. Cooke 48304) and Connecticut (DHP). The species is unreported in Europe. Donadini (1980) listed the authors as "(Berk. et Currey)" [sic]. Pachyella aquatilis (Berthet & Donadini) Donadini, see P. pseudosuccosa. Paehyella babitngtontt (Berk. & Br.) Boud., Hist. Class. DUSCOM ee LU lea Dime orl) ie Peztza extditformis Berk. & Br. may be added to the already lengthy synonymy. Donadini (1980) listed the authorseor PD. ipabirgton ease (Berke eh Bress)iue| sll. Pachyella elypeata (Schw.) Le Gale Prodr. Madagascar’ 4:)277 1953. This species is still known only from North America though the new species de : scribed below from F 1 quite simitar® a Noe Flore Mycol. Pachyella hydrophila (Peck ex Sacc.) Pfister, Mycotaxon 8335.) 19798 This species and Pachyella punetispora are quite Similar anatomically. Detailed field studies of P. hydrophtla should be made--it is thus far only known from dried material. 459 Pachyella megalosperma (Le Gal) Pfist., Canad. J. Bot. ime OL Ieee O73 No recent collections of this species have come to our attention. Comments on Pachyella pseudosuccosa should be seen below. Figure 1. Pachyella peltata. Apothecia from the type collection. Scale = 2 cm. Pachyella peltata Pfister §& F. Candoussau sp. nov. Figures 1 and 2 Apothecta 4-7 em diam, planum, sessilta planae, sim- tits Pachyella clypeata. Hymentwn brunneum. Exctpulum extertum ex hyphts 55 x 25 wm composttum. Hyphis exteris rectts vel undulatts vel ramtficans. Asct J+, 500 x 20- 22 um dtam. Ascosport elltpsoidet, laevt biguttulati 27-350 x 18-15 ym. Paraphyses rectae, brumnnae. Holotype: sur tronc de Saltx pourri, 4 terre, endroit inondé périodiquement, avec Pstlopezta nummtlartalts Pfist. §& Candoussau et Climacodon pulcherrtmus (Berk. §& Curt.) Nikol. ainsi que sur Alnus gluttnosa au méme endroit. 14. IX. 1980. Forét de Bugangue-Oloron, Pyré- nées Atlantiques FH. Isotypes CUP and herb. Candoussau. 460 SSS eon See Figure 2. Pachyella peltata. Outer excipulun. Scale = 50 pm. Apothecia gregarious 4 to 7 cm diam, sessile, flat, toward the center undulating--plicate as far as the margin. External surface dirty-white. Hymenium brown (chatain), shining, slightly wiscous hlesi se dirtyewa tesUupn ton camm thick at the center, tinted yellowish where broken. In external features agreeing closely with P. elypeata (Schw) theaGale Outer excipulum of longitudinally elongate cells with their long axes oriented perpendicularly to the outer Psurfacero ts theceapotiecium ao oece ll Smlecp acc larger of the *célissmeasuringe ome co impor es li ehie by Jarger..* Cells tichtly *packedeandvonl\es lish tlyecwollen in width. The outer cells are uniformly embedded in a gelatinous matrix and give rise to hyphoid hairs of 5 to 7 ym in width to more than 400 pm in length, which often branch and anastomose or become moderately coiled. Medullary excipulum of dense texturasintricata ce! sparingly present, hyphae 6-12 um in diam. Asci diffusely J+, nearly 500 pm long and 20-22 pm broad. : Ascospores at maturity hyaline, thin-walled, smooth or with some cytoplasmic granulation, biguttulate but often jobscurel yrsone 2/50 exe Solo anim Paraphyses septate, enlarged above, exceeding the asci, of granular contents which become dark brown in freshamatenivaluwheneplacedmins1odine. This species is quite similar to Pachyella clypeata; the spores are larger and the configuration of the hyphoid hairs differszaasinjhabital and erowth forme tars pdentical.; 461 Figure 3. Pachyeila punetrspora. Apothecia from material collected in France. Scale = 1 cn. Pachyveugapunertspora Prist, , Canad. Js Bot. ol: 2019. SR Figure 3 = Pezitza puncttspora (Pfister) Donadini. Doc. myco- logiques 11: 26. 1980. Pierre Gabard, Guy Roux and the junior author have collected this species in France (Pont du Goa, Vallée du poussucou, 900 m alt. 15.VII.1979; Crampes de Piet, Pallee d!'Ossau. s00 malt. Leg. Guy Roux, Pyr. Atlan- tiques, environs de Pau, s/Betula et autres bois, 28.VI1. 1979). These collections constitute the first report of the species outside North America. Figure 3 shows apothe- cia of the French collection. Pachyella pseudosuccosa (Le Gal) Pfister, comb. nov. Galaetinta pseudosuccosa Le Gal, Rev. Mycol. (Paris) 10: 91-95. 1945. Pegiza aquatilis Berthet & J.-C. Donadini. Bull. Mens. Soc. Linn. Lyon 47: 205. 1978. Pachyella aquitilts (Berthet & Donadini) Donadini, BuliSoce Linn. Provence 317.518251973, When Le Gal described Galactinta pseudosuccosa, she did so in such a way as to introduce some ambiguities, Dee) ll 462 particularly in clearly stating the name in association with the Latin description. It seems to be validly published. One form she described, forma macrospora, was the form upon which Galactinia megalosperma (Le Gal 1953) was based. The typical form, material of which has been unavailable for study, was described from dried material in the Boudier herbarium and was collected by Barla in NIC Cw iim ode A portion of the holotype specimen of Pegtza aquatt- its, kindly sent by M. Berthet wapnees Closclyswatiimtnat described by Le Gal as Galactinta pseudosuccosa. Both have a reddish brown hymenium and both are said to produce a yellow juice when broken. Moreover, the Barla collection and those of Donadini are all from the same general area in S. W. France: Pachyella vtolaceontgra (Rehm) Pfist., Canad. J. Bot. aig aval absyats Previously the combination of this species in Galac- LENCGaDY sBOUdL Cy (HUStyeG1asS mp Us COM mc Liem) m4 eel OU @) was overlooked. Also to be added to the synonymy is the combination: Pegziza vtolaceontgra (Rehm) Smits'Ka (Ret- SLUSOVIeGri Di Ukrayiniod KLCV) ep en oeLo >) mm onadi nd (1980) also proposed the combination Peztza vitolaceontgra. The junior author has made collections of Pachyella vtolaceontgra in France (Bois de St. Maur-Mirande, Gers, 13.VIII.1971; Forét de Bugangue, Oloron, Pyr. Atlantiques Do ee A975 HH HOLD ee Reeeb . ekhOF Geen el mt Ga NGOUSS ALI. Other collections were made in Switzerland by A. Nyffe- negger (near Kanton Bern: FHjZ)" and by iBruno Erb and Stappung Erlinsback (automne 1980, Herb. F. Candoussau). KevVeCOms DCClCSmO LE UaiiC11 a Hyphoid hairs forming a aca pean at the base of Giicma DOC Ne Cl aa sun ue A cea a, ey Hyphoid hairs not Revandlays a Hsccultayo Palisade layer aimee NemDaAsceOlmenceapOthectam.: 2). oll 2. Ascospores smooth 13-16 x 18- 25 (28) “pam. “Aieerdacosia UMDeRecOMChestiut color,.outer layer of exci pulum of radially arranged sileinage célls, hyphoid hairs neither branched nor erat ae N. America. : ve: fyi heels Ribena Ty Aecospannes or r apothecial aonstaneenan not as ADOVC Wms. 5 Ascospores snensal 13- 15 x 27- 30 si, ayterd ice ihe dese HMDeymOuL CG slay eCTsOLmexcapulumeot strontly packeds only slightly swollen cells, hyphoid hairs branched Wiel AWB Sie Wi. ONS. 6 ap 66 0 0 6 Wee jeune: nS COSDOLeSaVanlOUs Ly sOrlamenucdm., |.) eunsn po the cn ta 4, Ornamentations taking the form of iene eiheve topped warts, N. America, West Indies, and Japan. spat ean eC OnOLa A’ rene nar anes Ss ikiise in ane aovatl of warts or I MECTACO MNO EM NY LEMS 4 «6 Se Ae Spores ornamented with anaatetostins fi hemartiem reddish brown, known aes from Madagascar . : rae RRC LRECAE Sneiwes ermented rene agit CEeReS Bo 4 (ho ee Te 6. Hymenium dark brick to sepia, when damaged not producing a yellow ‘aig Busopemr : fee Se apererord sk 6. abareneiwi aati ovat nen damaged producing a velloW julce, burope’. 9... « .« 4. pseudosuccosa Spores smooth or punctate, margin attached to the substrate, apothecia Soon NO te Siea vet mtnanmt ac in diam, worldwide i ce Ney her Meme. DOD neLOnTe Spores ornamented with Se eens, margin free, apo- Checram ccictal |yelarcernsthanwl acm. mae 8 Sem opores el 2—1 55x. 21-25, 71m, marked neice smell occ discernable warts, el up to 9 pm, N. ANEGGECa, sEUrOpe wm. . .P. punettspora Seem porese | S=10exe22— 26 um, marked with warts which can, scarcely be seen with the light microscope, satis Ups tOmlo pithinadial iN. eAMCTC acme cme : Pe Ae ee oy ee Bo. tac hydrophila 463 464 ACKNOWLEDGEMENTS We wish to thank Richard P. Korf for his comments on this manuscript, seeror siberchets Kind lyescitemaveria imo t Peziza aquatilts for our study. We are grateful to Emile Jarias and Guy Roux. tor photographs. slhe senior author's work was supported in part by National Science Foundation Grant sDEBPs02340135- DL PERATURES CITED Berthet, P. and J.-C. Donadinuss l97o8 Un nouveaurlezrca Dy ex, Oe AManSe "lest cop dguatT Tom nOV eS) amebU aie Mens. Soc. Linn. Lyon 47: 202-205. Donadiniy J0-C8 919/87 a eLesyenresrez? sae epcimodl Nt Amans (II). Les Pezizes de Haute-Provence et de Dauphine-Savoie. Bull. Soc. Linn. Provence 31: 9-36. 1980. Le genre Peztaza (Dill.) Linne per Saint Amans sous-genre Pachyella (Boudier) nov. comb. Doc. mycologiques. 11: 25-26. Bilstein el pie OLS ees DSMLODeZLOLGm LUN Minmmmn VemmiC genus Pachye Lite Canad Jam ROU Od ez COG cUeoe ~ 819742) Ibid Vl Alew-taraunane bi care synonym of Pachyella adnata. Mycologia 67: 181. 1975s o€anningnel ectronemicrographcmores ascospores of Pachyella (Discomycetes). Mycotaxon 3: 105-108. 1979. Type studies in the genus Peztza VI. Species described by C. H. Peck. Mycotaxon 8: 333-338. MY COTA XON a Ee VOLS XIII," No. 3, pp. 465-468 October-December 1981 en ee ee ee pr ee Ra ee PITHOMYCES PAVGII, A NEW COMBINATION FOR TRICHOCLADIUM PAVGII AND PITHOMYCES FUNICULOSA MARY E. PALM and ELWIN L. STEWART Department of Plant Pathology Universtty of Minnesota SE Louse MN o0lUS USA Trtchocladtum pavgtt Nath was published as a new spe- cies from India on 16 July 1979 (1). Additionally, a new dematiaceous hyphomycete from Swaziland was described as Pithomyces funiculosa Palm, Stewart, & Rossman on 19 Octo- ber 1979 (2). We have determined that 7. pavgti and P. funteulosa are conspecific. Trichoeladtum pavgit has pri- ority according to the International Code of Botanical Nomenclature, Article 11.3 (3). However, based on conidium development and secession, 7. pavgtt can only be accommo- dated in Pithomyces Berk. & Br. Trichocladium Harz has been broadly circumscribed and at present contains a hetero- geneous assemblage of species which produce transversely septate conidia holoblastically. Pithomyces, in contrast, is characterized by the holoblastic production of dema- tiaceous conidia which are usually both transversely and obliquely septate. Moreover, rhexolytic conidium seces- sion leaves a characteristic denticle on the conidiogenous cell. Trichoeladium pavgit must be transferred to Pithomyces. The new combination is herein made and the taxon redescribed based on additional studies of isolates from India and the United States. Pithomyces pavgit (Nath) Palm, Stewart, & Rossman, comb. nov. Figs. 1-4 = Trichoeladium pavgit Nath, Sydowia 31: 122-125. 1978. (basionym) = Ptthomyces funiculosa Palm, Stewart, & Rossman, Mycotaxon 10: 246-250. 1979. COLONIES on malt agar white to grey becoming brown to black, floccose to appressed, dark ropy strands of conidio- 466 phores frequently present. HYPHAE septate, branched, of two types, hyaline to subhyaline, smooth, 1.5-5 (x=2.8) um diam, and subhyaline to pale brown, sometimes finely roughened, 2.5-6 (x=3.4) um diam. CONIDIOPHORES micronematous to semi- macronematous, mononematous, frequently funiculose, infre- quently branched, septate, subhyaline to pale brown. CONIDIOGENOUS CELLS holoblastic, monoblastic or polyblastic, integrated, intercalary, determinate or seemingly prolifer- ating percurrently to produce one or more conidia, cylin- dric to doliform, denticulate following secession of conidia, denticles short cylindric to doliform, 1-4 (x=2.4) xX 1.5-4 (x=2.3) um. CONIDIA solitary, pleurogenous, mostly verrucose, pale to dark brown, clavate to obovate to obpyriform to broadly or fusiform ellipsoid to reniform, transversely (0-)1-3(-4)-septate, mostly 2-septate, may be slightly constricted at the septa, rarely with 1-2 ob- lique or longitudinal septa, 10-17 (x=13.3) x 4.5-12 (x=7.4) um (1l-septate), 13-20 (x=17.2) x 5-12 (x=7.8) um (2-septate), 17-26 (x=22.1) x 6-12 (x=8.3) um (3-septate) , bases 2-4 (x=2.6) um diam, with a portion of the conidio- genous cell usually remaining as a basal frill. Collections examined: India: Hyderabad, Rajendranagar, V. Ravindra Nath. From kernels of Arachts hypogaea L. 3 i 1973 (IMI 205718 [holotype] and living culture received from V.R.N.). Swaziland: Luyengo, A.Y. Rossman. From . fungal comb of Macrotermes ukuzit Fuller. 15 1x 1977 (MPPD). United States: Alabama, Chambers Co., G. Morgan- Jones. From municipal industrial waste-water. v 1980 (AUAM 2405). In addition to its occurrence in India and Swaziland, G. Morgan-Jones (personal communication) has isolated P. pavgtt in the United States. Based on our observations using phase contrast and scanning electron microscopy, the conidiogenous cell either proliferates percurrently through the denticle, which remains as a short annellation, or the apparently inelastic outer wall ruptures and remains at the base of the exposed, elastic inner wall. Remnants of conidiogenous cells surrounding the possibly proliferating conidiogenous cell are seen infrequently and mainly in Older cultures. Ravindra Nath's line drawings of P. pavgtt indicate flared structures resembling annellations at the apex of some elongate conidiogenous cells which he de- scribed as "roughened". Examination of the type material (IMI 205718) and of a living culture indicate that the 467 FIGS. 1-4. Ptthomyces pavgit. 1. Photomicrograph (phase contrast microscopy for Figs. | and 2) of a developing co- nidium with intact outer wall. X2000. 2. Photomicrograph of a conidiogenous cell following rhexolytic conidium se- cession showing remnants of either the inner and outer walls or the walls from two successively produced conidia. X2000. Figs. 3-4. Scanning electron micrographs of conidium and conidiogenous cell with inelastic outer wall beginning to rupture (Fig. 3) and conidium and conidiogenous cell with ruptured outer wall or a possible annellation (Rid. 4s X3000. 468 fungus sometimes appears to proliferate percurrently, but, a conidiogenous cell so long and with more than one "annellation" was never observed by us. Final elucidation of conidiogenesis requires ultrastructural or time lapse Studies or both. ACKNOWLEDGEMENTS This work was made possible in part by funds provided by the USDA, Forest Service, Forest Products Division (Nos. 12-39 and 12-144). We thank Dr. G. Morgan-Jones for the opportunity to observe and cite his isolate of P. pavgtt and appreciate the helpful comments and suggestions of Drs. G. Morgan-Jones and W.J. Sundberg regarding the manuscript. Minnesota Agricultural Experiment Station, Scientific Journal Series, Paper No. 11,777. LITERATURE CITED 1. Nath, V. Ravindra. 1978. On Trtchoeladium and a new species, Trtchocladtum pavgtt. Sydowia 31: 122-125. ac) let Piy Wikteon pbs Sacaicteien cule ais MOSSE I Pithomyces funiculosa sp. nov. from fungal comb of Macrotermes ukugtt in Swaziland. Mycotaxon 10: 246-250. Sous Catlett Ases Ct a t0eds 1) melo / Cement erat Orde Code of Botanical Nomenclature adopted by the Twelfth International Botanical Congress, Leningrad, July 1975. Regnum vegetabile 97: 10. MYCOTAXON VOU HEL NOS spp.) 469-4:7.0 October-December 1981 VALIDATION OF THE HERPOMYCETINEAE AND HERPOMYCETACEAE IN THE LABOULBENIALES ISABELLE |. TAVARES Department of Botany, University of California, Berkeley, California 94720 When the Euceratomycetaceae in the suborder Laboulbeniineae was described (Tavares, 1980), a comparison was made between the development of the perithecia in the Laboulbeniineae and the Herpomycetineae I. Tav. (nom. nud.). Inadvertently, I omitted a reference to the second volume of Thaxter's monograph (1908) of the Laboulbeniales, in which the tribe Herpomyceteae was described in the key to genera within the family Laboulbeniaceae. In addition, there was no mention of the family Herpomycetaceae, nor was a diagnosis of the Herpomycetineae included (Tavares, 1980). Consequently, a Latin diagnosis for the Herpomycetineae follows, for which I am indebted to Dr. William J. Dress, Bailey Hortorium, Cornell University. Subordo Herpomycetineae nom. nov. Dioeciae. Thallorum femineorum perithecia uno vel pluribus axibus receptacular- ibus secondariis, qui una vel pluribus cellulis suprabasal- POUS@ileaxeupLimardiouproducti sunt, porntata. —Peritheci7 paries exterior ex seriebus quattuor verticalibus cellular- um constans, serie quaque ex cellulis numerosis brevibus altitudine subaequis composita. Cellula infima in una serie verticali cellularum parietalium perithecii juvenalis et introrsum et sursum per medium perithecii crescens, carpogonium formans. Antheridia sunt phialides. Familia Herpomycetaceae (Thaxter) stat. nov. Tribus Herpomyceteae Thaxter. Mem. Amer. Acad. Weta iesig TE ABW AER They Family Herpomycetaceae: As in the suborder. Dioecious, occurring on cockroaches (Blattaria). The ger- minating spore becomes transformed into a primary axis of four or more cells. In the female, perithecia are produced 470 on secondary axes arising from the primary axis. Vertical septa are formed in the perithecial primordium, producing four cells, each of which divides into a vertical row of wall cells. When there are two tiers of wall cells, one of the four lower cells grows inward and upward and forms a central carpogonial upgrowth (Tavares, 1980). At maturity, there are many short outer wall cells, equal or subequal in height, in each vertical row. Four vertical rows of inner wall cells surround the passage in the perithecial neck through which the ascospores emerge (Tavares, 1966). Eight spores having median septa are formed in each ascus. Male thalli bear elongate, simple antheridia. Thaxter's description (1908) of Herpomyceteae, which was only a brief statement in his generic key, neverthe- less served satisfactorily to distinguish the tribe, be- cause Herpomyces, the solitary genus, is the only dioecious genus having secondary receptacles and simple antheridia. Although Dimeromyces aberrans Thaxter (1924) may have simple antheridia, additional receptacular cells in the female thallus are produced in the primary axis and do not form a secondary axis as they do in the closely related genus Dimorphomyces, in which all antheridia are compound. In his poorly preserved material, Thaxter was unable to see the internal structure of the perithecium of Herpomyces (see Thaxter, 1931). Consequently, he (Thaxter, 1908) placed Herpomyces in the Laboulbeniaceae because of the simple antheridia, not realizing how strongly its peri- thecial development differed from that of the other genera invthis family. LITERATURE CITED Tavares, Isabelle I. 1966. Structure and development of Herpomyces stylopygae (Laboulbeniales). Amer. J. Bot. D3) BLI=3 18. - 1980. Notes on perithecial develop- ment in the Euceratomycetaceae fam. nov. (Laboulben- iales, Laboulbeniineae) and Herpomyces (Herpomycet- ineae). Mycotaxon 11: 485-492. Thaxter, R. 1908. ) Contribution toward, a monograph of the Laboulbeniaceae. Part II. Mem. Amer. Acad. Arts Sci. 133 217-4693. Pls. Px LP xXx, - 1924, Contribution towards a monograph of the Laboulbeniaceae, Part III. Mem. Amer. Acad. Arts Soin Pi 2309-406 2 ais elle « L931. “thrid Ss Part Ve" Mem. ‘Amer. Acad. vArts SCi ty elGst 1 —43 570 ee bs el bo ee MY COTAXON VON ELS eNO. 3, pp. 4/1=480 October-December 1981 ERYNIA (ZYGOMYCETES: ENTOMOPHTHORALES ) : VALIDATIONS AND NEW SPECIES RICHARD A. HUMBER USDA-SEA-AR Insect Pathology Research Untt Boyce Thompson Institute at Cornell HOUe TRO MECHA CUMmmuNeCW VOR K EL GO, OSA SUMMARY New specific names are proposed to validate three heretofore invalidly published species of Hrynta Nowakowski (Entomophthoraceae): £&. bullata Thax- ter & MacLeod in Humber, #. creatonott Yen in Hum- ber, and £. myrmecophaga Turian & Wuest in Humber. Erynta formtcae Humber & Batazy in Humber, sp. nov., iS proposed as a name for a fungus occurring on ants and differing from the incompletely char- acterized #. myrmecophaga. During recent studies on the entomopathogenic Entomoph- thorales, Hrynta Nowakowski and the closely allied or syno- nymous genus (depending on one's taxonomic opinion) Zooph- thora Batko have received much attention (Remaudiére and Hennebert 1980; Remaudiére and Keller 1980; Humber 1981, 1982; Humber and Ben-Ze'ev 1981; Ben-Ze'ev and Kenneth 1981, 1982). Several species referable to Hrynta sensu Humber & Be-Ze'ev (1981) are not validly published under the Inter- national Code of Botanical Nomenclature. This paper vali- 472 dates three of these species and proposes a second new spe- cies of Erynta from ants. ERYNIA BULLATA Thaxter & MacLeod, sp. nov. CORPORA HYPHALIA vartabtlts per tntttum contagtonts, hyphotdea postrem. NUCLEI persaepe consptcut, mtntmum 6-10 um tn dtametro, contentts gra- nulatts autem stne nucleolo consptcuo ovotdeo centralt. CONIDIOPHORA dichotome digttate ramosa ad apices, intertexta tn hymentum continuum; tn taentts emergens et itn integumenta super corpore hospttts coalescens. CONIDIA PRIMARIA untnucleata, bttuntcata, elltpsotdea ovotdea, 23.0-87.5 wn x2 12.0-16.5 wm (medto 29.5 x 14.0 um), expulsa vehementer everstone paptllae distinetae contra contdtophora. CONIDIA SECUNDARIA tnobser- vata. SPORAE PERDURANTES hyalinae vel stramineae vel palltdae alutaceae tn massa tn corpore hospttis; multtnucleata ubt maturae; eptsporits con- spteue vel bullatts vel tuberculatis, 37-62 \wm (medto 50 um) tn dtametro toto bullts tnelusts. RHIZOIDEA numerosa, stmplicta, hyphotdea, stne haptero disctform terminalt, ex abdomine et thorace emergentta; in praesentta vel conidtorum vel sporarum perdurantium formata. CYSTIDIA desunt. HYPHAL BODIES variable in size and shape during early stages of develop- ment in host, becoming hypha-like at time of the host's death. NUCLEI easily distinguished in most Vegetative or reproductive structures; large, usually 6-10 um or more in diameter; contents granular, without a prominent central ovoid nucleolus. CONIDIOPHORES branched digitately toward the apices, intertwining to form a continuous hymenium; emerging in bands from weak points of the host exoskeleton, coalescing to form more or less complete coverings over the abdomen and the junction of the neck and thorax. PRIMARY CONIDIA uninucleate, bitunicate, ellips- oid to ovoid, 23.0-37.5 um x 12.0-16.5 um (averaging 29.5 x 14.0 um), forcibly ejected by the eversion of the conspicuous basal papilla against the conidiophore. SECONDARY CONIDIA not observed. RESTING SPORES forming a compact, hyaline to creamy-yellow or light tan mass inside the host body; multinucleate when mature; individual spores with prominent, rounded bullations up to 5 Lm in height on the epi- spore; 37-62 um (averaging 50 um) in overall diameter (including the episporial bullations). RHIZOIDS numerous, unbranched, hypha-like, without any terminal disc-like holdfast, emerging principally from the abdomen and thorax; present whether the fungus produces conidia or resting spores. HOLOTYPE: CUP 58970 (Cornell Univ., Plant Pathology Herbarium). Resting spores in Sarcophaga aldricht. Coll.: D. Tyrrell, 235 June 1981. PARATYPE: CUP 58971 (Cornell Univ., Plant Pathology Herbarium). Conidia on Sarcophaga aldricht. Coll.: D. Tyrrell, 23 June 1981. TYPE HOST: Sarecophaga aldricht Parker (Diptera: Sarcophagidae). TYPE LOCALITY: Birch Creek Road, Webwood, Ontario, Canada. 473 This species appears to be a relatively common patho- gen of calliphorid and sarcophagid flies in North America. It was collected several times in the northeastern United States by Roland Thaxter (Riddle 1906, as Entomophthora "x") but never formally described. After Thaxter's death, Povah (1935) received permission from Dr. W. Weston at Harvard University to use Thaxter's unpublished herbarium name, Entomophthora bullata, for the fungus Povah found attacking bluebottle flies in Michigan. This name, attributed to Thaxter, was presented with scant details of the fungus's morphology except for the bullate nature of its resting spores (33-50 um diam. in Povah's collection) and the over- all similarity of the primary conidia of this fungus with those of Erynta amertcana (Thaxter) Remaudiére § Hennebert (1980). Unfortunately, Entomophthora bullata Thaxter in Povah appeared in print after 1 January 1935 and is, accor- ding to Article 36 of the International Code of Botanical Nomenclature, invalidly published without a Latin diagnosis or reference to any previously published Latin diagnosis. All subsequent combinations based on this name — JZooph- thora bullata (Thax. in Povah) Batko (1966) and Hrynta bul- lata (Thax. in Povah) Remaudiére § Hennebert (1980) — are also invalidly published. The most thorough study of £. bullata is that by Mac- Leod et al. (1973) who provide a complete morphological characterization. They suggest the host's age may affect WiectjerecOnralasor resting’ Spores. are produced at the cime of the fly's death. Kramer (1979) studied the time course Simin tectionsandstne spore states sprodcuccdsin whreesspectes Dimcagierpiottdmrt les mintcctedein tno =lavOraloLy se sicmCcOLLO- borated that the disease in younger flies does tend to pro- duce only conidia while the probability for resting spore production increases with the age of the flies at the time omeintecr one minecontrast to MacLeod et al (19/5), uwiic state that the fungus never produced both conidia and resting spores in the same cadavers of field-infected S. aldricht, Kramer (1979) found that, on rare occasions, both spore states may occur together in affected individuals of Phormia regina (Meig.) and Phaenicta sertcata (Meig.). It is significant, then, to note that in the field-collected examples of Sarcophaga aldrtcht in the holotype collection, tufts of conidiophores are apparent at the joints of the legs whereas the fly bodies are filled with resting spores. No bands of conidiophores are apparent on the abdomens of flies in the holotype collection. 474 Cytological studies by Riddle (1906) indicated that three-month-old resting spores of #. bullata are multinu- cleate. This condition was verified with resting spores from the holotype collectionestained wictieeci1 tncrsaccto- orcein or bismarck brown. Humber (198la) discusses the implications of this multinucleate condition in entomoph- thoralean resting spores. ERYNIA CREATONOTI Yen, sp. nov. "CONIDIA pyrtformia, uninucleata, med, 32 x 25 wm, bast paptllata, muro tenut, cytoplasmatae granoso; NUCLEI med. 12 x 9 \1m; CONIDIOPHORI ramost coalescentes tegitmenta suffusco vel roseo super totum hospitis corpus. CORPORA HYPHALIA brevia, segmentis hypharum tnecomposttis, 8-28 ym latis 20-24 ym Longts. CYSTIDIA raro tnventuntur. SPORAE PERDURANTES azygo- sporae (zygosporae pauca tnventae), globosae; eptsporto laevt atque crasso; cytoplasma granosum vel vacuolatus; dtam. 10-28 um. Hospes substrato RHIZOIDEIS ftxus." (Yen 1962) TYPE: Latin and English descriptions and figure 1-4, J. Insect Pathol. 4, 88-94 (1962). TYPE HOST: Creatonotus gangis (L.) (Lepidoptera: Arctiidae) TYPE LOCALITY: Kun-Kuan, Taipei, Taiwan. This species, originally proposed with the orthographi- cally incorrect name Entomophthora creatonotus Yen (1962), was invalidly published since no type was designated. The combinations Zoophthora creatonott (Yen) Batko (1964) and Erynta ecreatonott (Yen) Remaudiére § Hennebert (1980) are also, thererorem invalid! ye pubiisned mas! 1emCchOlCesO mete description and figures of #. creatonott to typify this spe- cies is motivated by the lack of any mention by Yen (1962) of the permanent deposition of either any specimens or the Slides from which his photomicrographs were prepared. The English description of all morphological characters of this fungus is meticulously detailed even if the figures are of Tim cCedauts li oye The placement of this species in Erynta is fully justified by its uninucleate (presumably bitunicate) coni- dia borne on digitately brached conidiophores. The large nuclei are readily detected, even in unstained material (Yen 1962), and correspond to those of all other Frynta Species. The presence of fascicles of hypha-like rhizoids and of a few cystidia supports this generic assignment. 475 ERYNIA FORMICAE Humber & Bazazy, sp. nov. CORPORA HYPHALIA trregularta per intttum contagtonts, 35-155 um x 8-16(20) wm, et posterius mantfestius hyphotdea et saepe parciter sep- tata. NUCLEI persaepe conspteut, magnt, contentis granulatis autem sine nucleolo ovotdeo centralt. CONIDIOPHORA in taentis distinctis emergentia, dtchotome vel digitate ramosa ad apices. CONIDIA PRIMARIA unitnucleata, bitunicata, obovata vel pyriformia, 18-25 um x 10-16 um, expulsa vehementer everstone paptllarum contra conidiophora; paptlla basalt altquantum distincta. CONIDIA SECUNDARIA contdia primarta stmu- Lantta quanquam parvtora, 16-19 um « 10.5-13.5 um, expulsa vehementer tn modum contdiorum prtmartarum. RHIZOIDEA sparsa crassa, haptera disetformta terminalia, ex junetura promesothoracica in fasciculum laxum emergentta. CYSTIDIA et SPORAE PERDURANTES tgnotae. HYPHAL BODIES irregular in length, diameter, and branching pattern in- side host body during early stages of development 35-155 um x 8-16(20) um, later becoming infrequently septate and more obviously hyphal in appearance. NUCLEI conspicuous in most vegetative or reproductive Structures «sarge, With granular contents but no central nucleolus: CONIDIOPHORES emerging in distinct bands from weak points on the insect exoskeleton, dichotomously or digitately branched at the apices. PRI- MARY CONIDIA uninucleate, bitunicate, obovate to pyriform, 18-25 um x 10-16 um, forcibly discharged by eversion of the more or less distinct papilla against the conidiophore. SECONDARY CONIDIA like the primary but smaller, 16-19 um x 10.5-13.5 um, forcibly discharged as in the primary conidia. RHIZOIDS relatively few in number, stout, terminating in a discoidal holdfast; emerging from the ventral thorax just behind the head, forming a loose fascicle. CYSTIDIA may be present; if so, tapering to a point from a relatively broad base. RESTING SPORES not observed. HOLOTYPE: CUP 58975 (Batazy 1285a), coll: J. WiSniewski and A. SOKOLOWSKin m1 9eSept 91977, slucholaspines forest, Poland PARATYPES: CUP 58976 (Batazy 1286), CUP 58977 (Batazy 2074a) and specimens retained by S. Batazy and R. A. Humber. TYPE HOSTS: Ants of the genus Formtca (Hymenoptera: Formicidae): F. polyectena and F. rufa. TYPE LOCALITY: Pine forests near Tuchola, Poland. A comparison of entomophthoraceous fungi reported to attack ants in Switzerland (Turian and Wuest 1969, 1977), the Federal Republic of Germany (Loos-Frank and Zimmermann 1976), and Poland (Batazy and Sokoktowski 1977) indicates the existence of two distinct species of Frynta. FErynta formtcae is described for the better characterized fungus (Loos-Frank and Zimmermann 1976; Batazy and Sokotowski 1977); these collections have been attributed to the inval- 476 idly published and less completely characterized Zoophthora myrmecophaga Turian & Wuest (which is validated below as Erynta myrmecophaga Turian §& Wuest). The conidia of the Swiss fungus are smaller (averaging 18 x 12 um), considerably more ovoid, and have a less con- spocuous papilla than those of F. formcae. The conidia of the Swiss fungus but not of #. formcae have a single large vacuole occupying most of the conidial volume. In addition to being held onto grass blades by the mandibles and clasped legs, #. formeae further secures the ant cadavers with a few stout rhizoids emerging in a loose fascicle from the ventral thorax (Loos-Frank and Zimmermann 1976; Batazy and Sokotowski 1977). This sort of rhizoid generally terminates in a discoid haptor (Batko 1966; Humber 198la); although Loos-Frank and Zimmermann (1976) refer to a holdfast disc ("Haftscheibe') on the rhizoids, neither they nor Batazy and Sokotowski (1977) illustrate the morphology of this holdfast. Erynta myrymecophaga has no such fascicle of rhizoids; Turian and Wuest (1977) report the presence of a few very fine’ rhi- zoids emerging from all portions of the host body. The presence or absence of cystidia in £. formticae re- mains uncertain. Loos-Frank and Zimmermann (1976) note the presence Of isolated, broadicystidlantaperingmtordampoiet, but do not state where these occurred on the host, or at. what stage of development. Batazy and Sokotowski (1977) observed no cystidia on any specimens allowed to develop and sporulate in damp chambers. Loos-Frank and Zimmermann (1976) provide a detailed study of the effects of #. formtcae on its host, and find the behavior of affected ants and fungus-mediated histo- pathology of the brain to be much like the effects in ants caused by the parasitic trematode Dicrocoeltum dendrtttcun. Humber (1975, 1976) also found a marked tendency for hyphae of Stronguellsea magna’ Humber to penetrate the thoracic gan- glion and brain of affected flies, but without causing any obvious histopathology or changes in host behavior. More complete information about the morphology and pathobiology of EF. fornteae is provided by Loos-Frank and Zimmermann (1976) and by Batazy and Sokotowski (1977). 477 ERYNIA MYRMECOPHAGA Turian & Wuest, sp. nov. CONIDIOPHORA paretter dtchotome ramosa; tn taenits ctreum abdomen inter abdomen et ocetptttum et etreum partes orts emergens. CONIDIA PRIMARIA uninucleata, bitunicata, late ovotdea vel pyrtformta, medio 18 um x 12 um, vacuolam centraltam magnam conspicuam et guttulas oleosas sudano- phtlas numerosas continentia; paptlla basalt indistineta. CONIDIA SECUNDARIA contdia primaria stmulantta stne vacuola quanquam parviora. Hospes ad substratum mandibults et crurtbus adhaerens. CONIDIOPHORES sparingly dichotomously branched, emerging in bands on the abdomen, between the head and thorax, and around the mouth. PRIMARY CONIDIA uninucleate, bitunicate, braodly ovoid to pyriform, averaging 18 um x12 um, containing a large central vacuole occupying most of the conidial volume and numerous sudanophilic oil droplets; basal papilla indistinct. SECONDARY CONIDIA similar to but smaller than primary conidia, but not containing the prominent central vacuole. Host at- tached to the substrate by the mandibles and clasped legs. HOLOTYPE: Figs. 1, 2, and 4, Mtt. Schwetz. Entomol. Ges. 50:285- 289" 61977)% TYPE HOST: Servtformica fusca L. (Hymenoptera: Formicidae). TYPE LOCALITY: Sparse grasslands near Geneva, Switzerland. This description of £. myrmecophaga is obviously frag- mentary. There remains enough reasonable doubt about cer- tain aspects included in the Latin description provided for Zoophthora myrmecophaga (Turian and Wuest 1977) that it was preferable to provide a wholly new description including only those characters which seem relatively well estab- lished. Turian and Wuest (1969) found no rhizoids attaching ants affected by EF. myrmecophaga to the substrate. These authors later described very fine rhizoids (Turian and Wuest 1977; Fig. 4) which are little distinguished from the epidermal hairs on the grass blades to which the ants are affixed. This characterization may actually note tru rhi- zoids of E. myrmecophaga, but their appearance is not typi- cal of other entomophthoralean rhizoids. The possibility remains that these structures, observed on insects collected in 1967 and 1968, might have been either strands of insect or spider silk or the fine hyphae of some saprobic deutero- mycetous contaminant. 478 Similarly, it is so unlikely that the two different types of restingespores notedeby. lurianeandeWues te GlJ0 7. 1977) actually represent any over-wintering spores which might be produced by #. myrmecophaga that these spore types were intentionally omitted from the description and typifi- | cation of ‘this fungus vabove. “lurlanvandeWuest 41969 shic: 3) first reported finding one solitary resting spore among the hyphaly bodiesmin’ onevant;, = this se lobosessporcenased Single-layered, relatively thin, smooth-surfaced wall, was 20 um in diameter, and contained numerous oil droplets. If | this was an entomophthoralean resting spore, it was immature | and neither its size nor structure could be considered to be | taxonomically significant. Turian and Wuest (1977; Fig. 3) later found a. few globose resting spores with thick walls | and rugose surfaces; the illustrated structure does not have | the thick, two-layered wall which is characteristic of ento- mophthoralean resting spores. Neither were these rugose spores found in the numbers or aggregated mass in the abdo- men that one might expect for naturally produced entomoph- thoralean resting spores. The best clue to the identity of these structures comes from the conspicuous furrows on the surface; this morphology is unprecedented in the Entomoph- thorales but is strongly reminiscentoot tricolipate polien grains like those produced by oaks and beeches (Fagaceae) (Faegri and Iversen 1975; Bassett et al. 1978). Erynta myrmecophaga obviously requires further study from both any existing specimens and any future collections. Important. differences between this species: ands thestarepet— ter characterized #. formtcae exist in the size and shape of the primary conidia, and the presence and nature of rhi- zoids. It should be noted, however, that the behavior and appearance of affected ants, the; relatively low desree of conidiophore branching, absence of cystidia, and absence of verifiable resting spores are common characteristics of these species. s Noneroreche simmanrtulesaromsuttucten til valine portant or convincing to suggest that £. formcae and E. myrmecophaga are not distinct species. ACKNOWLEDGEMENTS I wish to express my deep gratitude ‘to R. P. Karf for invaluable discussions about the nomenclatural problems of validating these species, to D. Tyrrell and S. Batazy for 479 providing the type material of #. bullata and FE. formicae, POs mmm GessmLOreconrecting the Latin descriptions, andsto Heo epooper and DE. McCabe for their review and criticism Dretheemanuscript. LITERATURE CITED BALAZY, S., and A. SOKOLOWSKI. 1977. Morphology and biology of Entomophthora myrmecophaga. Trans. Britt. Mycol. Soc. Bo melo 4-157, DADOEIT, I:.J., C-W:. CROMPTON, and J.W. PARMALEE. 1978. An atlas of atrborne pollen grains and common fungus spores of Canada. Canad. Dept. Agric., Res. Branch (Ottawa), Monograph No. 18. BATKO, A. 1964. Some new combinations in the fungus family Entomophthoraceae (Phycomycetes). Bull. Polon. Acad. Sct. SCrmmcctUm btOlw ic: 405-406. BATKO, A. 1966. On the subgenera of the fungus genus Zoophthora Batko (Entomophthoraceae). Acta Mycol. 2: 15-21. : BEN-ZE'EV, I., and R.G. KENNETH. 1981. Zoophthora ortenta- lts sp. nov., a new fungal pathogen of Aphts cttricola (Homoptera: Aphididae), and two new combinations of other species of Entomophthoraceae. Phytoparastttca 9: 33-42. BEN-ZE'EV, I., and R.G. KENNETH. 1982. Features-criteria of taxonomic value in the Entomophthorales: I. A revision of the Batkoan classification. In preparation for Mycotaxon. FAEGRI, K., and J. IVERSEN. 1975. Textbook of pollen analy- sts, 3rd ed. Hafner Press, New York. HUMBER, R.A. 1975. Aspects of the biology of an insect- parasitic fungus, Strongwellsea magna (Zygomycetes: Ento- mophthorales). Ph.D. dissertation, University of Washing- ton, Seattle. HUMBER, R.A. 1976. The systematics of the genus Strong- wellsea (Zygomycetes: Entomophthorales). Mycologia 68: 1042-1060. HUMBER, R.A. 1981. An alternative view of certain taxonomic criteria used in the Entomophthorales (Zygomycetes). Mycotaxon 13: 191-240. HUMBER, R.A. 1982. Strongwellsea vs. Erynta: the case for a phylogenetic classification of the Entomophthorales (Zygo- mycetes). In preparation for Mycotaxon. HUMBER, R.A., and I. BEN-ZE'EV. 1981. Erynta (Zygomycetes: Entomophthorales): emendation, synonymy, and transfers. Mycotaxon 18: 506-516. 480 KRAMER, J.P. 1979. Interactions between blow flies (Calli- phoridae) and Entomophthora bullata (Phycomycetes: Ento- mophthorales). J. New York Entomol. Soe. 87; 135-140. LOOS-FRANK, B., and G. ZIMMERMANN. 1976. Uber eine dem Dicrocoeltum-Befall analoge Verhaltensanderung bei Amei- sen der Gattung Formica durch einen Pilz der Gattung Entomophthora. 2. Parasttenk. 49: 281-289. MacLEOD sD eMan ep Deh RRELL eR Oe erOUP ER manda tem CCil len he 1973. Entomophthora bullata as a pathogen of Sarcophaga alarichi. Jay TRVveErteDV mM IPaciGl i 2amtz0e x 5-7 HB 70-176x 28-56 Zygnemomyces F echinulatus N AH FH 2-5 same same pendulatus N AH FH 5-12 same same Ballocephala sphaerospora TD AH 5-80 x 5=10 HB 40-80 x 5-10 verrucospora TD AH ; ie HB 20-40 x 10-12 pedicellata TD AH ? 2 HB 20-31 x 13-16 All measurements in pm. D=desmid algae AH=aseptate hyphae ° Davidson and Barron (1973) F=fern gametophyte HB=hyphal body + Drechsler (1940) N=nematode HC=hyphal cluster * measured from illustration TB=tabanid larvae FH=filling host TD=tardigrade SH=septate hyphae hyphal bodies at maturity. In Ancylistes, Ballocephala and Macrobiotophthora vimariensis, after an initial mycelial stage, the hyphae become regularly septate. The newly formed-cells may remain attached, or fragment into freely floating hyphal bodies, depending upon the physical activity of the host. In Completoria, the infection bladder germinates and forms numerous short hyphal lobes which form a radiating cluster of cellsstiilVingsthe host cellsicompletely .alhiseis reminiscent of growth patterns in some of the entomogenous Entomophthorales (Thaxter 1888). 485 TABLE 3. PRIMARY SPORE CHARACTERISTICS OF THE NONENTOMOGENOUS ENTOMOPHTHORALES. PRIMARY SPORE SPECIES shape ornam length width base disch #spore devl Ancylistes closterii PYR - 13.6 5-17.25 P + 1 4h pfeifferi PYR - 21-23.5 P + ay netrii PYR - 16-25 P + 1 Be Completoria complens PYR = foe p == 1 T Macrobiotophthora vimariensis OBV - 18-21. 5x10-11.5* TR Fy 1 db vermicola OBV = 16—2 26xXG9= 4 TReee 1 T Castaner (1968) OV/B - 6.5-9.5x6.5-8.-5 TR + 1 ae Amin & Web. (1980) OBV - 21-26 x 12-16 TR + i dW Meristacrum asterospermumt OBV = S= 119 xso-8 TR oh 5-15 B asterospermum? OBV a I=17) - 5xO—10 TR ae 2-8 B milkoi PYR - 12-16 x10-13 P + 4(3-1) B Zygnemomyces echinulatus SPH ECH 527-72 U = 10-20 A pendulatus SPH ECH Da U = t=3 A Ballocephala sphaerospora SPH - 3-0-4.5 U + 20-80 A verrucospora SPH ECH 4.5-6.0 U ae 4-12 A pedicellata SPH - 6-7-8.0 U ate 2-10 A All measurements in pm. ECH=Echinulate A=Acropetal *Measured from published OBV=Obovoid B=Basipetal illustrations. OV/B=Ovate-Obovate P=Papillate ° Davidson and Barron (1973) PYR=Pyriform T=Terminal t Drechsler (1941) SPH=Spherical TR=Truncate U=Unmodified 3. PRIMARY SPORES. Morphology. Shape. Three basic spore shapes are found in the non- entomogenous Entomophthorales; all spores are of the uni- tunicate type (Remaudiére and Hennebert 1980, Remaudiere and Keller 1980). Ancylistes, Tabanomyces, and Completoria have pyriform spores (i.e. globose spores with conical Dapillace(berdaans.L939,.) Couch, 19497, Leitgeb s16o1)* ancmare similar to those found in Conidiobolus. Meristacrum has obovate spores with truncate bases (Davidson and Barron LO Eee Anos tinal. y,s callocephala (Drechsler 1951, Rich- ardson 1970, Pohlad and Bernard 1978), Zygnemomyces echin- ulatus (Miura 1973) and Meristacrum pendulatum (McCulloch 1977) have globose spores lacking basal modifications. 486 Ornamentation. Spore ornamentation is uncommon in the Entomophthorales. Some species of Massospora Peck emend. Soper have verrucose spores, and the primary spores of Con- idiobolus coronatus (Cost.) Batko may become villose by the growth of appendages through the spore wall. Spore orna- mentations consisting of small echinulations are also found in Ballocephala verrucospora, Zygnemomyces echinulatus, and Meristacrum pendulatum. Karyology. The morphological characterisitics of nuclei are con- sidered important by Humber (1981). The presence or ab- sence of nucleoli and heterochromatin (and hence the stain- ing ability of the nuclei) are taxonomically useful at the familial level (see §I1V.3), and certain genera with similar spores (e.g. Conidiobolus, Entomophaga Batko) can best be distinguished using nuclear morphology. The number of nuclei per spore has been used to reveal the misplacement of individual species (e.g. Culicicola Nieuwland [Humber V98 15) yes Unfortunately, few descriptions include karyological data, although there are a few exceptions among the fungi considered here. Hyphal bodies, gametangia and zygospores of Ancylistes closterili apparently have several nuclei with central nucleoli’ (Dangeard 1906). Couch (1949) was able’ to Stain nuclei (weakly?) of A. netrii with lactophenol-cotton blue and confirmed Dangeard's earlier reports. Moorman (1976) also reported nucleoli in her study of mitosis in an unidentified species of Ancylistes. From these limited reports). it appears,) that sAncylistes Pis# mostaisimielanrm vo Conidiobolus in terms of karyology (Humber pers. comm.) Two nonentomogenous fungi, Completoria complens (Humber pers. comm.) and Entomophthora vermicola, possess. the heterochromatic nuclei lacking central nucleoli typical of Entomophthora Fresenius. 4. PRIMARY SPOROPHORES. Spore Number. Most entomogenous species produce only one spore per Sporophore. Species of the entomogenous genus Erynia Nowa- kowski, while having apically branched sporophores, still produce only one terminal spore per branch. The nonentomo- genous members, however, can be divided into two groups. Single-spored sporophores are found in Ancylistes, Complet- Ooria, and Macrobiotophthora Reukauf. Pleuri- or multispor- ed sporophores are found in Meristacrum, Tabanomyces Couch et al., Zygnemomyces Miura and Ballocephala. The number of spores in the multispored genera is actually quite variable (see Table 3). Ballocephala has two to 80 spores per sporophore, with most of this range attributable to variation between individual sporophores of B. sphaerospora. Similarly, Meristacrum asterospermum usually produces five to 15 spores (Drechsler 1940) but was found to produce as few as two (Davidson and Barron 1973). 487 Sporophore Septation and Spore Development. Ballocephala produces sporophores with crosswalls at the base of lateral sporiferous branchlets. The main axis of the sporophore remains nonseptate, the spores are formed terminally, and then are pushed aside as the sporophore continues to elongate. Thus spores are formed and mature in acropetal succession. Acropetal spore formation is also Observed in Zygnemomyces echinulatus and Meristacrum pendulatum (Figs. 1-2). In Meristacrum asterospermum and Tabanomyces. milkoi the sporophores become transversly septate, delimiting sep- arate cells, each of which produces a single spore. Septum formation, and hence spore maturation, is basipetal. 5. PRIMARY SPORE DISCHARGE MECHANISM. Ingold (1934, 1971) distinguished three types of spore discharge in the Entomophthorales: papilla-eversion (Conid- iobolus), sporophore cannon (Entomophthora), and sporophore rocket (Basidiobolus). The method of spore discharge found in Ballocephala (see below) should be recognized as a fourth type, and fungi that have lost spore discharge could be placed in a fifth category. The basic, and putative ancestral method of spore dis- charge (papilla-eversion) involves the separation of the wall layers of a columella which projects into the spore. The domeshaped columella that remains (more or less intact) after spore discharge and the everted conical or rounded papilla at the spore base are diagnostic for this release mechanism. Ancylistes, Completoria, and Entomophthora vermicola (with simple sporophores), Tabanomyces milkol and Meristacrum asterospermum (with septate sporophores) have this type of spore discharge. The method of spore discharge was not seen in the "Conidiobolus-like fungus" (Castaner 1968) or Macrobiotophthora vimariensis, but can be reasonably inferred from their spore shape. All other spore discharge mechanisms are modifications of this basic method. The second type involves a sporophore having a more or less swollen apex, and which ruptures just below the spore. In Entomophthora s. str., the spore is campanulate, has a flat base, and is carried away in a stream of sap shot from the collapsing sporophore. It should be noted that in Entomophthora muscae (Cohn) Fresenius, the secondary spores are discharged by the papilla eversion method. The third discharge mechanism, found in Basidiobolus, involves the rupture of the base of a subspore swelling. This swelling tapers apically to a point forming a minute columella within the spore. The spore rockets away as the swelling expels fluid from its torn end, and lands with or without the remnants of the vesicle still attached to the everted spore base. Theseclaimeathate tht Semismeid si tWwO-stage rocket is probably more fanciful than accurate, as the decrease in turgor pressure within the sporophore must be accompanied by the simultaneous release of spore turgor by the eversion of the papilla. 488 TABLE 4. SPOROPHORE AND SECONDARY SPORE CHARACTERISTICS IN THE NONENTOMOGENOUS ENTOMOPHTHORALES- SPOROPHORE SUBSPORE CELL 2° SPORES SPECIES length width} septn length width replic disch Ancylistes closterii 3-725 - - - ++ + pfeifferi 4-7 = - - ae a netrii 25=100°xX95S - - - + + Completoria complens 60-80 ? - - - - ? Macrobiotophthora vimariensis 4-5 - - - + - vermicola 27-50 x 6-8 = - - + - Castaner (1968) <60 X16. 5=3 - - - +* + Amin & Web. (1980) <60-90 x ? - - - + + Meristacrum asterospermumt 200-500x6-9(3-5) +(S) - - + - asterospermum® 60-500x5-9(3-5) +(S,U)- - + - milkoi 90-200x7-8(4-6*) +(U) - - + - Zygnemomyces echinulatus 100-750x2.7-5.7 - 1-5-3 x «5-1-1 - - pendulatus 18= 71x2-3 a 1=3 = = Ballocephala sphaerospora 50-175x4-5 = 4-7 x 3-4.5 = ~ verrucospora 50-150x5-7 - ew oe Mr - - pedicellata 33— 9735 = 2-1 xX 1-6-2.6 = = (1-1-1-6) All measurements in pm. S=Spiralled sporophores * measured from published illustrations U=Unspiralled sporophore + Drechsler (1941) ++=produces 2°+ 3° spores ° Davidson and Barron (1973) **=produces microspores § (narrowest width) The sporophore discharge mechanism found in Ballo- cephala constitutes a fourth type. All species of Ballo- cephala have spherical spores lacking basal modifications, but are subtended by a special cell (Separated from the sporophore by a crosswall) whose rupture discharges the spore. Drechsler (1951) proposed that this cell was, in fact, the undischarged primary spore which produces a pre- mature secondary spore, a hypothesis which is consistent with the basic developmental patterns of the Entomophthor- ales. In B. sphaerospora and B. verrucospora, the spore bearing cells are utriculate, asymmetrical, curved upwards, and as large as the spore (Drechsler 1941, Richardson LOST MVD Vs Ballocephala pedicellata has ovoid to broadly Obpyriform pedicels which are much smaller than the spores. Spore discharge in this species is likely, although the evidence is circumstantial (Pohlad and Bernard 1978). 489 Spore discharge is lacking in Zygnemomyces echinulatus and Meristacrum pendulatum. In both species, the apex of the sporophore gives rise to a terminal spore lacking basal modifications, and then proliferates sympodially. The resulting pedicels are not delimited by a crosswall, although they may become devoid of cytoplasm. Thus, they are of distinctly different origin, than the sporiferous cells of Ballocephala. 6. SECONDARY" SPORES. If forcible spore discharge is a key characterisitic GfBcheseEnvcomophcthorales;—so too is the ability of thesdis— charged spores to produce ‘secondary' spores (Table 4). In almost every member of the order, primary spores that land On hostile substrates can produce replicative spores which aresetOorciblysdischarged. These spores are also called repetitive, Or secondary (2°), tertiary (3%), etc. spores. Exceptions to this rule are Ballocephala spp., Zygnemo- myces echinculatus, and Meristacrum pendulatum. Two additional types of ‘'secondary'" spores have been taxonomically useful. Microspores ("multiplicative spores" Drechsler 1952) are formed and discharged from single primary spores of various species of Conidiobolus. Basid- iobolus microsporus Benjamin produces elongate passively detached "microspores" by a totally unrelated method (Ben- jamin 1962, Tucker unpubl.). Microspores are also found in the "Conidiobolus-like fungus" (Castaner 1968) but are otherwise absent 1n nonentomogenous species. Capillispores ("capilloconidia" Remaudiére and Henne- bert 1980; "elongated secondary conidia" Drechsler 1955) are usually slightly thick walled, ovoid, ellipsoid (or Slightly irregular in shape), without modifications of the spore base and borne on long slender sporophores ("capil- lary conidiophores" Thaxter 1888). The loss of spore dis- charge is apparently counterbalanced by the increased in- fective potential or likelihood of dispersal by a passing host. True capillispores are produced only by Basidio- bolus. Nondischarged spores borne on tapering sporophores, are found in Meristacrum asterospermum, Macrobiotophthora vimariensis, and Entomophthora vermicola. 7. RESTING SPORES. Morphology and Development. Resting spores, whose function is one of perennation rather than dispersal, are usually thick walled and formed within the vegetative mycelium. The distinction between zygospores (developing following conjugation) and azygo- spores (developing without conjugation) has been questioned (Humber 1981), but is here considered provisionally useful (see below). Virtually all of the nonentomogenous Ento- mophthorales produce resting spores of one form or another (see Table 5). Ancylistes, as in Conidiobolus, forms zygospores after scalariform or lateral conjugation of unequal gametangia 490 TABLE 5. RESTING SPORE CHARACTERISTICS OF THE NONENTOMOGENOUS ENTOMOPHTHORALES ZYGOSPORANGIUM ZYGOSPORE / AZYGOSPORE SPECIES con form length width shape color orn diameter Ancylistes closterii U GB APPRESSED SPH BRN none 14. 5-20 pfeifferi U G 30-40 SPH BRN none 18. 5-22 netrii U cr. APPRESSED SPH fs none 19-26 Completoria complens E? GB APP RESSED SPH - none 1 Wis Macrobiotophthora vimariensis - - - - - - - vermicola E CE APPRESSED DDC - none 19-20 Castaner (1968) - - - - - - Amin & Web.(1980) - - - - = - - Meristacrum asterospermumt - HB LOOSE SPH YEL UND 20 asterospermum® - HB LOOSE SPH UND T= 20 milkoi E G APP RESSED OVD ~ none 12-20x23-30 Zygnemomyces echinulatus E HB/CT 22-28* OCT BRN none 16-24 pendulatus ? HB POL - none 18-24* Ballocephala sphaerospora E HB 28-33x14-16.5 DOL BRN none 17-20x13-16 verrucospora - - - - - - - - - pedicellata E HB APPRESSED DOL BRN PUNC 20.6-26.6 x 17-3-18-6 All measurements in pm. con=conjugation (color) *Measurements from pub=- E=equal gametangia BRN=Brown lished illustrations. U=unequal gametangia YEL=Yellow tDavidson & Barron (1973) dev=zygosporangia devel- “Drechsler (1940) ope from or within: (shape) CT=Conjugation tube DDC=Dodecahedral orn=ornamentation G=Gametangium OVD=Ovoid PUNC=punctate GB=Gametangial bud POL=Polygonal UND=Undulate HB=Hyphal body SPH=Spherical (PELtzer 81872))s. The presence of numerous protrusions on the zygosporangial wall is used to separate A. pfeifferi from other species of the genus (Berdan 1938, Couch 1949). Completoria forms = one. toy.tens. (or. 20 )peroundmtom oval thick walled resting spores within the enlarged cells in the central portion of the hyphal mass. (The peripheral cells of the thallus are by this time emptied by the forma- tion of primary sporophores or infective hyphae.) Conjuga- tion has not been observed (Atkinson 1884, 1885). zygnemomyces echinulatus forms brownish, thick walled, octangular zygospores following scalariform conjugation. The octangular shape results from the position of the zygo- Spore in the conjugation tube between the two parental hyphal bodies (Fig. 1E). The resting spores of Meristacrum 491 pendulatumsy are, similarsin=shape (Fig. 2F), but the mode of development is unknown. Entomophthora vermicola forms zygospores after scalariform or Lateral conjugation (Fig. 3K-P). At maturity, these zygospores are dodecahedral, each face with 5 (4-6) raised edges and a central depression (Fig. 3P). Conjugation in Ballocephala has not been proven, although its occurrence was inferred by Drechsler (1951) from the presence of two slightly inflated locules at either end of the zygospore. The zygospore itself is COMrormesOusecy Lindrical, sbrOwn atom dark—bDrown sine coLlor, smooth or punctate, and is preferentially formed at reduced temperatures (5° C) (Pohlad and Bernard 1978). Meristacrum asterospermum was described by Drechsler (1940) as forming azgyospores by the rounding up of the cytoplasm of a thalloid segment, laying down of retaining walls at either end, and deposition of a new undulate or scalloped wall around the cytoplasm. The star-shaped rest- ing spore is colorless or slightly yellowish, is about 15- 20 um in diameter, and contains a single large oil drop- let. Finally, in Tabanomyces milkoi, uninucleate, spherical gametangia fuse in pairs with the contents of one gametan- gium migrating into the other and forming a smooth, color- less ovoid zygospore with a single diploid nucleus (Couch et al. 1979, Nam and Dubitskii 1977). zygospores vs. Azygospores: A Different Perspective. Resting spores have been used for taxonomic purposes to some extent, but recent treatments of the family (Remaudiere and Keller 1980, Humber 1981) have cautioned against some traditional applications of resting-spore characteristics. In particular, the distinction between zygospores and azygospores (with respect to conjugation) was discussed at great length by Humber (1981), who. dismis- sed it for two reasons. First, the development of (a)zygo- Spores is often difficult to follow or interpret, and hence published accounts may be misleading. This) Ob Jections is valid, and the presence or absence of conjugation must be used with caution. However, the second objection, that the nuclear events of karyogamy and meiosis are of more import- ance to the fungus than whether conjugation occurs, poses an interesting question. Of what importance are the sexual events of conjuga- tion, karyogamy and meiosis to the entomophthoralean fungi? It must first be noted that all "sexual" Entomophthorales are homothallic (automictic) and homomictic (i.e. no mating types exist). Conjugation, when it occurs, jis between cells of the same branch or between hyphal bodies. Second- ly, since most spores and hyphal bodies are multinucleate, genetic recombination may occur somatically rather than during meiosis. There are very few species in the Ento- mophthorales for which evidence of meiosis exists (excep- tions include Conidiobolus thromboides Drechsler= Entomoph- thora virulenta Hall & Dunn, Tabanomyces milkoi). The lack Of data on basic aspects of sexual events and genetics is a 492 great, hindrancel=ins attempts. lO, assess u Cicer iilpOLcaicemson zygospores or to understand evolution within the order. Retaining the ability to coevolve with host organisms is jlikely; to be important sto =thesemtingl, but Ltesisrealso apparent that the presence or absence of conjugation or of karyogamy and meiosis do not seem to be as critical. Perennation and infective ability, however, continue to be important as a basic requirement of the entomophthoralean Ti Lemcy Cle. The formation of the resting spore, whether induced by conjugation or not, involves the phenotypic expression of extensive portions of the genome, and thus the shape, size, or ornamentation of resting spores may ultimately be of more taxonomic significance than nuclear SVCtl tors 8. SUMMARY OF TAXONOMIC CHARACTERS. From the preceding discussion of morphological charac- ters in the nonentomogenous Entomophthorales, it should be clear that character diversity within the order is greater than that encompassed by the entomogenous members alone. Nevertheless, the three characters which Humber (1981) pro- posed as most valuable in delimiting genera: "nuclear num- ber wand (nature, " “sporophore branching; =| andy =.methiodmor spore discharge," apply as well to the nonentomogenous genera, and the assessment of these characters needs only minor revision. 1. Spore karyology, i.e. nuclear morphology and num- ber per primary spore, remains important in distinguishing natural groups (i.e. families) within the Entomophthorales (see §I1V.3). This may be of practical value for separat- ing morphologically similar genera (e.g. Conidiobolus from Entomophaga; or Ancylistes from Completoria). 2. Spore discharge 1s virtually universal within the order and thus is presumably an ancestral character. The Spore discharge "theme" has five variations: papilla- eversion (Entomophthoraceae, Ancylistaceae), sporophore- cannon (Entomophthoraceae), Ballocephala-type (Ancylist- aceae), and sporophore-rocket (Basidliobolaceae), plus the loss of discharge (Entomophthoraceae [Massospora], Ancyl- istaceae [Zygnemomyces]). 3. Sporophore architecture now includes the presence or absence of sporiferous branchlets or transverse septa- tions as well as branching of the sporophore. In addition, the mode of spore formation (basipetal, acropetal, or term- inal) which is linked to sporophore septation, can be used to distinguish genera. It is presumed that the unispored sporophore is ancestral, and that multispored sporophores are derived. 4. Zygospore morphology, which Humber (1981) dis- counted but actually used as a secondary characteristic to delimit Triplosporium (Thaxter) Batko, is most appropriate- ly used for delimitation of species or subgenera. III. GENERIC TAXONOMY AND REVISIONS. 493 ANGYEISTEOMPriLezer, Monatsb. KoOnigl. Preuss. Akad. Wiss. Berlin. 396. 1872. myper) Species; Ancylistes closterii Pfitzer, Monatsb. Konigl. Preuss. Akad. Wiss. Berlin. 396. 1872. Other species: Ancylistes pfeifferi Beck, Verh. Zool.-Bot. Ges. Wien 46: PEvemieec9'o. Ancylistes netrii Couch, J. Elisha Mitchell Sci. Soc. 30: 251% 1949. Validation of this species is here accomplished with the publication of the following Latin diagnosis: Hyphae asumentes primum aseptatae 2.2-4.8 pm crassae, tum ramosae et septatae, cellulis 15-30 X 8-15 pm; hyphae exitibus unica in cellula una factae, 2-4 pm crassae, parietem hospitis juxta tumidae usque ad 4-9 pm emergentes et hyphas externas facientes, quae hospitem proxi- mum obnoxium inficunt, aut si ex aqua emergunt sporophora 25-100 ym faccunt; sporae primigenae 16-25 pm crassae papillis conicis; zygo- sporae intra hospitem inter aliquot hyphas longitudinales seriales formatae, ut videtur conjugatione cellulis duabus hyphis oppositis, zygosporis intra tubum conjugentem, sphaericus vel subsphaericis 19-26 pm crassis. Hospe: Netrium (Desmidiaceae). Ancylistes is in many ways, Similar to Conidiobolus, especially in spore shape and karyology (see §IV.2, IV.3). The species are divided on the basis of host (Saccoderm or placoderm desmid algae), size of the primary spore, and zygosporangial morphology. See Berdan) (1938) “and? Couch (1949) for a treatment of this genus. BALLOCEPHALA Drechsler, SOUL - BWeperathyy sendy (ONhisy RI Aer lee die Type species: Ballocephala sphaerospora Drechsler, Bull. Weyatente iweyen (agi sy 7Ase AU ile jonk: Other species: Ballocephala verrucospora Richardson, Trans. Brit. Mycol. SOCHIES 3 7mO0 Cian LO 7 Oe Ballocephala pedicellata Pohlad & Bernard, Mycologia 70: Tsuba Cals ey, Tivcmqenus;, =stound sonly sin Stardigrades ; miss unique min the manner in which the spores are formed and discharged. As previously mentioned (see §II.4, II.5), the discharged spore may. actually be a secondary spore developing neoten- ously from the undischarged primary spore, which is present in the form of a utriculate vesicle or a reduced pedicel. The species are separated on the basis of spore ornamenta- tion and the morphology of the sporiferous cell. 494 COMPLETORIA Lohde, Tagebl. Versamml. Deutsch. Naturf. Aertze 47: 206. 1874. Type species: Completoria complens Lohde, Tagebl. Versamml. Deutsch. Naturf£. Aertze 47: 206. 1874. This rare parasite of fern prothalli has been reported only twice since the original description (Leitgeb 1881, Atkinson 1884, 1885). The presence of large stainable nuclei lacking central nucleoli and a thallus composed of short hyphal branches indicate a distant relationship with most other nonentomogenous Entomophthorales. It is placed in the Entomophthoraceae (see §IV.3). MERISTACRUM Drechsler emend. Tucker & Humber, Us Wash. (ACaAd oC) 30m 25 eer O40. Type species: Meristacrum asterospermum Drechsler, J. Wash. Negeiely Sebo ys, PGbhs AO. Vegetative hyphae developing within living animals, Simple to somewhat branched, at first continuous, later dividing into disjointed segments which put forth a sporophore into the air, the terminal portion dividing basipetally, forming several to few cells, each of which produces a Single spore forcibly discharged by papilla- eversion; secondary spores replicative or passively detach- ed; resting spores azygospores formed from hyphal segments or zygospores formed from conjugation of spherical uni- nucleate gametangia. Other species: Meristacrum milkoi (Dudka & Koval) Humber, Mycotaxon 13: 213 239 Galt. Basionym: Coelomomyces milkoi Dudka & Koval apud Dudka, cOval & Andreeva, Novitates Systematicae Plantarum nonaVascularumelO<956 9-597 37. Synonym: Pseudocoelomomyces milkoi Nam and Dubitskii, Insts 2Z00l-mAlmawrAtayimAcadsii6cl). skazakhr woo Re Gv NEL no. 1309-77 Dep.). 1977. (Lacking Latin diagnosis’) Synonym: Tabanomyces milkoi (Dudka & Koval) Couch, Andreeva’, / Laird’ Wx, \Nolani,;,. "Proc, Natl SAcad-mocis. (U mero) Jie Body of the infested nematode containing unsegmented allantoid thalli, 2-5 pm wide; sporophcres erect, unseg- mented, 18-71 pm long, after penetrating the host cuticle forming an apical hook upon which a single terminal spore develops, growth continuing by sympodial proliferation; sterigmata formed from the sporophore hook, becoming emptied of cytoplasm, 2-3.5 pm wide proximally, tapering to 0.8-1.2 pm wide at the spore; spore spherical, echinulate, 5-8 pm in diameter, adhering to the nematode when detached; resting spores 18-24 pm, thick walled, with a single large oil drop, found within dead nematodes. 496 ZYGNEMOMYCES ECHINULATUS Miura. Fig. 1. (A-E) Stages in sporophore development, showing acropetal maturation of the spores; (F) zygospores formed following scalariform conjuga- 2E represents a portion of tion. Redrawn from Miura the iconotypus. Bar=10pm. (1973). Fig. 497 ZYGNEMOMYCES PENDULATUS (McCulloch) Tucker. Fige 2. (A-E) Stages in sporophore development, showing the apical hook that becomes the pedicel, and continued growth by sympodial proliferation; (D-E) mature echinulate spores, the pedicel making an acute angle with the sporophore base; (F) zygospores within the nematode. Drawn from type slide. Bar=10pm. 498 Zygnemomyces was established by Miura (1973) who distinguished Ze echinulatus from Ballocephala and Meristacrum by three characters: echinulate spores (though this is no longer valid with the discovery of Ballocephala verrucospora by Richards [1970]), spores borne on pedicels (and hence not discharged), and octangular zygospores formed by scalariform conjugation. The first two characters also are found in ae pendulatus, and a careful reexamination of spore development has shown that both species have sympodially proliferating sporophores (Figs. 1 & 2). The two species can be distinguished on the basis of sporophore size (Table 4), spore number (Table 3), and orientation of the pedicel. The three multispored genera: Meristacrum, Balloceph- ala, and Zygnemomyces, as here treated are coherent genera, and each is quite easily distinguished from the others. Meristacrum, with basipetally septate sporophores contrasts with the nonseptate sporophores of Ballocephala and Zyg- nemomyces. While spores develop acropetally in both of these genera, the manner in which the spores are formed and discharged (i.e., borne on propulsive sporiferous cells vs. pedicels formed from the sporophore apex) is significantly different. MACRO BIOTOPHTHORA Reukauf emend., Centralbl.. Bak tewParasitt mintekt 2 Abe. 0 ly 6S roo Ue oie Type species: Macrobiotophthora vimariensis Reukauf. Cen- tralbl.) Bakt @ Parasit) SIntekt Ww AD tee W637 90 ee Ol. Vegetative thallus developing internally within host, giving rise to external hyphae and sporophores on which Single terminal spores are borne; spores obovate, flat based, forming secondary spores or germinating vegetative- ly; conjugation scalariform or lateral forming zygospores, resting spores absent, or producing chlamydospores. This |genus was established by (ReukaulegCl 9 PZ) ee fOu uM. vimariensis, a fungus which he found parasitizing Hypsibius dujardini (Doyere) (=Macrobiotis lacustris Duj.) living near a spring in Belvederer Parks in Weimar, Germany. This fungus produced hyphal bodies which digested the fat bodies of the tardigrade and then gave rise to a multitude of sterile aseptate hyphae which penetrated the cuticle. Primary spores were obovoid, uninucleate and were produced terminally. These spores either germinated directly or gave rise to secondary spores on slender’ sporophores. While no other fungi similar to M. vimariensis have been isolated from tardigrades, the following nematode parasite is sufficiently similar to warrant transfer to this genus. Other species: 499 MACROBIOTOPHTHORA VERMICOLA (McCulloch), comb. nov., Basionym: Entomophthora vermicola McCulloch. Trans. Digiueme My .COlWmOOC O08 al/.3 ame OP TGS.0l—2 )i.ue lO? ye Body of infested nematode filled with nonseptate hyphae, 5-7 wm in diameter, developing erect hyphae which penetrate the nematode cuticle; external hyphae hyaline, unsegmented, flexuous (40-) 110-160 x 3-6 pm; sporophores Py oURXmO-G NM, eStralght (Or bent at, the point of emergence (OZ Us) aul oranched s(branchedtsaccordings to McCulloch): primary spores terminal, obovoid, 16-22 x 9-14 pm (rarely to 30 x 16 pm), pleurinucleate and unitunicate, discharged at maturity, adhering to the cuticle of a passing nematode and initiating new infections; secondary spores obovoid medialy constricted, nonpapillate, 14.4-15.5 um long, 9- iS soeaulleeineadiameterns narrowing at a ineck ™to= /-4—¢. 9° um; secondary sporophores 17-20 pm long, 3.5-4.5 pm at base, tapering to 1.8-2.2 wm at spore; zygospores 17-22 um, dodecahedroid, faces centrally depressed with 4-6 edges. Of the three type slides, only the one containing the nematode illustrated in McCulloch (1977: Fig. 1) remains DncaCt ma Ghigs., 3A~—J))% External hyphae are present, but these are not the morphologically modified cystidia found in species of Erynia (Remaudiére and Hennebert 1980, Humber 9S 1s) s Figure 3C shows the appearance of several of the developing spores with a thin, hyaline outer wall layer separated from the spore wall proper, and the: retraction of the cytoplasm from the spore apex. While the separated wall resembles the image seen in spores with bitunicate walls, these effects are probably due to the lactophenol mounting medium. The secondary spores are obovoid and medially constricted, with a distinct neck below a more globose region (Fig. 3H-J). A basal papilla is lacking making secondary spore discharge unlikely. The sporophores On which they are borne taper to a narrow point of attach- ment, but they are not the true capillary sporophores of Other genera. Macrobiotophthora vermicola was originally placed in Entomophthora, a genus to which it is no longer referable. Living material of this species obtained from E. Bernard, permitted additional characterization of the species, and the following observations are in conflict with the origin- aly description. Thies pramany SspOlese:contal na 4—O sin tomo— phthora-type nuclei and are forcibly discharged by papilla eversion. Secondary spores are medially constricted and passively detached. Dodecahedral zygospores are formed following scalariform or lateral conjugation. Two Conidiobolus-like fungi reported from nematodes (Castaner 1968, Amin and Webster 1980) are probably species of Macrobiotophthora. However, nuclear morphology and num- ber are unspecified, making placement of these two fungi provisional. The method of thallus and primary spore form- ation in all three nematophagous fungi is similar, but dif- ferences include the absence of zygospores (although these will probably be found), forcible discharge of secondary spores, production of microspores (?), and variation in 500 SUL spore Size (see Tables 2-5). Additional reports of para- sites on terricolous animals may increase further the size of Macrobiotophthora by at least two, if not more species. IV. FAMILIES OF THE ENTOMOPHTHORALES. At present, the order is variously divided, with one family, Entomophthoraceae (Waterhouse 1973); two families, Entomophthoracee and Basidiobolaceae (Alexopoulos and Mims 1979, Benjamin 1979); or three families, Entomophthoraceae, Ancylistaceae, and Basidiobolaceae (Ubrizsy and V6érés 1966, Batko 1974). Although three characters: karyology, spore discharge mechanism, and conjuation might logically be used to delimit these families, only karyology results in a division consistent with current views of the order. Lee Separation of the genera on the basis of spore discharge method would result in four groups. The basic papilla-eversion method has been modified in Entomophthora to form a sporophore-cannon. The significance of this is reduced, Since the secondary spores of E. muscae are still dispersed by the papilla-eversion method. Samia ray, Basidiobolus has a vestigial spore papilla, although the propulsive force is provided by the rupture of a subspore swelling. Less obvious is the origin of the Ballocephala dis- charge vesicle. The suggestion that the vesicle is homo- logous with the primary spore (Drechsler 1951), and gives rise directly to sessile secondary spores that (like capil- lispores) are not dispersed, is consistent with the basic developmental patterns of the Entomophthorales. Lt.also should be noted that spore discharge was lost independently by Massospora and Zygnemomyces. From the above observa- tions, it can be concluded that spore discharge mechanism cannot be used alone for family delimitation. 2ie Details of gametangial conjugation and resting spore morphology are also possible criteria for family delimitation. Zygospore characteristics have proven to be useful and conservative within the other orders of the Zygomycotina (Benjamin 1979). The Basidiobolaceae, with its beaked gametangia, is distinct from all other members of the order. The tendency of Conidiobolus and Ancylistes to form zygospores in line with the unequal gametangla Fige 3. MACROBIOTOPHTHORA VERMICOLA (McCulloch) Tucker. (A-H) Spores and sporophores drawn from the type slide. (A-B) Devel- opment of primary sporophores; (C) sporophore with retracted cyto- plasm and separated outer wall layer; (D) primary spore with distinct columella; (E) spore with everted papilla; (F) germinating primary spore; (G-J) development of secondary spores, (I-J) medially con- stricted obovoid secondary spores on tapering secondary sporophore. (K-P) Conjugation and zygospores from living culture. Scalariform (K) and lateral (L) conjugation; (M-N) developing zygospores; Mature Zzygospore in optical section (0) and surface aspect (P).- Bar=10pm. 502 contrasts with the laterally budded zygospores of Entomo- phthora. “Yet edivision, of the Order einto slaniliccmonmtiis character alone is problematical due to the difficulties in interpreting zygospore development in many species, and the absence of conjugation in others. Until more research is done on Entomophthorales, the utility of zygospore charac-— ters will remain suspect. oie Karyology is here regarded as the most useful character, and three distinct lines can be delimited within the Entomophthorales: Basidiobolus with large nuclei with central nucleoli; Entomophthora, Entomophaga, Exrynila, Strongwellsea Batko & Weiser emend. Humber, Massospora, Completoria (Humber pers. comm.), and Macrobiotophthora have small, readily staining nuclei with heterochromatin but without a central nucleolus; and Ancylistes, Conidio- bolus, Meristacrum, Ballocephala (?), and Zygnemomyces (?) have small, poorly staining nuclei with a central nucleo- lus. The elevation of these three groups to family status is not only consistent with the currently accepted segrega- tion of the Basidiobolaceae, but also unites those genera which have similar spore discharge and zygospore character- istics and provides a logical framework for the perception of evolutionary trends within the Entomophthorales as a whole. V.- CONCLUDING REMARKS. The taxonomic revisions proposed here are based upon the evaluation of basic morphological characteristics. Although important, the host organism is deemphasized at the generic level. While research in the Entomophthorales has been and probably will continue to be centered upon the parasites of insects, the nonentomogenous fungi must not be overlooked. It is hoped that as new’ reports of nonentomogenous fungi are forthcoming, they will include full descriptions of all aspects of nuclear morphology and number, method of spore discharge, sporophore architecture, zygospore development and morphology, and thallus characteristics. Only then will a fuller understanding of the order as a whole be obtained. ACKNOWLEDGMENTS . I would like to thank R. A Humber and R. K. Benjamin for reviewing this article, and for their many comments, criticisms, and encouragments; J. Ammirati for reading the manuscript; and D. Stuntz for correcting the Latin diagnosis and proofreading the final draft. Types were lent by the following institutions: Commonwealth Mycological Institute (CMI) and Plant Pathology Herbarium, Dept. of Primary Industries, Queensland (BRIP). E. Bernard kindly supplied the new isolate of Macrobiotophthora vermicola. 503 i KEY TO THE SPECIES OF NONENTOMOGENOUS ENTOMOPHTHORALES we OPOLODNOres producing One. terminal Sporeeesecesseccacecececee 1". Sporophore producing several spores per sSporophoreseccecccce 7 N Ww 2 e Spores IDV GASROL MN ohelelelersiorene’eleferere cictetoters cele ietere etererctereisis etetercietoretetttate ON Spores ODOVat Ceres crovelercloic ec clceicie ele cierercicreteio crercioichete cvs erie ter teen 6 3 - Nuclei staining readily, parasitic on fern prothalli eeocccccceccecccsvccccccccee ecOmpletoria complens 3". Nuclei poorly staining, parasitic on desmid algae...cccreeee 4 4. Zygosporangium with several protuberances; zygospore loose within the zygosporangium........e.-eAncylistes pfeifferi 4'. Zygosporangium SMOG Mote orevere evellelel eo ololerere cle eiarcleherelererenereneresiorere ctene rs 5) 5 - Host Closterium (Placoderm Desmid); primary spores 13-17.5 PMecevvccccveccvcescccscccves eAncylistes closterii 5'. Host Netrium (Saccoderm); spores 16-25 pm....Ancylistes netrii 6 - Vegetative thallus consisting of hyphal bodies; parasitic on tardigradesSecccccecceeseeeee eMacrobiotophthora vimariensis 6'. Vegetative thallus septate-mycelial; parasitic on nematodes eeccccccceccvceseee eMacrobiotophthora vermicola 7 - Sporophores becoming transversly septate; spores developing hbasipetallyeccccccccccccvvevcccccsccsccsesevesscesessseees 8 7'. Sporophores remaining nonseptate, or producing lateral, basally septate sporiferous cells; spores developing ACLOPeCALLYcccececvccccveccrscscccvcccevvcescesvssevessocs FY 8 . Spores pyriform; zygospores ovoid, punctate; 2° spores YEPLicativeccecescccccecvvccccceccceee oMeristacrum milkoi 8'. Spores obovate; azygospores spherical, walls undulate; 2° spores passively detached.......Meristacrum asterospermum 9 . Spores produced on basally septate sporiferous cells, the rupture of which discharges the Sporeececccccscccevcee 10 9'. Spores produced terminally on the sporophore axiSececercesee 12 10 - Sporiferous cells utriculate, as large as spore or larger; spores formed along entire sporophore axiSecerceccececee 11 10'. Sporiferous cells not inflated, smaller than spore; spores clustered at the tip....eeeeeeeeeBallocephala pedicellata 11. Spores smooth, unornamented....+.e...+-eBallocephala sphaerospora 11'. Spores verrucoS€ecccccccsecevceceeseeeBallocephala verrucospora 12 . Sporophore apex straight, becoming transformed into a spore- bearing pedicel forming an acute angle with the distal end of the sporophore..-.eeeeeeeeeZygnemomyces echinulatus 12'. Sporophore apex becoming hooked, transformed into a pedicel which forms an acute angle with the proximal end of the SPOLOPhOLe oeececoescccesscssecesssuzygnemomyces pendulatus 504 LITERATURE CITED. AINSWORTH, Ge C. 1971. Dictionary of the Fungi, 6th ed. Commonw.e "Mycol. Inst., Kew. 663 pp. ALEXOPOULOS, C. Je, and C. W. Mims. 1979. Introductory Mycology, Third Eds John Wiley & Sons, New York. 632 pp. AMIN, Ee. Ne Ee, and J. WEBSTER. 1980. British Records. Trans. Brite Mycol. Soc. 74: 633-635. ARNAUD, Ge 1952. Mycologie Concréte: Genera. Bull. Mycol. Soc. Fr. 58: 181-223. ATKINSON, G. Fe. 1894. Completoria complens. Bot. Gaz. 19: 467-468. ATKINSON, Ge Fe 1895. Damping Off. New York (State) Agr. Exp. Sta. (Ithaca), Bull. 94: 231-272. BARRON, Ge Le 1977. The Nematode Destroying Fungie Topics in Myco- biol. noe 1. Canad. Biol. Publ.-, Guelph. 140 pp. BATKO, A. 1974. 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Sci. U.S.A. 703° 2299=2302. DANGEARD, Pe. Ae 1906- Recherches sur le développement du périthéce chez les Ascomycetes. Le Botanist 9: 157-303. DAVIDSON, Je Ge Ne, and Ge Le BARRON. 1973. Nematophagous fungi: Meristacrum. Canad. J.» Bote 51: 231-233. DAVIDSON, J» Ge Ne, and Ge Le BARRON. 1973a- Nematophagaous fungi: Haptoglossa. Canad. J- Bot. 51: 1317-1323. DRECHSLER, Ce 1940. Three fungi destructive to free-living terric- Olous nematodes. J. Wash. Acad. Sci. 30: 240-254. DRECHSLER, C. IRIE An entomophthoraceous tardigrade parasite producing small conidia on propulsive cells in spicate heads. Bull. Torrey Bot. Club 78: 183-200. DRECHSLER, C. 1955. A small Conidiobolus with globose and with elongated secondary conidia. J. Wash. Acad. Sci. 45:114-117. HUMBER, R. A. 1981. An alternative view of certain taxonomic cri- teria used in the Entomophthorales (Zygomycetes). Mycotaxon 13: 191-240. INGOLD, C. T. 1934. The spore discharge mechanism of Basidiobolus ranarume New Phytol. 33: 274-277. 905 INGOLD, C. Te. 1971. Fungal spores: their liberation and dispersal. Claredon Press. Oxford. KERRY, Be Re 1974. A fungus associated with young females of the cereal cyst-nematode Heterodera avenae. Nematologica 20: 259-260. KERRY, Be Re, and De He CRUMP. 1980. Two fungi parasitic on females of cyst-nematodes (Heterodera spp.). Trans. Brit. Mycol. Soc. 74: 119-125. LEITGEB, M. He. 1881. Completoria complens Lohde, ein in Farnpro- thallien schmarotzender Pilz. Sitzungsber. Akad. Wiss., Math.-Naturwiss. Cle, Abt. 1. 84: 288-324. LOHDE, G.- 1874. Uber einige neue parasitische Pilze. Tagebl. Ver- samml. Deutsch. Naturf. Aerzte 47: 203-206. MACLEOD, D. Me, and Es. MULLER-KOGLER. 1970. Insect pathogens: species originally described from their resting spores mostly as Tarichium species (Entomophthorales: Entomophthoraceae). Mycologia 62: 33-66. MCCULLOCH, Je Se 1977. New species of nematophagous fungi from Queensland. Trans. Brite Mycol. Soc. 68: 173-179. MIURA, Ke 1973. Three entomophthoralean parasites of nematodes collected from Japan. Rep. Tottori Mycol. Inst. (Japan) 10: 517-522. MOORMAN, Ge We 1976. Mitosis in Ancylistes spe Mycologia 68: 902- 909. NAM, E. A., and A. M. DUBITSKII. 1977. A review of the systematic status of the fungus Coelomomyces milkoi and a description of a new genus of the entomophthorous fungus Pseudocoelomomyces (Entomophthorales). Inst. Zool. Alma Ata Acad. Nauk Kazakh. SSR. (VINITI no. 1309-77 DEP., UDK 591.69-577.1:576). PFITZER, Ee 1872. Ein neuer Algen-Parasit aus der Ordnung der phycomyceten. Monatsb. Konigl. Preuss. Akad. Wiss. Berlin. 379-398. POHLAD, Be. Re, and E. C. BERNARD. 1978. A new species of Entomoph- thorales parasitizing tardigrades. Mycologia 70: 130-139. REMAUDIERE, Ge, and G. C. HENNEBERT. 1980. Révision systématique de Entomophthora aphidis Hoffm. in Fres. Description de deux nou- veaux pathogénes d'aphides. Mycotaxon 11: 269-321. REMAUDIERE, Ge, and S. KELLER. 1980. Revision systématique des genres d'Entomophthoraceae a potentialité entomopathogene. Mycotaxon 11: 323-338. REUKAUF, Ee 1912. Ein verderber des Wasserbadren Macrobiotus lacus- tris Duj., Macrobiothophthora vimariensis (Reukauf). Central- bl. Bakt. Parasite. Infekt. Abte 1, 63: 390-393. RICHARDSON, Me. Je 1970. Ballocephala verrucospora Spe nove, para- sitizing tardigrades. Trans. Brit. Mycol. Soc. 55: 307-309. SOPER, Re Se 1974. The genus Massospora, entomopathogenic for cica- das. Part Ie Taxonomy of the genus. Mycotaxon 1: 13-40. THAXTER, Re 1888. The Entomophthoreae of the United States. Mem. Boston Soc. Nature Hist. 4: 133-201. TRIBE, He Te. 1977. A parasite of white cysts of Heterodera: Caten- aria auxiliaris. Trans. Brit. Mycol. Soc. 69: 367-376. UBRIZSY, Ge, and Je VOROS. 1966. A new conception in the review on the phylogeny and system of fungi. Acta Botan. Acad. Sci. Hunge 12: 199-220. MYCOTAXON Vols) XL Bie NOt opp D eS UO sono October-December 1981 ERYNIA (ZYGOMYCETES: ENTOMOPHTHORALES ) : EMENDATION, SYNONYMY, AND TRANSFERS RICHARD A. HUMBER USDA-SEA-ARS Insect Pathology Research Untt Boyce Thompson Instttute at Cornell Tower Road, Ithaca, New York 14853, USA DS RAG IG BN beers, NSERC/Agrtculture Canada, Research Branch Researeh Statton Harrow Harrow, Ontarto NOR 1G0, Canada SUMMARY The emended sense of Erynia Nowakowski (Entomophthor- ales: Entomophthoraceae) is limited to species having uni- nucleate, bitunicate primary conidia borne on (digitately- and apically) branched conidiophores and forcibly discharged by the eversion of the basal papilla. This emendation spe- cifically (1) rejects a separation of Zoophthora Batko from Erynta based on the presence of capilliconidia or any other criterion, (2) rejects any synonymy of Strongwellsea Batko & Weiser emend. Humber with Zrynta, and (3) excludes from Erynta any species having conidia which are multinucleate, unitunicate, or borne on simple (or rarely or basally branched) conidiophores. Zoophthora Batko is rejected as a later synonym of Erynta Nowak., and the remaining species of Zoophthora not previously placed in Erynta are trans- ferred. All recognized species of Erynta sensu Humber & Ben-Ze'ev are cited. The recognition of two genera, Zoophthora Batko (1964a) and Erynta Nowakowski (1881), separated principally by the formation of capilliconidia (secondary conidia formed atop Capillary conidiophores) by Remaudiére and Hennebert (1980) has raised a problem in synonymies (Humber 198la): If the type species of Zoophthora, Z. radtcans (Brefeld) Batko (= 507 Entomophthora sphaerosperma Fres. sensu Thaxter 1888), and the type species of Erynta, EF. ovtspora (Nowakowski) Nowak- Owski, are accepted on any basis to be in distinct genera, then both of these generic names may stand. If, however, one accepts Batko's sense of Zoophthora for all species with uninucleate, bitunicate conidia borne on branched conidio- pnores (Batko 1964a-b, 1966a-b, 1974; Humber 198la-b, 1982; Ben-Ze'ev and Kenneth 198la-b, 1982a-b), then both of these species must be in the same genus. Further, the Interna- tional Code of Botanical Nomenclature (ICBN) requires that Batko should have adopted the older generic name, Erynta, rather than propose a new name for this large group of Species. Humber (198la-b; 1982), Ben-Ze'ev (1980), and Ben-Ze'ev and Kenneth (198la-b, 1982a-b) find no basis for the sepa- rate and simultaneous usage of Zoophthora and Erynta espoused by Remaudiére and Hennebert (1980) and Remaudiére and Keller (1980). Among the possible resolutions of this synonymy, the only one that is both in accord with the ICBN and the prior transfer of most Zoophthora species to Erynta (Remaudiére and Hennebert 1980; Humber 198la-b) is now to take up £Zrynta Nowakowski as the correct name of the genus which includes Entomophthora radtcans Brefeld. Accordingly, we present an emended circumscription for Erynta, and regard Zoophthora to be a younger synonym. ERYNIA Nowakowski emend. Humber & Ben-Ze'ev, emend. nov. Hl Erynta Nowakowski, 1881, Datenn. III Zjazdu Lek. Przgyr. Polak. Krakow., Seke. Bot. 6, 67-68. Zoophthora Batko, 1964, BULL ACAd aeRO LON weoCt. (NOs (Nei, Meh, MAL Gas. VEGETATIVE STRUCTURES usually hyphal bodies or mycelium, but exact form depends upon stage of development. NUCLEI easily observed, 4.5-15 um diameter; without a prominent central nucleolus; filled with conspicuous heterochromatic threads or granules. CONIDIOPHORES apically branched, forming a (usually digitate) cluster of conidiogenous cells; each branch basally septate, each conidiogenous cell usually uni- nucleate, producing a single apical conidium; some conidio- phores unbranched, producing a single terminal conidiogenous cell and one conidium. PRIMARY CONIDIA uninucleate, bituni- cate (with a separable outer wall layer except over the basal papilla); shape variable but usually ovoid, clavate, turbi- 508 nate, or elongate, radially symmetrical or curved; papilla centered on spore axis or displaced laterally; forcibly dis- charged by eversion of the papilla against the conidiophore. SECONDARY CONIDIA more or less similar to primary conidia (or with short radiate arms), formed singly on a short coni- diophore arising from a primary conidium and forcibly dis- charged by eversion of the papilla or else amygdaliform to elongate, clavate, falcate, or lanceolate, usually curved, and borne on an extended, thin capillary conidiophore aris- ing from a primary or secondary conidium. RESTING SPORES usually globose with two thickened wall layers; epispore colorless or variously colored, smooth or variously orna- mented; budded laterally or terminally from parental hyphal body whether as zygospores or azygospores. RHIZOIDS present in most species, either (1) hypha-like, simple or branched, solitary or aggregated into pseudorhizomorphs, terminating in more or less branched (but only rarely discoid) holdfast of (2) stout, unbranched, solitary or loosely fasciculate (but not aggregated in pseudorhizomorphs), terminating in a richly branched discoid or irregularly expanded holdfast; rhizoids may not form when resting spores are produced. CYSTIDIA absent in most species; simple, usually aseptate, Spear-like, extending slightly above level of conidiophores or else thick and columnar, aseptate, usually branching api- cally, extending far beyond level of conidiophores; cystidia usually absent when resting spores are formed in tne host. TYPE SPECIES: Erynta ovtspora (Nowakowski) Nowakowski, 1881, Datenn. IIT Zjazdu Lek. Prayr. Polak. Krakéu., Seke. Bot. 6, 67-68. Basionym: Entomophthora ovtspora Nowakowski, 1877, Bot. Zettg. (Letpzig) 35, 20. This emendation rejects Zoophthora as a separate genus for species forming passively detached secondary capilli- conidia as advocated by Remaudiére and Hennebert (1980). It also rejects a definition of Hrynta based on conidial morphology (Remaudiére and Hennebert 1980; Remaudiére and Keller 1980), thereby excluding Erynta carolintana (Thax.) Rem. §& Henneb. (with unitunicate, plurinucleate conidia on simple conidiophores) and restoring Strongwellsea Batko § Weiser emend. Humber (with bitunicate, uninucleate conidia on unvaryingly simple conidiophores; cf. Remaudiére and Keller 1980, Humber 198la, 1982). This emendation also acknowledges the formation of multiradiate secondary coni- dia by some riparian species (Descals et al. 1981). 509 The formal adoption of Erynta as the nomenclaturally correct name for the genus circumscribed above requires the transfer of all remaining names placed in Zoophthora but not yet formally acknowledged as species of Frynta. ERYNIA APHIDIS (Hoffmann in Fresenius) Humber § Ben-Ze'ev, comb. nov. BASIONYM: Entomophthora aphidis Hoffmann in Fresenius, 1856, Ab- handl. Senekenberg. naturf. Ges. 2: 208 emend. Remaudiére § Henne- bert, 1980, Myecotaxon 11: 290, NON Entomophthora aphtdis Hoffm. in Fres. sensu Thaxter, 1888, Mem. Boston Soc. Nat. Htst. 4: 175. ERYNIA CANADENSIS (MacLeod, Tyrrell, § Soper) Humber § Ben-Ze'ev, comb. nov. BASIONYM: Entomophthora canadensis MacLeod, Tyrrell, §& Soper. 19/9, Candd..d. (Bot, 67: 2664. ERYNIA CRASSITUNICATA (Keller) Humber § Ben-Ze'tev, comb. nov. BASIONYM: Zoophthora crassttuntcata Keller, 1980, Sydowta Ann. MYCOU. OCP a lan moos 1102 ERYNIA COLEOPTERORUM (Petch) Humber §& Ben-Ze'ev, comb. nov. BASIO- NYM: Entomophthora coleopterorum Petch, 1932, Trans. Brit. Mycol. OCm ue CiaRL es ERYNIA CRUSTOSA (MacLeod §& Tyrrell) Humber §& Ben-Ze'ev, comb. nov. BASIONYM: Entomophthora crustosa MacLeod §& Tyrrell, 1979, Canad. Enconmol edd sel 138. ERYNIA ELATERIDIPHAGA (Turian) Humber, Ben-Ze'ev, §& Kenneth, comb. nov. BASIONYM: Entomophthora elateridtphaga Turian, 1978, Mtt. Schwetz. Entomol. Ges. 51: 398. ERYNIA GEOMETRALIS (Thaxter) Humber § Ben-Ze'ev, comb. nov. BASIO- NYM: Empusa (Entomophthora) geometralts Thaxter, 1888, Mem. Boston DOC NOL BOLSC. a4 ce 1/0. ERYNIA HENRICI (Molliard) Humber § Ben-Ze'ev, comb. nov. BASIONYM: Entomophthora henrict Molliard, 1918, Compt. Rend. Hebd. Séances ACG OCL en earre 16757958; ERYNIA JACZEWSKII (Zaprometov in Jaczewski § Jaczewski) Humber, comb. nov. BASIONYM: Tartehtum jaczewskitt Zaprometov in Jaczewski § Jac- zewski, 1931, Opredelttel grtbov 1: 225. ERYNIA LANCEOLATA (Keller) Humber § Ben-Ze'ev, comb. nov. BASIONYM: Zoophthora lanceolata Keller, 1980, Sydowta Ann. Mycol., Ser. II, 6059 155% ERYNIA NEBRIAE (Raunkiaer) Humber & Ben-Ze'tev, comb. nov. BASIONYM: Entomophthora nebriae Raunkiaer, 1892, Bot. Tidsskr. 18: 108-111. ERYNIA OCCIDENTALIS (Thaxter) Humber §& Ben-Ze'ev, comb. nov. BASIO- NYM: Empusa (Entomophthora) occidentalts Thaxter, 1888, Mem. Boston DOC MMIC EU BHESt ee al7.1. ERYNIA ORIENTALIS (Ben-Ze'ev & Kenneth) Humber, Ben-Ze'ev, & Kenneth, comb. nov. BASIONYM: Zoophthora ortentalis Ben-Ze'ev §& Kenneth, 1981, Phytoparastttca 9: 35. ERYNIA PHALLOIDES (Batko) Humber § Ben-Ze'ev, comb. nov. BASIONYM: Zoophthora phalloides Batko, 1966, Acta Mycol. 2: 7. ERYNIA PHYTONOMI (Arthur) Humber, Ben-Ze'ev, & Kenneth, comb. nov. BASIONYM: Entomophthora phytonomt Arthur, 1886, Bot. Gaz. (Craw- FOVUSeL Lie) s11si4. ERYNIA RADICANS (Brefeld) Humber, Ben-Ze'ev, § Kenneth, comb. nov. BASIONYM: Entomophthora radicans Brefeld, 1870, Bot. Zeitg. (Letpaztg) 28: 186. 510 Entomophthora coleopterorum Petch (1932) is included in the above list because of the conjunction of long, narrowly | oval conidia (32-44 x 8-14 um), two fascicles of stout rhi- zoids emerging from specific points on the ventral thorax, and cystidia (Petch 1944). This set of characters occurring together is known only from Hrynta species although it must still be determined that £. coleopterorum has uninucleate, bitunicate conidia and branched conidiophores. Petch (1944) regarded this species to be a probable synonym of Entomoph- thora carpenttert Giard (1888) whose description was based wholly on the unique mode of rhizoidal attachment of affect- ed elaterid beetles. Turian (1957) identified as &. car- penttert a fungus he found on beetles which died head-down and were attached to the plant by two fascicles of rhizoids; he claimed this fungus produced uninucleate, globose conidia. Humber (198la) raised doubts about Turian's interpretation and discussed the possible synonymy of F£. carpenttert and E. coleopterorum. Until any extant material of these collec- tions can be re-examined, there is no doubt that the best characterized of these collections is FE. coleopterorum, and that this species is more certainly a species of Erynta than of any other entomophthoralean genus. Entomophthora henrtct Molliard (1918), which was de- scribed from mosquitoes (Culex pipiens), 1S accepted here as an Erynta species by virtue of its much branched conidio- phores, presence of cystidia (which Molliard stated were also produced in culture), and of rhizoids with discoid or irregu- larly ramified holdfasts. It remains to be established whether the conidia of this species are uninucleate and bi- tunicate as may be predicted from this constellation of other characters. Hrynta henrict resembles EF. ovtspora, but insuf- ficient information about the morphological plasticity of either species currently €xistsPto supports thespropost trons, Waterhouse (1975) of their synonymy. Entomophthora nebriae Raunkiaer (1892), which attacks carabid beetles (Webria brevtcolis), has elongate conidia (28.0-37.0 x 10.0-13.0 um) borne on richly branched conidio- phores. Rhizoids are present, but their morphology is not described. No secondary conidial forms or cystidia are men- tioned by Raunkiaer (1892). The resting spores of this spe- cies are formed outside of the host body, and are spherical, 36-50 wm in diameter, with a pale brown, smooth, and rather thin wall. Batko (1966b) regarded this species to belong in Zoophthora subg. Zoophthora. Zoophthora jacezewskit (Zaprometov in Jaczewski §& Jaczew- Ski) Batko (1964b) was not discussed by Remaudiére and col- on leagues in their revision of entomophthoralean taxonomy. We have been unable to obtain the literature to confirm either the nomenclatural validity of Tartchtum jaczewskit Zaprometov in Jaczewski § Jaczewski (1931; see also MacLeod and Muller- Kogler 1970) or the appropriateness of Batko's acceptance of Entomophthora sabrit Rozsypal (1951; an invalidly published nomen nudum) as the conidial state of T. jaczewskit. One of the present authors (RAH) feels that it is not unreasonable to accept both Batko's acknowledgement of 7. jaczewskit to be validly published and of his synonymization of £. zabrit with this species. Those species previously described as or transferred to Erynta and accepted according to the above emendation in- clude the following: Erynta amertcana (Thaxter) Remaudiére §& Hennebert (1980) Erynta aquattca (Anderson & Ringo ex Anderson § Anagnosta- kis) Humber (198la) non Zrynta aquattca (Anderson § Ringo) Remaudiére § Hennebert (1980) Erynta blunktt (Lakon ex Zimmermann) Remaudiére §& Hennebert (1980) Erynta brahminae (Bose & Mehta) Remaudiére §& Hennebert (1980) (possibly synonymous with Lrynta echtnospora) Erynta bullata Thaxter §& MacLeod in Humber (1981b) non Erynta bullata (Thaxter in Povah) Remaudiére §& Hennebert (1980) Erynta calltphorae (Giard) Remaudiére & Hennebert (1980) Erynta eontca (Nowakowski) Remaudiére § Hennebert (1980) Erynta ereatonott Yen in Humber (1981b) non E£rynta creato- nott (Yen) Remaudiére §& Hennebert (1980) Erynta curvtspora (Nowakowski) Nowakowski (1881) Erynta delptntana (Cavara) Humber (1981a) Erynta delphacts (Hori) Humber (1981la) Erynta diptertgena (Thaxter) Remaudiére §& Hennebert (1980) Erynta echinospora (Thaxter) Remaudiére §& Hennebert (1980) Erynita erinacea (Ben-Ze'ev §& Kenneth) Remaudiére §& Hennebert (1980) Erynta formtcae Humber §& Batazy in Humber (1981b) Erynta gloeospora (Vuillemin) Remaudiére §& Hennebert (1980) Erynta gractlts (Thaxter) Remaudiére §& Hennebert (1980) Erynta montana (Thaxter) Remaudiére & Hennebert (1980) Erynta myrmecophaga Turian & Wuest in Humber (1981b) non Erynta myrmecophaga (Turian §& Wuest) Remaudiére §& Henne- bert (1980) Erynta neoaphidts Remaudiére §& Hennebert (1980) [= Entomoph- thora aphtdis Hoffmann in Fresenius sensu Thaxter (1888) ] Erynta nouryi Remaudiére §& Hennebert (1980) [= Entomophthora 5i2 exitttalts Hall §& Dunn sensu Gustafsson (1965) ] Erynta ovtspora (Nowakowski) Nowakowski (1881) Erynia phalangtetda (Lagerheim) Remaudiére § Hennebert (1980) Erynita rhtzospora (Thaxter) Remaudiére § Hennebert (1980) Erynia sepulchralis (Thaxter) Remaudiére §& Hennebert (1980) Erynta vartabtlts (Thaxter) Remaudiére § Hennebert (1980) Erynta virescens (Thaxter) Remaudiére §& Hennebert (1980) Erynta vomttortae (Rozsypal) Remaudiére & Hennebert (1980) Doubtful or) excluded species. Erynta carolintana (Thaxter) Remaudiére §& Hennebert (1980) = Entomophthora carolintana (Thaxter) Keller, 1978, Sydow- ta Ann. Mycol., Ser. II, 31: 88 = Empusa caroltntana Thax- terl 188Syilen. BOSton pOc mNOe. HUScem love Erynta castrans (Batko §& Weiser) Remaudiére & Keller (1980) = Strongwellsea castrans Batko & Weiser, 1965, J. Invert. Pathol. 14: 463 emend. Humber, 1976, Mycologia 68: 1056. Erynta magna (Humber) Remaudiére & Keller (1980) = Strong- wellsea magna Humber, 1976, Mycologta 68: 1057. Zoophthora extttalis (Hall & Dunn) Batko, 1966, Bull. Polon. AGadieSets, eoecryusct. Biola io 40 Sma pmeiLomopiT ora exttralts Hall & Dunn, 1957, Htlgardta 27: 163, Zoophthora ferruginea (Phillips in Houghton & Phillips) Batko, 1966, Acta Mycol. 2: 19 = &ntomophthora ferrugtnea Phillips in’ Houghton @ Phillips, 1886, Ari. Mao. Nacvweeer. Ser. 5, 18: 6. | Zoophthora forftculae (Giard) Batko, 1964, Bull. Potlon. Acad. Sev. , Sen. SCt.Biol. “12: A040= | Empusasforficulae(Gianw Petch, 1944) Trans. Broce Mycol. WSOC. 427-8 =e LOmO pi thora forftculae Giard, 1889, Bull. Set. France Belgtque AOS PANE Empusa carolintana and the two species of Stronguellsea are excluded from Zrynta for reasons mentioned above and discussed extensively by Humber (198la, 1982). Remaudiere and Hennebert (1980) provide strong reasons to regard both Entomophthora extttalts Hall § Dunn (1957) and Entomophthora ferruginea Phillips in Houghton and Phil- lips (1886) as nomtna confusa; we agree with and accept these determinations. It is impossible to know exactly what fun- gus is described or illustrated by Hall and Dunn (1957); this Situation is not eased by the fact that no type was declared for E. exttralts (nor did the ICBN require typification until the year after the publication) of ‘this species) or thatethe cultures deposited by the authors of this taxon in various major culture collections are found to be either Contdtobolus 513 thrombotdes Drechsler (= Entomophthora virulenta Hall § Dunn) or Erynta radicans (see Remaudiére and Hennebert 1980). The fungus regarded to be EF. exttialts by Gustafsson (1965) was redescribed by Remaudiére and Hennebert (1980) as Erynta nouryt. Similarly, the real identity of Entomophthora fer- rugtnea remains indeterminable in view of the three separate senses in which this name has been applied (see Remaudiére and Hennebert 1980). Batko (1964b) offered no explanation for his assignment of Entomophthora forftculae Giard to Zoophthora. Neither Remaudiére and Hennebert (1980) nor Remaudiére and Keller (1980) mentioned this species in their treatments of Zooph- thora and Erynta, possibly because Giard (1889) mentioned no secondary conidia (the character used by Remaudiére et al. to separate these two genera), and possibly because of the uncertainty which surrounds the identity of this fungus. PhemconlulamoOteGlardusmspeC1eSma tem exe OuLmuupet Onc om Ga in, with a length/width ratio of = 3; the conidia in the only othexrscollections attributed) to this species: are 18=21)x 8-10 Um (Rostrup 1893) and 20-30 x 15-18 um (Petch 1944), with L/W ratios of € 2. Giard, who did not recognize the widely used distinction between Empusa (with simple conidio- phores) and Entomophthora (with branched conidiophores) , noted the conidiophores of Entomophthora forftculae to be only infrequently branched (''peu ramifies''); those of the fungi reported by Rostrup and Petch could not have been pro- minently or digitately branched since Petch (1944) reassigned this species to Hmpusa. Rhizoids were not noted in any of these three collections (and note that Batko 1964a required rhizoids to be present in species of Zoophthora!). Despite the fact that L/W ratios of 3 or more are known only from Erynta species (with the sole exception of Balazy's undescri- bed species noted by Humber 198la), the absence of informa- tion regarding the wall structure and nuclear number of the Conidid.sand of the actual nature of the yconidiophores:sug- gests that no definitive generic assignment for Entomophthora forfteulae is advisable in the absence of other characters so frequently noted in species of Erynta, It seems doubtful thatethe collections of fungi by Rostrup (1893) and Peteh (1944) are conspecific with EF. forftculae Giard; the true identity of each of these fungi remains to be established by re-examination of any existing herbarium specimens. At least two aspects of this emendation of Erynta de- serve further comment. It is important to re-emphasize that this nomenclaturally required change from Zoophthora to Ery- nia has also necessitated the shift of the type species from 514 the very common Erynta radtcans (= Entomophthora sphaero- sperma Fres. sensu Thaxter) (see Batko 1964a; Remaudiére and Hennebert 1980) to the less well known Zrynta ovtspora. Re- gardless of its nomenclator, the fungus which is still most widely known as Entomophthora sphaerosperma has served as the type for a genus distinguished primarily by the presence of rhizoids and branched conidiophores ever since Nowakowski (1883) affirmed a real taxonomic difference between kmpusa and Entomophthora. Entomophthora aphtdis Hoffm. in Fres. was originally described only from its brown, roughened resting spores; the conidial state of this species was not found or described by Hoffmann. In his influential monograph, Thaxter (1888) ap- plied the name Empusa (Entomophthora) aphtdts to an excep- tionally common fungal pathogen of aphids which produces conidia and whose resting spores may be assumed to exist but have never been found. Remaudiére and Hennebert (1980) were successful in rediscovering Hoffmann's fungus and demonstra- ted unequivocally that Thaxter had misapplied Hoffmann's spe- cific name: The conidia of #. aphtdts Hoffm. are elongate- fusoid and produce secondary amygdaliform capilliconidia atop Capillary ‘conidiophoreés. Ihe) presence of capil liconidia in EH. aphtdis but not in Thaxter's fungus was used by Remaudiére and Hennebert to place these two fungi into separate genera as Zoophthora aphtdits (Hoffm. in Fres.) Batko and Erynta neo- aphtdts Rem. §& Henneb. (the new name provided for Thaxter's misapplied sense of FE. aphtdis). The emended sense of Erynta provided here places both of these species together in the Same genus. » It wil lebevespecially amportanty then, storsstus dents of these fungi not to confuse these two similar speci- fic names and to be certain that they do not apply the name Erynta aphtdts (a very uncommon fungus with a seemingly nar- row host range and localized distribution) when the fungus they refer to is actually Erynta neoaphtdts (a very common Species with a wide host range and occurring in most parts of the world). In addition to the species of Hrynta discussed above, we are aware of two additional new species awaiting publi- cation: Kramer (1981) is proposing a new species of Hrynta from snipe flies (Diptera: Rhagionidae). Ben-Ze'ev and Ken- neth (1981b) are proposing a new species of Zoophthora which affects froghoppers (Homoptera: Cercopidae); this species will have to be transferred to Erynta after its formal pub- lication. ets) ACKNOWLEDGEMENTS Wemwis mC Omtlianicelrs eR So stSODeEL Rh er at korn a andeD: E. McCabe for their review and constructive comment on this Manuscripts anders. .o. Kelier, RR: G. Kenneth, and J. PL. Kramer for providing invaluable information, some of which Hsm@ettnereiny presspOreinepreparationetor publications REFERENCES CITED BATKO, A. 1964a. On the new genera: Zoophthora gen. nov., Trtplospo- ytum (Thaxter) gen. nov. and Entomophaga gen. nov. (Phycomycetes: Entomophthoraceae). “Bull. Polon, Acad. Set., Sér. Sci. Biol. 12: 323-326. BATKO, A. 1964b. Some new combinations in the fungus family Entomoph- thoraceae (Phycomycetes). Bull. Polon. Acad. Set., Sér. Set. Biol. 12: 403-406. BATKO, A. 1966a. A new aphidicolous fungus from Poland — Zoophthora phallotdes sp. nov. Acta Mycol. 2: 7-13. BATKO, A. 1966b. On the busgenera of the fungus genus Zoophthora Batko 1964 (Entomophthoraceae). Acta Mycol. 2: 15-21. BATKO, A. 1974. Filogeneza a struktury taksonomiczne Entomophthoraceae. In Ewolucja btologtcana: Szktce teorettczne t metodologtczne (C. Nowinskiego, ed.). Polska Akad. Nauk, Inst. Filozof. Socjol., Wroclaw. BEN-ZE'EV, I. 1980. Systematics of entomopathogenic fungi of the "sphaerosperma group" (Zygomycetes: Entomophthoraceae) and their pro- SPeci com Oluceuineol10lOvUCalepest CONCTOL. sun. Dem Gissentdation. Hebrew University of Jerusalem. BEN-ZE'EV, I., and R. G. KENNETH. 198la. JZoophthora ortentaltis sp. nov., a fungal pathogen of Aphis cttrtcola (Homoptera: Aphididae), and two new combinations of other species of Entomophthoraceae. Phytopaga- eevee 92) 55-42) BEN-ZE'EV, I., and R. G. KENNETH. 1981b. JZoophthora radtcans and Zoophthora petcht sp. nov. (Zygomycetes: Entomopnthorales), two spe- cies of the "sphaerosperma group" attacking leaf-hoppers and frog- hoppers (Homoptera). Entomophaga 26: 131-142. BEN-ZE'EV, I., and R. G. KENNETH. 1982a. Features-criteria of taxonomic value in the Entomophthorales: I. A revision of the Batkoan classifi- - cation. In preparation for Mycotaxon. BEN-ZE'EV, I., and R. S. KENNETH. 1982b. Features-criteria of taxonomic value in the Entomophthorales: II. A revision of the genus Hrynta Nowakowski 1881 (= Zoophthora Batko 1964). In preparation for Mycotaxon. DESCALS, E., J. WEBSTER, M. LADLE, and J. A. B. BASS. 1981. Variations in asexual reproduction in species of Entomophthora on aquatic insects. Dane eer. Lee Yi COL.820C,) (728807102. GIARD, A. 1888. Sur quelques entomophthorées. Bull. Set. France Bel- gtque 19: 298-309. GIARD, A. 1889. Sur quelques types remarquables de champignons entomo- phytes. Bull. Set. France Belgique 20: 197-224. GUSTAFSSON, M. 1865. On species of the genus Entomophthora Fres. in Sweden. I. Classification and distribution. Lantbrukshogskol. Ann. 61:8 103-2127, HALL, I. M., and P. H. DUNN. 1957. Entomophthorous fungi parasitic on the spotted alfalfa aphid. Hilgardta 27: 159-181. 516 HOUGHTON, W., and W. PHILLIPS. 1886. On Aphte rwmicts, Linn., as a » pest on the Mangel-Wurzel crops in Shropshire in the autumn of 1885, and on a fungus destructive of the same Aphis. Ann. Mag. Nat. Htst. Sera o, Lore lay. HUMBER, R. A. 198la. An alternative view of certain taxonomic criteria used in the Entomophthorales (Zygomycetes). Mycotaxon 18: 191-240. HUMBER, R. A. 1981b. Erynta (Zygomycetes: Entomophthorales): valida- tions and new species. Mycotaxon 18: 471-480. HUMBER, R. A. 1982. Strongwellsea vs. Erynta: the case for a phylo- genetic classification of the Entomophthorales (Zygomycetes). In preparation for Mycotaxon. JACZEWSKI, A. A., and P. A. JACZEWSKI. 1931. Opredelitel grtbov, vob) b, Leningrad: KRAMER, J. P. 1981. A mycosis of the blood-sucking snipe-fly Sympho- romyta hitrta caused by Erynta tthacensts sp. n. (Entomophthoraceae) . Mycopathologta: in press. MacLEOD, D. M., and E. MULLER-KOGLER. 1970. Insect pathogens: species originally described from their resting spores mostly as Tartchtum species (Entomophthorales: Entomophthoraceae). Mycologia 62: 33-66. MOLLIARD, M. 1918. Sur la vie saprophytique d'une Entomophthora (E. henrict n. sp.). Compt. Rend. Hebd. Acad. Set. Parts 167: 958-960. NOWAKOWSKI, L. 1881. O grupie owadomorkéw (Empusaceae). Datenn. III Zjazdu Lek Przyr. Polak. Krakow., Seke. Bot. 6: 67-68. NOWAKOWSKI, L. 1883. Entomophthoreae. Przyczynek do znajomésci pasor- zytnych gryzbkow sprawiajacych pomor owadéw. Pamtetn. Wydz. Akad. Unteg. w Kradow. 8: 153-183. PETCH,= I. 1932.) A list) of the entomogenous (fungivot Greate Brigain: TONS MET me MUCOL MRCOG it? LOL 18s PETCH, T. 1944. Notes on entomogenous fungi. Trans. Brit. Mycol. plete, VAS “oalestels) 4 ; RAUNKIAER, C. 1892. Et Par nye Snyltesvampe. Bot. Tidsskr. 18: 108-111. REMAUDIERE, G., and G. L. HENNEBERT. 1980. Révision systématique de Entomophthora aphtdits Hoffm. in Fres. Description de deux nouveaux pathogenés d'aphides. Mycotaxon 11: 269-321. REMAUDIERE, G., and S. KELLER. 1980. Reconsidération systématique des genres d'Entomophthoraceae 4 potentialité entomopathogénes. Myco- taxon 11: 323-338. ROSTRUP, 0. 1893. Medd. naturf. Foren. (Copenhagen), p. 87, ROZSYPAL, J. 1961 Prispevek k biologickému boji proti skudcum obili- novi parasit larev hrbace osenniho. Sborn. Ceskosl. Akad. Zemed. 24: 85-94. THAXTER, R. 1888. The Entomophthoreae of the United states) Mem: BOSCOn SOC NA ws vet.) 4.) 55-201 TURIAN, G. 1957. Entomo-mycoses dans la région de Genéve.. Mitt. Sehwetz. Entomol. Ges. 30: 93-98. WATERHOUSE, G. 1975. Key to the species of Entomophthora Fres. Bull. Brit. Myeol. (50C., 9: loreal. MYCOTAXON VO ett eeN On opp 5 1 5.74 October-December 1981 eee ae ee ee oe ee ee Se EE a ee ee ee REVUE DES LIVRES par G.L. HENNEBERT Book Review Editor, Croix du Sud 3, B-1348 Louvain-la-Neuve Belgique THE COELOMYCETES, Fungi imperfecti with pycnidia, acervuli and stromata, by Brian SUTTON, 696 p., 397 fig., 8°, hard cover, 1980. Commonwealth Mycological Institute publication, Commonwealth Agri- cultural Bureaux, Farnham, Slough SL2 3BN, England. Price UK £920 .005abroadss 133.60. The Coelomycetes by Sutton is the first comprehensive and modern contribution on that important wide spread group of fungi. After numerous publications on the Coelomycetes, including seven issues in the CMI Mycological Papers and the fundamental and synthetic chapter on Coelomy- cetes in The Fungi IVA (1973), we are really thankful for such a fine monograph covering 375 genera and 750 species. The inestimable progress realized by Dr Sutton in the taxonomy of these fungi is the recognition of conidiogenesis as a basic taxonomical criterion to be used at the level of classes and orders, the conidiomatal structure (pycnidia, acervuli, sporodochia, etc.) and morphology being useful as a secondary criterion at the suborder level or lower. Although, in that concept of the taxonomy of conidial fungi, Hyphomycetes and Coelomycetes would ideally be and are close to being unified and classi- fied according to the conidiogenesis, the conidioma from the pycnidium to the obsolete exposed conidial cell representing a continuum, the author still uses the term Coelomycete here for the practical reason of deli- miting a workable field. Accepting the major modes of conidiogenesis as defined at the lst Kananaskis Conference (1969), Sutton proposes two classes in the Deuteromycotina, the Thallodeuteromycetes and the Blasto- deuteromycetes, and five orders, the Thallales, Enterothallales, Blastales, Phialidales and Tretales, instead of the traditional Sphaerop- sidales, Melanconiales and Moniliales. The structure of the conidioma ' delimits eventually the suborders, f.i., in the Blastales, the Blasto- hyphineae (blastic Moniliales), the Blastopycnidineae (blastic pycnidial), the Blastopycnothyrineae (blastic Pycnothyriales) and the Blastostroma- tineae (blastic Melanconiales). Notice that two orders, the Enterothallales and the Tretales, are not reprensented amongst the coelomycetous fungi so far but in the Hypho- mycetes only. The Coelomycetes traditionally classified in the Pycno- thyriales and now representing the Thallopycnothyrineae, the Blastopycno- thyrineae and the Phialopycnothyrineae, are not treated for the simple reason that almost no information on their conidiogenesis is available. Sutton discuss the hypothesis of Morgan-Jones, Nag-Raj and Kendrick (1972) that a disinction might exist between an annellidic conidiogenous cell and percurrently proliferating phialide but is not convinced . He prefers to base the distinction between annellidic and phialidic 518 percurrently proliferating conidiogenous cell on the single or repetitive conidium production by each of the successive cells. There is the reason why some fungi having conidiogenous cells looking like annellides are classified as repetitively proliferating phialidic fungi. In regard of the amount of new information to be included, the author chose with reason to avoid any already published data and to refer to good descriptions, illustrations or comments published elsewhere when possible. Quite a number of genera are therefore not illustrated. But it would have been of great help to the users to reproduce, at least, an illustration of those genera, to give at oncea full picture of the group.The text of the book is concise but clear. The paper, rather thin but of good quality and flexible, makes the size of the book reasonable and its manipulation easy. The book is completed with adequate indices and a glossary. There is no need to say that the Coelomycetes are economically im- portant plant pathogens and material deteriogens. Not only should any mycologist welcome this major contribution to the systematics of fungi, so also should any plant pathologist, forest manager or microbiologist concerned with soil biology or material degradation be aware of it. Furthermore, taxonomists will appreciate being stimulated in their investigations by an author who, at almost every page of his book, points out some ways of progress. A REVISION OF CHRYSOSPORIUM AND ALLIED GENERA, by C.A.N. van OOR- SHOTS studiessin Mycology n = 20,690 =pann JON toon) amon eens De cover, 1980. CBS Publicaticon,..Baarn Nederlands PricesHrin Z5,—- Chrysosporium Corda and the related genera Myceliophthora, Emmonsia, Zymonema, Trichosporiella, Blastomyces, Glenosporella and Geomyces are revised, including 38 species and varieties. The redescribed anamorphs have either thallic or blastic conidiation and rhexolytic dehiscence. Teleomorphs belong to the Gymnoascaceae, Onygenaceae, Ascosphaeraceae ° and Sordariaceae. The genus Chrysosporium Corda is quite an homogenous grouping now, Sporotrichum pannorum Link having a distinct position in Geomyces. COELOMYCETES. VII. STEGONSPORIUM, by K.T. Van WARMELO and B.C. SUTION, Mycological’ Papers n- 145, 46 p. 7913) pl... °8 =, paper) covers 1981. Commonwealth Mycological Institute, Kew Surrey, England. CABLPublicabironasericesf.3).00. The original spelling Stegonsporium from Corda in Opiz 1926 is accepted instead of Fries's spelling Stegonosporium 1849. Two species are maintained, S. pyriforme and S. acerinum. Sixty five species are ex- cluded or questioned. The excluded species belong to Stigmina, Myxocyclus, Coryneum, Camarosporium, Trimmatostoma, Dictyodesmium, Neohendersonia and Camaropellum, with five new combinations. Two new genera, Stegonsporiopsis and Kaleidosporium are proposed with two new combinations. Stegonsporium produces annellidic conidiophores with dictyoseptate conidia amongst filiform paraphyses. CONTRIBUTION TOWARDS A RATIONAL ARRANGEMENT OF THE CLATHRACEAE, by D.M. DRING;, 96 p., 2/7 ifie=, 8 . paper cover, 1981, srepranted from Kew Bulletin, 35(1), 1980. Royal Botanic Gardens, Kew, England. Price Loe Oor These are the conclusions of a long-term interest and research in the Clathraceae by the late Dr. Dring (f 1978) and of his personal se observations in West Africa. It has been fortunate that the unfinished manuscript could be completed from the author's large collection of notes and drawings. Dring recognized 8 genera, Azeroé, Blumenavia, Clathrus, Colus, Ideodictyon, Laternea, Lysurus and Pseudocolus, and 36 species. These are fully described and illustrated, with synonymies and references to other valuable illustrations. The specimens examined are all cited, demonstrating the often restricted geographical distribution. The genus Clathrus, with 16 species, is the largest one and appears to the author as the ancester of a diversifying evolution. All the species of the Clathraceae, the exception of Clathrus ruber and C. hirudinosus repre- sented in Europe, West Asia and North Africa, are known from the other parts of the world and many are from tropical areas. THE GENERA OF FUNGI SPORULATING IN PURE CULTURE, by J.A. von ARX, Chardeedi tion. s424e peels pie oS ee kardacovers =| 96170 mCramer: Fl 9490 Vaduz, Lichtenstein. Price DM 120.-, for subscribers 100.-. This third edition of Dr von Arx's book is a fully revised and ex- panded version of the previous ones. The classification of the fungi has been emended according to recent advances. The Dothideales include the Myriangiales, Dothiorales and Pseudosphaeriales. The Pezizales include the Tuberales. The Tilletiales are reintroduced. The Ustomycetes (Ustila- Sinales, Exobasidiales, Taphrinales, Sporobolomycetales and basidiomy- cetous yeasts) are distinguished from the Endomycetales on morphologi- cal grounds and wall composition. Basidiomycetes are not otherwise trea- ted. All together 853 genera are accepted, of which only 370 are given an illustration, one third of these illustrations being new and based on Yecent studies. About 50 genera, some of them being recently described, are reduced into synonymy. Some genera rejected in recent years are re- introduced, e.g. Endomycopsella, Karakulina and Bipolaris. Illustrations and dichotomous keys result from studies of the fungi in pure culture, independent of their natural substrate. Keys have been adjusted to up- dated terminology. The 1100 literature titles include the most recent ones published in 1980. This edition, certainly like the previous ones, will be appreciated by those who have to identify fungi from sporulating cultures. MARINE MYCOLOGY, The Higher Fungi, par Jan KOHLMEYER et Erika KOHLMEVER (t210).090 tpe, 1270 fie, eo a erelaeyto1lé (1979. Academic Press, N.Y. Price US $ 71.50. 149 Ascomycétes, 4 Basidiomycétes, 56 Deuteromycétes filamenteux et 177 levures sont les champignons supérieurs récoltés 4 ce jour prés de et dans la mer. Des 209 espéces filamenteuses rencontrées, 191 sont spécifiques au milieu marin. C'est dire qu'un ouvrage intitulé Marine Mycology ne saurait @étre de pure taxonomie sans aussi considérer 1'éco- logie. C'est cette alliance qu'ont fort bien réussie les deux auteurs. Aprés un exposé des méthodes (chap. 2) 1'écologie des champignons cétiers et marins met en évidence les relations champignon-substrat chez les espéces de pleine eau, d'estuaires, de sables, des écumes, des marais salés, des algues , des lichens, des mangroves et des débris animaux et végétaux (chap. 3-19 et 21). Dans les chapitres touchant la taxonomie (chap. 20, 22-28; 421 p.) les auteurs proposent d'abord des clés dicho- tomiques basées sur la morphologie sporale ou conidienne et finement il- lustrées par Erika Kohlmeyer. Le principal chapitre 26 comporte la des- cription taxonomique des 209 champignons filamenteux, avec synonymie, photographies et bibliographie. Une nouvelle espéce et 7 combinaisons sont proposées. 49 rejections de noms douteux ou invalides sont énoncées. 5.0 Dans le dernier chapitre,les levures sont répertoriées, sans description mais avec indication de leur habitat.Prés de 1400 références bibliogra- phiques complétent l'ouvrage. En mycologues avertis, les auteurs témoi- gnent dans ce livre d'un sens aigu du détail et ume maniére esthétique de le mettre en valeur. Ce livre est une "somme" de ce domaine particu- lier de la mycologie. Sa qualité suscite 1'admiration. MICROBIAL POLYSACCHARIDES AND POLYSACCHARIDASES, par R.C.W. BERKE- LEY, G.W. GOODWAY and D.C. ELLWOOD édit., Publication de la Society for general Microbiology, xvinte4v79'p.,) 211 oe oe relic. ag 19s Academic Press, N.Y. Prix US $947.00. Ce livre est la synthése de deux symposiums tenus 4a Aberdeen en 1978, l'un sur The microbial extracellular Polysaccharides du Groupe pour 1'étude des surfaces et membranes de la Cellule microbienne, l'autre sur The microbial Degradation of Polysaccharides de la Society for general Microbiology. Sans doute, cet ouvrage sort du champ habituel de la mycologie. Cependant la composition différente des parois des champignons suivant leur classe taxonomique est une raison de considérer cet ouvrage de portée aussi fondamentale qu'appliquée. La détection, l'analyse et la production de polysaccharides et des enzymes qui les dégradent sont les objectifs d'une recherche de plus en plus active. Ce sont surtout les polysaccha- rides extracellulaires, comme le pullulan de 1'Aureobasidium pullulans, qui retiennent l'attention. La paroi fongique est donc un organe essen- tiel. Sa composition, sa structure microscopique fine, sa perméabilité, la structure biochimique, l'activité et la solubilité de ses polysaccha- rides sont autant d'aspects traités dans ce livre. La paroi se détruit autant qu'elle se construit. Les polysaccharidases produites par les champignons eux-mémes ou les bactéries associées agissent au niveau de la paroi fongique et dans le milieu, transformant les polysaccharides en nutrients recupérables. Ainsi les cellulases, chitinases, amylases, glu- canases, pullulanases des champignons ne sont pas les moins connues. Ce livre s'adresse non seulement au microbiologiste et biochimiste indus- triel, mais aussi le mycologue curieux de la nature exacte des champignons. STEROLOGICAL METHODS. VOL. I. PRACTICAL METHODS FOR BIOLOGICAL MORPHOMETRY,--par ©aR, WEIBEL, ©515ep. , milli Seretie9 toile, 31979. Academic Press. Price US $ 66.50. La premiére approche d'un étre vivant est d'abord d'en saisir la forme. Au travers du microscope, ot le relief disparait, la forme est celle d'une image, d'un profil, d'une section. Le microscopiste, le bio- logiste, le mycologue traduit la vision de la forme (morphoscopie) en un tracé d'abord (morphographie), en une description ensuite (morphologie), en une mesure enfin (morphométrie). La promotion récente de la taxonomie numérique accentue la nécessité d'une morphométrie méthodique et tradui- sible en termes de réalité. La recherche de méthodes morphométriques valables est le but delastéréologie. Ces méthodes sont mathématiques et visent a relier les paramétres tridimensionnels d'une structure spatiale aux mesures bidimensionnelles des sections de cette structure. Inversé- ment, elles fournissent la réponse a la question: comment interpréter l'image vue au microscope comme expression de la réalité spatiale. L'auteur ne traite pas des fondements mathématiques des méthodes, les réservant au volume II. Le livre est donc d'un niveau mathématique acces- sible & qui posséde des notions de mathématique moderne. Par voie d'exemples, de photos et de graphiques, 1'auteur explique les méthodes d'échantillonage des structures A mesurer et la réalisation de mesures ork qui ne faussent pas la réalité. Il serait intéressant que des mycologues expérimentent les méthodes morphométriques proposées par la stéréologie au profit de la taxonomie des champignons. BULLETIN OF MYCOLOGY. VOL. I. N°l, edited by Sultan AHMAD and S.H. IQBAL, Department of Botany, University of the Punjab, New Campus, Lahore, Autumn 1980. We welcome the appearance of this new journal devoted to taxonomic mycology from Pakistan, a country where surely much waits to be disco- vered and described. The first issue of the Bulletin of Mycology contains interesting papers on freshwater Hyphomycetes, on coprophilous Ascomyce- tes, on Agaricales and other fungi from Pakistan. We hope that the new journal receives the attention that it deserves. SEXUALITY AND PATHOGENICITY OF FUNGI, edited by R. VANBREUSEGHEM andeCh.s DEWVROEY. 2505p... 111, ,) 8°, hard «cover, 1981. Masson, 120 Bd, Saint Germain, 75280 Paris. This is the Proceedings of the Third international Colloquium on Medical Mycology organized by the Prince Leopold Institute of Tropical Medecine, Antwerp, Belgium. The question raised at the beginning of the colloquium "Is there a relationship between sexuality and pathogenicity to man in fungi?" recei- ved an answer far beyond any hope. That answer is that no relation between mating type and symptoms has been observed so far, but that a close rela- tion between the mating type and the host or biotope is demonstrated and that that relation is determining particular geographical distribution. Beside the pathogenicity of mating types, serotypes, + and - types, sexual and asexual spores, many correlative aspects of the pathogenic fungi have heen treated by the 32 contributors, like taxonomy and the relation ana- morph-teleomorph, gentics, somatic hybridization, sexual stimulation, sexual test strains, filamentization of yeasts. The book answers a number of questions raised in the taxonomy and ecology of the fungi. It offers valuable readings for any mycologist and for plant pathologists. YEAST GENETICS AND MOLECULAR BIOLOGY 1980. Reports, by A, GOFFEAU and J.M. WIAME. 102 p., 8°, paper back, 1980. Louvain-la-Neuve, Lab. Enzymology, Croix du Sud 1, 1348 Louvain-la-Neuve, Belgium, Price Workshop reports $ 10, Abstracts book $ 20. This is the Workshop Reports of the tenth International Yeast Con- ference held at Louvain-la-Neuve, September 1980. It contains 24 plenary presentations covering the research progresses on Saccharomyces cerevi- siae about its chromosome structure and replication, RNA, protein synthe- sis, cell cycle, sporulation, recombination, mutagenesis, radiation repair, expression of cloned genes, mating, regulation of C and N, metabolism, mitonchondria, killers, genetics engineering, evolution and taxonomy. It certainly leads to conclude that Saccharomyces cerevisiae is a favorite model for studying the structure and the operation of eukaryo- tic cells. The Abstracts Book was distributed at the Conference but is still available, it reproduces the abstracts of each of the 200 posters presented. INTRODUCTION TO FUNGI, by John WEBSTER, 2d edition, 669 p., 331 fig. 8°, paper or hard cover, 1980. Cambridge University Press. P.O. Box 110, Cambridge, England. Price h.c. £ 30, p.c.. 9.95. 522 After the publication of the first edition, in 1970, advances have been made in all fields of mycology. Where necessary the text has been altered or expanded. The literature cited is twice in number now. The author adopts the Ainsworth, Sparrow and Sussman's system of the Fungi, as a separate kingdom. The Myxomycetes are more extensively treated. The anamorphic fungi, the Deuteromycotina, receive a special chapter with a description of their conidiogenesis. New and detailed accounts are added on three ecological groups of fungi, the aquatic, the preda- cious and the seedborne imperfect fungi. This reinforces the aim of the author to be not purely taxonomical but to describe the fungi in their life behaviour. The clarity of Webster's writing, the precision of the data, the excellent selection of original photographs and drawings are prominent characteristics of the book. The book is an excellent manual for the university level. For reaching constantly the present limits of knowledges, it concerns all students of mycology. HOW TO KNOW THE TRUE SLIME MOLDS, by M.L. FARR, in Pictured Key Nature, Series 5132 px loostiec.., 2oxl/ecm spirale binding goLe Wm. C. Brown Co. Publ., 2460 Kerper Bd., Dubuque, Iowa 52001, USA. This "pictured key" to the Myxomycetes is intended to be introduc- tory and practical. The introduction shortly explains what Myxomycetes are, how they live, where they growth, how to collect them, how to keep them alive, how to look at them, how to describe them, how to preserve them. The keys to orders, families, genera and species are all dichotomous Specific descriptions, comments and illustrations are given at every spe- cies keyed out. Illustrations are mostly line drawings picturing frui- ting bodies and, eventually, capillitium and spores. At every taxonomic level, indeed, the keys are constructed from both macroscopic and micro- scopic characters, They thus are not to be considered as field keys. They cover a selection of 276 common North-American species of Myxomycetes. TAXONOMY OF THE INDIAN MYXOMYCETES, by T.N. LAKHAMPAL and K.G. MUKERJI.- in Bibliotheca Mycologica vol= 9785 "53l=p.5, 40Mfiee,es a, hard cover, 1981. J. Cramer, Fl 9490 Vaduz, Lichtenstein. Price: DMV1205 subscription DM 96. The authors have realised two aims. In the first part of the book, they describe, in a taxonomical order, all the species of Myxomycetes that they recorded in 1000 specimens from their respective collecting fields, i.e. Himalahal Pradesh and Delhi. The Taxonomy is that of Martin and Alexopoulos (1969) modified by Alexopoulos in Ainsworth, Sparrow and Sussman (1973). The illustrations are assembled in 28 full-page plates of black and white photographs of standard quality and 12 plates of ra- ther poor line-drawings. In the second part of the book, the authors pro- duce an updated index of all the Indian Myxomycetes records. disposed in taxonomical order, with reference to the pertinent literature. The contri- bution is important. Out of the 450 presently known species of Myxomyce- tes, 293 are recorded in India and, out of those, 111 are recently recor- ded, by the authors mostly, after the publication in 1977 of Dr Thind's monograph. BIODEGRADATION ET HUMIFICATION, ATLAS ULTRASTRUCTURAL, par G.M.OLAH, 0; REISINGER et*G.) KILBERTUS 33334 0p. 51998 fie.5..4 eo cartonna.s19/6. Les Presses de l'Université Laval, Québec. Librairie Vuibert, Paris. — En 1974, était organisé, a Nancy, le ler Colloque international sur le théme "Biodégradation et Humification" (Mycotaxon IV(1):317, 1976). 523 Les auteurs, organisateurs de ce Colloque, ont rassemblé en un atlas leurs documents de microscopie électronique illustrant d'une part les agents de la biodégradation dans le sol (bactéries, champignons, algues bleues, et microfaune) et d'autre part les victimes, la matiére organique vivante en dépérissement (celle des algues, champignons, bryophytes et phanéro- games). L'atlas est divisé en chapitres introduits par un texte et un schéma expliquant le processus de la biodégradation dans le sol. Les illustrations font face a une légende bréve, & mon avis trop bréve, mais suivie d'un grand nombre de références & la littérature. Les photographies sont de bonne qualité et suscitent l'intérét par leur choix et leur ori- 1 ginalité. On y voit des vues inattendues de bactéries du sol ou coloni- sant des hyphes fongiques. Des images en ombrage et en transmission il- lustrent la sporulation de divers champignons, un asque operculé béant, des annellides, des conidies en grappe, des pores septaux, la perforation bactérienne de parois fongiques, la désorganisation enzymatique de celle- ci, et la repousse interne d'une hyphe de régénération. Cet atlas est didactique et permettra de visualiser, dans l'enseignement, un des grand phénoméne de la biologie du sol, le "turn-over" de la matiére organique. COMPENDIUM OF COTTON DISEASES, par G.M. WATKINS, éd., in The Disea- SemCOmpendiasSeries pallu, Pomp. 64ers oo Oy pile COL. Ga BLOG Le The American Phytopathological Society, 3340 PilotKnob Road, St Paul, Minnesota 55121. Prix US$ ll. Ce compendium, comme les précédents sur les maladies du soya, du froment, de la luzerne, du mais, continue une . série fort utile 4 la clinique des plantes cultivées. Plus de quarante spécialistes ont contri- bué 4 la rédaction et l'illustration de celui-ci. Prés d'une cinquantaine de champignons responsables d'une pathogénése fongique du cotonnier sont envisagés, la symptomatologie et 1'épidémiologie décrites et le traitement indiqué. A coté des maladies fongiques, les autres maladies biotiques et abiotiques sont aussi décrites et illustrées. Une clé syn- thétique des symptOmes permet l'identification rapide des maladies. THE CORTICIACEAE OF NORTH EUROPE. Vol. 6. PHLEBIA-SARCODONTIA, par John ERIKSSON, Kurt HJORSTAM et Leif RYVARDEN, p.1051-1276, fig. DEI -ODShmO wee Droche sl Iolerunes tl ora, er..07 eboxno> se bkindern , Os OmS se Norway. Prix gNkresl20. Sont déja parus et recensés dans Mycotaxon les volumes 2 (Aleuro- discus - Confertobasidium), 3 (Cornicium - Hyphoderma), 4 (Hyphodermella - Mycoacia) et 5 (Mycoaciella - Phanerochaete). Dans le présent volume, les auteurs décrivent les genres Phlebia (31 espéces), Repetobasidium (8 espéces), Piloderma, Resinicium (3 espéces chacun), Phlebiopsis, Ra- dulomyces (2 espéces) etPhysodontia, Plicatura, Plicaturopsis, Podoscypha, Pseudomerulius, Pseudoxenasma, Pulcherricium, Punctularia, Radulodon, Ramaricium, Repetobasidiellum et Sarcodontia monospécifiques. Les auteurs proposent 1 nouveau genre (Repetobasidiellum), 9 nouvelles espéces dont 3 en Phlebia, 14 nouvelles combinaisons dont 5 en Phlebia et un nom nou- veau. Certainement, ce volume est attendu, comme le sont aussi les suivants. THE FISCHER-SMITH CONTROVERSY: ARE THERE BACTERIAL DISEASES OF PLANTS?, par Erwin F. SMITH et Alfred FISCHER, introduit et tra- duit de 1'allemand par C. Lee CAMPBELL, Phytopathological Classics n° 13, xviii + 65 p., 40 fig., 8°, broché, 1981. The American Phytopathological Society, St Paul, Minn., USA. Prix US $ 8.50. Erwin F. Smith, American plant pathologist, USDA, Washington, who has demonstrated experimentaly the pathogenicity of bacteria to plants 524 refutes theassertion of Alfred Fischer, a German professor in Botany at the University of Leipzig, of the non-existence of bacterial diseases. It was in 1897. Fischer answered by unjustifiable critics about the Smith's observations and experiments. Smith then took his own defence, ina long and convincing paper (1902). As a result, the existence of bacterial diseases was accepted, and the rivals were both promoted, the one as director of the USDA Plant Pathological Laboratory, Washington, the other as professor and director of the Botanical Gardens at the Universit ty of Basel.'The facts ardé\God's, the theories‘are human" (Lord Ripon, 1874). TROPICAL MYCORRHIZA RESEARCH, par Peitsa MIKOLA édit., Oxford SciencesPublicatsonsiyexiy +2 /ONp., oil Gere lie lL OOOmshOrethe International Foundation of Science, Oxford University Press, Walton ‘Street, (Oxford s0xX2,6DP; Englands Prix=e ilo. Ce volume est constitué des 31 communications présentées a 1'Inter- national Workshop on Tropical Mycorrhiza tenu au Ghana, en 1978, par 1' International Science Foundation (IFS). Elles se regroupent en quatre thémes: les ectomycorhizes en région tropicale, les mycorhizes de la végétation naturelle, nutrition mycorhizique des plantes tropicales et mycorhizes des plantes agricoles. Il est intéressant de constater le role grandissant des ectomycorhizes dans la reforestation des zones tro- picales dénudées. D.H. Marx fait ume revue trés large des possibilités d'inoculation de champignons mycorhizogénes, a4 travers le monde, et des méthodes utilisées. D'autres donnent les résultats particuliers obtenus en leur pays. La recherche explore aussi les mycorhizes naturelles des essences forestiéres et des plantes cultivées tropicales, en particulier de 1'Hevea, de 1'Elaeis, du cacaoyer, des Citrus et des graminées. Si on recherche les facteurs qui peuvent leur @tre favorables, on s'inquiéte de plus en plus de l'effet néfaste sur les mycorhizes des fertilisations chimiques et des herbicides en forét comme en agriculture. La mycorhization est un phénoméne tel qu'il importe de le connaitre pour le maintien du monde végétal et du notre. DEVELOPMENTAL MICROBIOLOGY. par John PEBERDY, in Tertiary Level Bi- ology Series, 230yp-,,11U., 12° ,apaper back,, 1980... Blackie-& Son Ltd Bishopbriggs, Glascow G64 2NZ. Prix £ 8.50. Ce petit livre de microbiologie est axé sur les phénoméne de crois- sance et de reproduction. Il en montre les aspects les plus fondamentaux de structure, de biochimie et de génétique cellulaires et les plus appa- rents de la morphologie, de l'accroissement et de la formation sexuée et asexuée de spores, avec le souci d'expliquer la relation entre ces aspects. Si dans un tiers du livre il est question de bactéries et virus vus sous les mémes aspects, ce livre est particulier par le choix des champignons comme matériel privilégié d'étude du développement (levures, myxomycétes, champignons filamenteux, champignons sexuées et asexuées). Ce livre est destiné au niveau du baccalauréat ou "undergraduate". BIOLOGICAL MEMBRANES, THEIR STRUCTURE AND FUNCTION, par R. HARRISON et G.G. LUNT, Tertiary Level Biology Ser«, 2e.ed., 286 po gelll oe toe L980... Blackiess SonelLtd. «Glas cow. .er isaac 3 La structure de la membrane cellulaire, sa composition biochimique, les mécanismes fonctionels de reconnaissance des macromolécules et de leur transport par la membrane cellulaire sont les sujets traités par les auteurs dans un langage clair et adapté au niveau universitaire. Cependant nous regrettons que ce livre n'envisage que la cellule animale etqu'ilnementionne qu'en passant les membranes végétale et fongique. MYCOTAXON Onc ar Ome 5 a OO October-December 1981 NE Omi ia Ce eaS XV PACIFIC SCIENCE CONGRESS The XV Pacific Science Congress will be held February 1-11, 1983, in Dunedin, New Zealand. The Pacific Sci- ence Association is responsible for the program, which is sponsored by the Royal Society of New Zealand. A fisrt circular may be obtained from The Secretary Gene- ral, 15th Pacific Science Congress, P.O. Box 6063, Dune- din North, New Zealand. IMC: ON HOUSING COSTS IN JAPAN Though it is indeed true that Japan is currently a very expensive place to visit, the organizing committee of IMC3 is well aware that most mycologists will not be able to spend great sums of money on housing for the congress. Dr. Keisuke Tubaki, a member of the Mycolo- gical Society of America's ad hoc committee on IMC3 ar- rangements, has assured us that though the meetings are planned for a downtown Tokyo hotel-convention faci- lity, there will be many hotel rooms available close by in fess prestigious hotels in the $15-20/night (1981 prices) range. There have apparently been rumors that housing costs would be substantially higher. Our Japa- nese hosts will do all they can to keep costs down for participants. The same concerns will be kep in mind for pre-congress and post-congress tours, workshops, and field trips. The Tokyo meetings will be from 28 August through 3 September, 1983. 526 AUTHOR INDEX, VOLUME THIRTEEN ALCORN, J. L. Cochliobolus ravenelit sp. nov. and C. trtpogonts sp. nov. 339-345 ALCORN, J. L. Ascus structure and function in Cochltobolus species 349- 360 ARENDHOLZ, WOLF-RUDIGER & RICHARD P. KORF. The publication date of Arend- holz's thesis on leaf-inhabiting Helotiales 187-190 BAKER, JOHN G., IRA F. SALKIN, DAVID H. PINCUS & RICHARD F. D'AMATO. Can- dida paratroptealis, a new species of Candida 115-119 BALAZY, STANISLAW, see MIETKIEWSKI & al. BEN-ZE'EV, ISRAEL, seé@ HUMBER §& BEN-ZE'EV BORELLI, DANTE, see McGINNIS §& BORELLI CANDOUSSAU, FRANCOISE, see PFISTER §& CANDOUSSAU CRANE, J. L., see HEWINGS & CRANE D'AMATO, RICHARD F., see BAKER & al. DUMONT, KENT P. Leotiaceae III. Notes on selected temperate species re- ferred to Helottum and Hymenoscyphus 59-84 GAMS, W. §& V. HOLUBOVA-JECHOVA. Chlortdtwn and some other dematiaceous hyphomycetes growing on decaying wood. Corrections and additions 257-258 GOCHENAUR, S. E. Cyrenella elegans gen. et sp. nov., a dikaryotic ana- morph 267-277 GRUFF, SUSAN C., see KORF & GRUFF GUZMAN, GASTON, see SCHROEDER §& GUZMAN HENNEBERT, G. L., Revue des Livres 278-286, 517-524 HENSSEN, A. & B. RENNER. Studies in the lichen genus Psoroma 1: Psoroma tenue and Psoroma etnnamomeum 433-449 HEWINGS, ADRIANNA D. & J. L. CRANE. The genus Codinaea. Three new species from the Americas 419-427 HJORTSTAM, KURT. Notes on Corticiaceae (Basidiomycetes). VIII. Two new species of Tubultecrtnts 120-123 HJORTSTAM, KURT. Notes on Corticiaceae (Basidiomycetes). IX. Three new combinations in Hypochnitctellum 124-126 HJORTSTAM, KURT §& LEIF RYVARDEN. Studies in tropical Corticiaceae (Basi- diomycetes) III. Two new species of Laxttextuwmn 35-40 HOLUBOVA-JECHOVA, V., see GAMS §& HOLUBOVA-JECHOVA HUMBER, RICHARD A. An alternative view of certain taxonomic criteria used in the Entomophthorales (Zygomycetes) 191-240 HUMBER, RICHARD A. Erynta (Zygomycetes: Entomophthorales): validations and new species 471-480 HUMBER, RICHARD A. & ISRAEL BEN-ZE'EV. Erynta (Zygomycetes: Entomophthor- ales): emendation, synonymy, and transfers 506-516 JAHNS, H. MARTIN. The genus PtZophorus 289-330 JENKINS, DAVID T. A new species of Amantta 112-114 KANE, JULIUS, IRA F. SALKIN, IRENE WEITZMAN & CATHERINE SMITKA. Trtcho- phyton raubitschekit, sp. nov. 259-266 . KOHN, LINDA M. A preliminary discomycete flora of Macaronesja: Part 3, Hyaloscyphaceae subf. Trichoscyphelloideae 145-149 KOHN, LINDA M. Sclerotinia bresadolae Rick, a taxonomic synonym of Ctbo- rtnta candolleana (Lév.) Whetzel 405-406 KORF, RICHARD P. Marcelle Le Gal, a reminiscence 1-4 KORF, RICHARD P. A preliminary discomycete flora of Macaronesia: Part 2, Hyaloscyphaceae subf. Arachnopezizoideae 137-144 KORF, RICHARD P. A preliminary discomycete flora of Macaronesia: Part 6, Geoglossaceae 361-366 KORF, RICHARD P. & SUSAN C. GRUFF. Discomycetes Exsiccati, fasc. IV 5-15 KORF, RICHARD P., see ARENDHOLZ & KORF LEGER, J. C. Les Hymenochaete a éléments hyméniens pinnatifides 241-256 LOWY, B. A new species of Daeryoptnax from Brazil 428-430 LUNGHINI, D., see ONOFRI & al. LUSTRATI, L., see ONOFRI & al. McGINNIS, MICHAEL R. §& DANTE BORELLI. Cladosporiwn banttanum and its synonym Cladosportum trtchotdes 127-136 MIETKIEWSKI, RYSZARD, RICHARD S. SOPER §& STANISLAW BALAZY. Notes on Zo- ophthora oeetdentalts (Thaxter) Batko (Entomophthorales: Entomophthor- aceae) 41-49 NAKASONE, K. K. Cultural studies on Porta etnerascens, P. rivulosa, and P. subvermtspora (aphyllophorales, Basidiomycotina) 105-111 OLIVER, LOLA K. & KEITH VAN CLEVE. Notes on soil fungi isolated from a 15-year-old aspen stand in interior Alaska 369-372 ONOFRI, S., D. LUNGHINI, A. RAMBELLI & L. LUSTRATI. New dematiaceous hy- phomycetes from tropical rainforest litter 331-338 PALM, MARY E. & ELWIN L. STEWART. Pithomyces pavgti, a new combination for Trichocladtum pavgtt and Pithomyces funtculosa 465-468 PFISTER, DONALD H. & FRANCOISE CANDOUSSAU. The psilopezioid fungi. VII. A new species of Pstlopezta from France 367-368 PFISTER, DONALD H. §& FRANCOISE CANDOUSSAU. The psilopezioid fungi. VIII. Additions to the genus Pachyella 457-464 PINCUS, DAVID H., see BAKER & al. RAMBELLI, A., see ONOFRI & al. REDHEAD, S. A. §& R. SINGER. Restnomycena gen. nov. (Agaricales), an ally of Hydropus, Mycena and Baeospora 150-170 REDHEAD, S. A. §& J. A. TRAQUAIR. Coprtnus sect. Herbicolae from Canada, notes on extralimital taxa, and the taxonomic position of a low tempera- ture basidiomycete forage crop pathogen from western Canada 373-404 RENNER, B., see HENSSEN §& RENNER RYVARDEN, LEIF. Type studies in the Polyporaceae 13. Species described by J. H. Léveillé 175-186 RYVARDEN, LEIF, see HJORTSTAM §& RYVARDEN SALKIN, IRA F., see BAKER & al., see KANE & al. SCHROEDER, ROBERT F. § GASTON GUZMAN. A psychotropic fungus in Nepal 346- 348 SIMMONS, EMORY G. Alternaria themes and variations 16-34 SIMMONS, EMORY G. dHalystomyces, a new dematiaceous genus from Arizona's Sonoran Desert 407-411 SINGER, R., Seé@ REDHEAD §& SINGER SMITKA, CATHERINE, see KANE @ al. SOPER, RICHARD S. New cicada pathogens: Massospora cteadettae from Au- stralia and Massospora pahartae from Afghanistan 50-58 SOPER, RICHARD S., see MIETKIEWSKI @ al. STEWART, ELWIN L., see PALM §& STEWART TAVARES, ISABELLE I. Validation of the Herpomycetineae and Herpomycetace- ae in the Laboulbeniales 469-470 TERADA, KATSUYUKI. Osortomyces, a new genus of the Laboulbeniales from Taiwan 412-418 TOYAZAKI, NORIHIRO. An undescribed pleomorphic species of Codinaea 450- 456 TRAQUAIR, J. A., see REDHEAD §& TRAQUAIR TUCKER, BRUCE E. A review of the nonentomogenous Entomophthorales 481-505 VAN CLEVE, KEITH, see OLIVER §& VAN CLEVE WEBER, WILLIAM A. Lichenes Exsiccati distributed by the University of Co- lorado Museum, Boulder: Fascicles 1-15, Nos. 1-600, 1961-1979 85-104 WEITZMAN, IRENE, see KANE & al. ZANG, MU. Sinotermttomyces, a new genus of Amanitaceae from Yunnan, China 171-174 528 INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME THIRTEEN This index includes genera, specific taxa. New Taxa are they are published. itself anh index, reference HEX Acarospora see 86-101 Acaulopage 276 Actinomyces 370 Agaricus decurrens 156; rhododendri 155, WSS Uiskikelkkeolitel sists}, silo Agrocybe tibetensis 347 Alectoria see 86-101 Aleuria annamitica 458 IMEC WO, iWsh AO, 245 Wh, 7S, 74). 150- 327 407 oe GiheehOSsthatac we noltinicantes 20; '‘gr' Erostratae, Solitariae 16; al- ternata 26, 29, 30; chlamydospora 16, iS, 24, 282 CaevySselaiiaeiml iO, ws, wes helianthi 16, 18, 19; limaciformis 16, 24, 25; longipes 16, 28-33; molesta 16, 17-19; mouchaccae 16, 18, 20, 21; ra- dicina 16; solani 28; tenuis 30; tenu- issima 16, 26, 29, 30; 27; zinniae 16 Ama miliice a ill Zemin tesSeGL 114; subsect Solitariae 112, 114; stirps Polypyramis 114; stirps Strobiliformis 114; stirps Virgineoides 114; marginata 112-114; muscaria 347 Amauroderma schomburgkii 181 Amyloathelia 124 Anaptychia see 86-101; multifida f circinalis Herwienine Wea, AS, Eepiidellia ws! 12; leucomelaena v 101; f verruci- fera 102; podocarpa v stellata 102 Ancylistes 201, 210, 481, 483-487, 489, 492, 493, 501, 502; closterii 482, 484- 486) 488") 490, 49Sn OOS ee netrite 401 482, 484-486, 488, 490, 493, 503; pfeif- feri 482, 484, 488, 490, 493, 503 Anthracobia macrocystis 7 Anthracothecium ochraceoflavum 86 Anzia see 86-101 Arachnopeziza 6, 137; '‘gr' Anomalae 14; ‘gr' Typicae 14; aranea f. aranea 138; f. monilipila 138, 139; aurata Uv, UG, WS eile 1S, Wests, Wekale Cangido=fUilVagOuss lS:au conmUtamsO alo. 14; leonina 6, 13; obtusipila 138, 141, 142; trabinelloides 6, 14; zonulata WSteh. RAID. aie Arachnoscypha 139; aranea 138 Arctomia fruticosa 98 Arthonia see 86-101 Arthopyrenia halodytes 87 Arthothelium see 86-101 Arthroderma benhamiae 260, 261; simii 260, 261 Aureobasidium 407 Bacidia see 86-101 Baeomyces see 86-101, 328; acicularis 298 Baeospora 9150505 151-95 168: 3 169 pallida LO loG infrageneric taxa, in boldface as are the pages where The Lichenes Exsiccati to which is species, and infra- paper (pp. 85-104) is indicated by the notation Ballocephala 200, 204, 210, 481, 483-489, 491-493, 498, 501, 502; pedicellata 482, 484, 485, 488,, 490, 493, 503; sphaerospora 482, 484-486, 488, 490, 493, 503; verrucospora 482, 484-486, 488, 490, 493, 498, 503 Basidiobolus 195, 198, 201, 204, 206-208, 210, 481, 487, 489, 501, 502; -micro= sporus 195, 208, 489 Belonia americana 87 Beltrania mangiferae 335; muelleri 335; onirica 331, 333-335, 337; querna 335 Bipolaris 339, 344; micropus 349; rave- nelii 340, 341, 345; tripogonis 344 Blastocladia pringsheimii 275 Botryobolus parasiticus 482 Boudiera dennisii 5, 7 Bryoria see 86-101 Buellia see 86-101; straminea 103 Calicium see 86-101 Caloplaca see 86-101 Caloplacopsis submexicana 88 Candelariella see 86-101 Candelina submexicana 88 Candida 115; paratropicalis 115-118; stell- latoidea 117; tropicalis 116, 117 Catenaria auxiliaris 483 Catillaria see 86-101 Cavernularia lophyrea 88 Cenomyce acicularis 298 Cetraria see 86-101 Cetrelia chicitae 88 Chaenotheca brunneola 88 Chaetopsina 333 Chaetosphaerella 257 Chaetosphaeria 450; callimorpha 426; dingleyae 425; fusiformis 257; fusi- spora 257; talbotii 426 Chiodecton see 86-101 Chloridium 257, 450; codinaeoides 450, 455 Chondropsis semiviridis 88 Chryseidea 331, 333; africana 331-333, SG Chrysosporium pannorum 371 Ciboria 79 Ciborinia 405, 406; candolleana 405, 406 Cladia see 86-101; galapagosensis 101; polia 101 Cladina galapagosensis 85, 89, 101; po- lia 85, 89, 101; subtenuis 89 Cladonia see 86-101; pileata 328 Cladosporium 127, 407; bantianum 127- 130 01325 eee 4 ee SORT Ghoncdesmen| 2a. W285 S2 ela hos Clavaria nigrita 362 Clavariana 276 Clavatospora 276 CoelhMmi@nelwis Ss SYA, sya GYAG) eit). 35050953 00-358 NDI Colom 5350: carbo— num 350, 351; cymbopogonis 344, 349, 35), sd, A, esses Cwineclomvls 350, 351; geniculatus 350, 351; hawai- iensis 344, 353, 356; theterostrophus 34953515, 352, 3545" homomorphus 349— 35 le intermedius 350, 351; kusanoi Syl. S285 Sse, See VWMMEUS SEO, Se)e miakei 350; nodulosus 349-351; palmi- vora 350; ravenelii 339, 341, 342, S44 S4 ODS GOO, ESatIIVUSEES49—35)I) S5S6 sitharamii S50; 351; spicifer 350, 351; sporoboli 342; tripogonis OS), oy), GNA sik, S58, seyh, sSe tiei= ie) Sis Wievoriae GO, sei, see Codinaea 419, 425, 426; apicalis 426; apiculata 425, 450; aristata 426; as- samica 426; botulispora 426; brevisetu- la 425; britannica 426, 450; clavulata 426, 451; cylindrospora 426, 450; di- morpha 451, 453-455; eucalypti 425, 450; fertilis 426, 455; glauco-nigra 426; gonytrichodes 426; hughesii 426, 450; illinoensis 419, 420, 425; longi- spora 426; lunata 426, 450; lIunulospo- ra 421, 422, 425; maharashtrensis 450; matsushimae 423-425; novae-guineensis 425, 450; obesispora 425; pakhalensis 426, 451; parva 426; septata 450; se- tosa 426; simplex 426; unisetula 425, 450; vulgaris 426 Coelocaulon see 86-101 Coelomomyces milkoi 232, 482, 483, 494 Coenogonium see 86-101 Collema see 86-101 Coltricia 183; sideroides 183 Completoria 200, 210, 481, 483-487, 490, 492, 494, 502; complens 482, 484-486, 488, 4905 9494, 5503 Conidiobolus 192, 196, 199, 201, 203- DOS meet =200 su 2iie ae 2i2, G21, Z2h7. 2219, 22 2 2 2G AEN re Oe, EO) ou aOe, AO Om CU OO Rm Ao 2pm Oo oO OP OOOO 2. adiaeritus 204; apiculatus 231; corona- iws AO, WSs imaler Als Ooscurus Aly, 22S MCC eS OAD Abuse > lneatinicOm— sorceSs 4B, BOO, Bie, Al, Bs 2A. Zo lpm Al, O12 13 Coniothyrium 371 Conotrema see 86-101 Coprinopsis friesii 376; phaeosporus 394 Coprinus:374, 386, 390, 401; sect Herbi- COlaeua3 73. 374, oor C90, 392, , 400, AmMoNiIwS B/S, SM, BO, s/s eirceMue= ss /4050 380. 98000, 3990;,1295, .990% au strofriesii 373, 385, 378,. 392, 394, 395; brassicae 373, 378, 386, 388, 39069 394.0 3972 DUrKIIg S15, 5925) ClInCho= Nensis, 3765, 38858392, (395; friesit 373, S75 ee OMNES (On msO LO) oC Os Ie, 393, 398, 400, 404; v microspora 398; herbivorus-— 375, 378, 388, 392, 395, 404: -herinkii 374, 376, 394, 397; kubi- ora Gree S73, SUA, Ss, Sele, ceil, Sear maysoidisporus 373, 374, 380, 381, 382, 384, 386, 387, 389, 390, 393, 396, 400-402; melo 373, 378, 388, 390, SE" s microspermus 398; neotropicus 373, '375, 390, 394, 395; paleotropicus $37 o 55 OD, 1090), 394. ©3952" pallidisporus 0985 phaeosporus 373; 375, 9376.) 386, Sh, SS, SS, S87, OO, 2O49 w sollitie= rius 398; picosporus 400; platypus 398; pseudofriesii 373, 376, 394, 396, S973) | psychromorbiduss 373, 95379, moro, 378, 382, 384, 386, 387, 390-393, 396, 400-402; pusio 400; rhombisporus 373, SUD, Sil 398, 404s Roecinesi Zoe ScileiMice 37/5, S73, SH Sse sweiliitzeyei= us 398; subtigrinellus 376, 380, 395, 3965) “subunticicolal 373, )13/8. 386.1590, S73 iieirinelius S76, S60, S85, cess triangulospora SfeVy40 urticicola SHS SH), S73, S82, BS, cei, S80, aA, 397, 399, 400; willapaensis 381; xan- INNCMS SHS, SHS, SA, Sa, sey Cora pavonia 90 Coriolopsis asper 179, 180; brunneo- leuca 181; caperata 179, 182; polyzona (Sie sancguinatiag | 7Chun 1S lof estrhu— mosa 185 Cornicularia see 86-101 Corticium cremeoisabellinum 125; subilla— queatum 126 Corynetes 363, 365; arenarius 365; atro- purpureus 364, 365; globosus 365; pur- purascens 365; robustus 365 Corynophoron colensoi 326 Crocicreas 69 Cullicicoia $86, 19, As, Bil, BAsiy 2s. 486 Curvularia 411 Cyclomyces 181; fuscus 181; setiporus 177; tabacinus 182 Cylindrotrichum 257; curvatum 258; el- lisii 257; gorii 258; helisciforme 258; oblongisporum 257; proliferum 257; tri- septatum 257; zignoéllae 258 Cypheliopsis bolanderi 90 Cyphelium inquinans 90 Cyrenella 268, 275, 276; elegans 267, 268-270, 272-276 Cystocoleus ebeneus 90 Cytospora 371 Dacryopinax 428, 430; dennisii 430; ele- gans 430; indacocheae 429, 430; mar- tinii 429, 430; maxidorii 428-430; spa- thularia 430; yungensis 430 Dactylina see 86-101 Daedalea 176; aulaxina 176; flavida 176; fuliginosa 176; lurida 176; micro- zona 176; plumbea 176; pruinosa 176; splendens 176; violacea 176 Darbishirella gracillima 90 Dasyscypha coerulescens v dealbata 148 Dasyscyphus 6, 14 Delacroixia 207 930 Dendrographa see 86-101 Dermatocarpon see 86-101 Desmazieria see 86-101 Dictyochaeta 451; fuegiana 451 Dictyonema see 86-101 Dictyopanus 177 Dimelaena see 86-101 Dimeromyces aberrans 470 Dimorphomyces 470 Diploicea canescens 90 Diploschistes see 86-101 Dirina see 86-101 Dirinaria subconfluens 96 Dolichocarpus chilensis 90 Drechslera 339, 344; tripogonis 342, 344 Embellisia 18, 24 Empusa (41; 229,;_, 513; 914; subg Triplo— sporium 219; aphidis 514; apiculata 7, Pile AKO calrolinieine we, Als. 227, 512; dipterigena 219; forficulae 512; fresenii 219, 227; geometralis 509; major 201, 211, 216; muscae 227; occidentalis 509; papillata 211, 216; thaxteriana 223, 224; Endocarpon see 86-101 Enterographa see 86-101 virescens 213 EntomopHhaga 192, 196, 199, 201, 209- 7a), alc. ASS. ASH), ES. | AS. §=— BOWE grylli 199, 221; obscura 224; thaxter- jana 224 Entomophthora 192, 195, 196, 198-200, 203- 20S ePZO9=2 0, e219 e220, 225, 220 eco, 231-234, 473, 483, 486, 487, 499, 501, 304, SiS, Bille eplyicis 2OA, BWA, AS. SOST OL, @ SI4ee eapieulata ) 204-ae225. 231; aquatica 213; batkoi 232; bulla- ta 473; canadensis 509; caroliniana SIZ ecarpentl er immcO4 see 205 seal ae acle. Zi Gee e2iiiam OlOs ms coleopterornummrclleam lide 218, 509, 510; creatonotus 474; crusto- Se 809s euilieis 198, ZOO, Bl@, eile curvispora 229, 230; delphacis 212; delpiniana 212; elateridiphaga 509; e- rupta 211; exitialis 511-513; ferrugi- nee SA, lds ieriiculles OY, Sis cle gantea 232; gloeospora 194;) grylli 228, 232; henrici 509, 510; ignobilis Z\\ 9 2S) 2 ch | © (pe kOe 2 ASS TAUISCES IS. Wet ZO, IO, Avil. 487, 501; nebriae 509, 510; obscura USE), 7, 7S, 27IND CreciceiMMalis ils ovispora 229, 230, 508; papillata 204, 225, 231; phytonomi 509; planchonia- met 70, APS [peKeli@eins 4s, 2742), 10%. 509; sphaerosperma 41, 47, 202, 210, 228, 507, 514; thaxteriana 199, 219, Jixs PA “ANeltovclithin ZBlile TWHslinaie yey, Gish, SIS), 6 AO), «= AIG weetrrinie@lle 481, 482, 486, 487, 489, 491, 499; vi- euler: A> Aol, Ail, AiGs Ae “Pil. 513; weberi 210; zabrii 511 Ephebe lanata 90 Erynia 192, 194, 196-198, 201-205, 208, ZN 213 ee zlO=2 One 2.0) Gee COZ ZO =a 5 Or AT, 472s aa GO 480. 499 O02 ee D06— 514; americana 473, 511; aphidis 509, 514; aquatica 213, 511; arrenoctona 203; blunkii 511; brahminae 511; bul- lata 471, 472-474, 479, 511; callipho- rae 511; canadensis 509; caroliniana IG, USS), ZAOZ=70, AG, 2S, Sls, SilAs castrans 512; coleopterorum 509; coni- ca 225, 511; crassitunicata 509; crea- tonoti 471, 474, 511; crustosa 509; curvispora Slits delphacis 212-215, 511; delpiniana 212-214, 511; dipteri- gena 511; echinospora 511; elateridi- phaga 509; erinacea 217, 511; formi- cae 471, 475, 476, 478, 479, 511; geo- metralis 509; gloeospora 511; gracilis 511; henrici 509, 510; ithacensis 212, 217; jaczewskii 509; lanceolata 509; magna 512; montana 511; myrmecopha- Gel CYAl, EMS. Uy, Mis. Sis meoriac 509; neoaphidis 42, 202, 214-216, 511, Byi[ dive mouryt =) 511, 513; occidentalis 509; orientalis 509; ovispora 507, 508, 510, 512, 514; phalangicida 512; phal— loides 509; phytonomi 509; radicans 509, 513, 514; rhizospora 512; sepul- chralis 512; variabilis 512; virescens 512; vomitoriae 512 Evernia prunastri 90 Everniastrum see 86-101 Exserohilum 339 Favolus 176; brasiliensis 176, 177; fi- brillosus 176; fissus 176; granulosus 176; guadeloupensis 176; junghuhnii 177; multiplex 177; peltatus 177; phi- lippinensis 176, 177; spatulatus 177; tener 177; tenuissimus 177 Fibuloporia subvermispora 109 Filobasidium 271 Fomes fasciatus 183 Fomitopsis 183; rhodophaeus 183; scutel- lata 183; supina 181, 183 Fulgensia desertorum 87 Fusarium roseum 370; semitectum 371 Galactinia 462; megalosperma 462; pseu- dosuccosa 461, 462; f macrospora 462 Ganoderma Ad, 182-184; applanatum 182, 184; lucidum 182-184 Geoglossum 361, 365; glutinosum 362; ni- gritum 362; v nigritum 362; umbratile v umbratile 362 Globifomes graveolens 178 Gloeocystidiellum 36; furfuraceum 39; propinquum 36; sibiricum 36 Gloeocystidium ltacticolor 36 Gloeoglossum 361, 362; glutinosum 362, 363 Gloeoporus 177; leptopilus 177; pusillus lia Glypholecia scabra 91 Graphis see 86-101 Gymnoderma see 86-101 Gyrostomum scyphuliferum 91 Haddowia 182 Haematomma see 86-101 Halysiomyces 407, 408; saxatilis 408 Haptoglossa 483 Helminthosporium 339 Helocarpon 290; crassipes 291 nelionivuun S85 CO, CA, @s, Ws GOR flllexo= punctatum 62; carpinacola 69; cauda- tum 62, 83; conocarpi 62; dearnessii 62; epiphyllum 66, 68, 74, 76; errati- Clin (9s, We, Gee ieuiclicswinn GE, 740). 72; fraternum 72; immutabile 68-69, 73, 74, 76, 83; linderi 76; midlanden- SO Wos55, GD, UWS, Hes 80; phyllophilum 79; translucens 80, 83 Heppia lutosa 91 Herpomyces 470 HERPOMYCETACEAE 469 Herpomyceteae 469 HERPOMYCETINEAE 469 Heterobasidion annosum 183 Heterodea muelleri 91 Heterodermia see 86-101; phyllogenon 79, rufo-corneum 59; barbifera 85, 101; circinalis 85, 101; stellata 85, 102; verrucifera 85, 86, 102 Heteromyces 328 Hexagonia 177; blumei 177; cingulata 177; dregeana 177; glabra 177; hirta 185; molkenboeri 177; pulchella 177; tabacina 177 Hubbsia lumbricoides 91 Humaria gregaria 5 Hydropus 150, 168, 169 Hydrothyria venosa 91 Hymenochaete 241, 254; acanthophysata MES). PAM). PI, PY, Ae, HSS Clininetino— mea 254; digitata 241, 246-249, 254, 256; harpago 241; hauerslevii 241, 253-256 ; pinnatifida 241-246, 254, 256; separabilis 241; spathulata 241 Hymenoscyphus 59, 60, 62, 65, 69, 72- (1 (9. SOO S ealiDopuinetatusso2.cau— Cais CO=54, G5, Wa, C8, 74%, Us, cereus 61; dearnessii 60, 62-64, epiphyllus 60, 65, 66; erraticus 61, 67, 69; fraternus 72; immutabilis 60, 65, 73, 74; lasiopodius 61; leucop- 61; musicola 61; phyllogenon 69; phyllophilus 79, 80; rufocorneus 80, sclerogenus 61; scutulus 61; serotinus 61, 72; translucens 61, 80, 82; umbili- catus 60 Hyphodiscosia europaea 455 Hyphodontia setulosa 111 Hypocenomyce friesii 92 Hypochniciellum 124, 125; cremeoisabelli-— num 124, 125; molle 124, 125; ovoide- um 124, 125; subillaqueatum 124-126 Hypogymnia see 86-101 Hypotrachyna bostrychodes 94 Icmadophila ericetorum 91 Ingaderia pulcherrima 91 Knightiella splachnirima 100 Koerberia biformis 91 Laccaria trullisata 267, 269 So Lachnellula 145, 148; pittospori ssp azo- rica 145, 146; ssp pittospori 146; pul- veracea 145, 147; viridi-glauca 145, 148 Lachnum 139; aranea 138 Lamia culicis 217 Lamprospora ovalispora 7 Lasallia see 86-101 Laxitextum = 535, G36" sbicolor SS ams6.9s9. 40; incrustatum 35, 37, 39; lutescens 39536, 938-40 Lecanactis see 86-101 Lecanidion 350 Lecanora see 86-101, 103; pinguis 102; pseudopinguis 85, 102, 103; sulphurea 104; texana 85, 103, 104 Lecidea see 86-101, 291; crassipes 291 Lecidella elaeochroma 93 Leciophysma finmarkicum 447; furfur— ascens 447 Lenzines vrs ecule WHS, I77, Ue, Webbe mel Melevi uve lesvuiliine: 177, vee ceilt= ata 178; elegans 178; guilleminiana 178; junghuhnii 178; juvenile 178; mu- rina 178; myriophylla 178; platyphyl- las ee platy podalsl(Seaestemtisms izes vespacea 176-178 Leprocaulon see 86-101 Leptogium see 86-101 Leptosphaeria 350 Leptosporomyces 124; ovoideus 124, 125 Letharia see 86-101 Leucogyrophana 124; cremeoisabellina 124; mollis 124; subillaqueata 124 Lichen cereolus 302 Lichina see 86-101 Lignosus sacer 183 Lobaria see 86-101 Lopadium pezizoideum- 94 Macrobiotophthora 481, 486, 498, 499, 501, 502; vermicola 484, 485, 488, 490, 499, 501-503; vimariensis 482, 484, 485, 487-490, 498, 503 Macrosporium longipes 28 Marasmius 150, 168; decurrens 152, 153, (56, eee ineasiinesws WSi0),, Yo4%, ies y CangdiiGissiimusml 2pm OsmE Vann VeUSmtoZ. 15350 IS6serhododencipia io Ipmlos Maronella laricina 94 Masonhalea richardsonii 88 Massospora 50, 51, 53-55, 198, 200, 205, 206, 210, 211, 227, 486, 492, 501, 502; cicadettae 50, 51-53, 56; cicadimal 51) 535 85am cZilteachticeroproc— tae 56; diminuta 55; dorisianae 53, Bas wielhiiintee Sse llewiSjevels SiO), Sih carinetae 55; 55; ocypetes 55; pahariae 50, 54-56; platypediae 56; spinosa 55; tettigatis 53855 Mastodia tessellata 94 Melanaria melanospora 94 Melanelia see 86-101 Menegazzia see 86-101 Meristacrum 192, 200, 204, 208, 210, 2320 481, 483, 485, 486, 494, Oe! [Meristacrum] 495, 498, 502; asterosper- mum 482-491, 494, 495,, 501, 503; milkoi 232, 484, 485, 488, 490, 494, 495, | 503; pendulatum, 481-483, 485— 487, 489, 491, 495 Micarea denigrata 94 Microglossum 361, 363, 365; atropurpure- um 364; olivaceum 363, 364 Microporus 182; affinis 182; scopulosus 182 Microthelia micula 94 Miyoshia fusispora 257 Miyoshiella fusispora 257 Moellerodiscus 62, 79 Monoicomyces leptochiri 416 Monosporium minutissimum 371 Mortierella alpina 370; gracilis 370; bellina 370; nana 370; vinacea 370 Mucor fragilis 370; varians 370 Muiogone 207 Mycena_ 150, 168, 150, 156; rorida 168 Mycoglaena myricae 94 isa- IGS: kalalochensis Nematophthora gymnophila 483 Neofuscelia see 86-101 Neophyllis melacarpa 94 Neozygites 196, 198, 208, 219, dis 219; turbinata 194 Nephroma see 86-101 Nephromopsis see 86-101 Neuropogon see 86-101 Nia 276 Nigroporus durus 183; vinosus 184 Normandina pulchella 94 228; aphi- Ocellularia alba 94 Ochrolechia see 86-101 Omphalaria kansana 94 Omphalia rhododendri 150, Omphalodium arizonicum 91 Omphalopsis rhododendri 152 Opegrapha saxicola 94 Osoriomyces 412, 416-418; UWB WA\Sy We AT Oxyporus cervino-gilvus 185 152 rhizophorus Pachyella 367, 457, 458, 463; adnata 458, 463; aquatilis 458; babingtonii 368, 458, 463; clypeata 458, 460, 463; hydrophila 457, 458, 463; megalosper- ma 459, 463; peltata 457, 459, 460, 463; pseudosuccosa 457-459, 461, 463; punctispora 457, 458, 461, 463; viola- ceonigra 462, 463 Panellus 177; pusillus 176 Pannaria see 86-101, 434, 445; pezizoi- des 434; rubiginosa 445 Paracnhnopeziza miniopsis 14 Parathelium see 86-101 Parmelia see 86-101 Parmelina galbina 95 Parmeliopsis see 86-101 Parmotrema see 86-101 Peccania see 86-101 Peltigera see 86-101 Peltula see 86-101 Penicillium 370, 371; asperum 371; chry- sogenum 370; coryophilum 371; freque- tans 370; funiculosum 370; kKapuscin-— skii 371; pinetorum 370; restrictum 370; solitum 370; soppi 370; thomii 370 Pertusaria see 86-101 RPezizal 45/7, 4585) ~aquati lism 457.5) 460i, 462, 464; atroviolacea 368; candido- fulva 13; caudata 62; epiphylla 66, 74, 76; exidiiformis 458; leonina 13; phyllophila 79, 80; violaceonigra 462 Pezoloma laricina 14 Phaeographina see 86-101 Phaeographis see 86-101 Phaeotrichoconis 338; 338; crotalariae 338; urariae 338 Phellinus 178-180; appositus 178; calli- morphus 179; chryseus 179; extensus 180; fastuosus 180; gilvus 179, 184; pectinatus 181; senex 178, 181 Phialea dearnessii 62, 65 Phlebia subochracea 111; Phylliscum see 86-101 Phylloporia chrysita 184 Physcia see 86-101; barbifera 101 Physconia see 86-101 Physma byrsinum 97 Pilophoron 290; sect Eupilophoron 316, 317; sect Nigricaulia 316, 317; cereo- lus 315; polycarpum 317; robustum 97 Pilophorus 289-293, 295, 316, 317, 326, 328; sect Eupilophorus 291; sect Nigri- Cauilew 29S acictilainiisu 292.2 o Sc Or SO), S05, S07, 409), SI@, s/, S20, Se5s v conjugens 299; f hallii 308; awas-— thianum 292-298, 300-302, 316; cario- SUM S495 CGareoliUis S10, 7274, 7885, 2S. 297, 302-307, 31355320," 324; Vo cephalo— diferus 305; v_ hallii 308; clavatus 292-298, 308-310, 316, 317; colensoi 295, 326; v reagens 326; conglomera- JU Zk, C86, 329, seis euriullum. 2O2Z; M28), Vise), PSO, COA, SNO=SUF, SMA, Siles Cistems ss Wilowile 287, 288, 286) PNG, 7A}. SIVA, S07, SUES. S28 inal= fii 308-310; japonicum 308, 310; nigri- caulle 2912297. 30419 9315-3170 325, 3205 pileatum 328; robustus 290, 292-294, 296-3005 9306-8 1317-32 Ihe 328-9252 Can distans 321; f magnus 320; staufferi 292,295, 93283) strumaticus 292-2947 296-298, 304-307, 313, 316; 320-324: vegae 292-298, 300, 304, 316, 324-326 Pithomyces 465; funiculosa 465; pavgii 465-468 Placynthium nigrum 97 Platismatia see 86-101 Plectania sect Plicosporae 6, 8; oides 6, 8 Pleospora infectoria 26 Poculum 73 Polychidium muscicola 97 punctispora 461; aurata 331, 335- subserialis 111 cyttari- Polydesmia 143; fructicola 143; pruinosa 143, 144 Polyporus 178; abnormis 178; aculeatus 178; albomarginatus 178; anisopilus 178; apalus 178; appositus 178; aty- pus 178; auriculaeformis 178; blumei Wee bonplandensis 73) botryoides (Sse DRacy pUSmalicomecal limonpinusml) 29) callochrous 179; candicans 179; candi- dulus 179; chryseus 179; cineraceus 179; cinerascens 179; cohaerens 179; confertus 179; connexus 179; convolu- tUS sl 9e ecOomiaceus a 79-5 corprugatus 179; cyathiformis 179; demidoffii 179; dermatodes 180; dilitatus 180; discifor- mis 180; dissectus 180; dozyanus 180; elatus 180; extensus 180; fastuosus 180; flabelliformis 182; fuscellus 180; fuscus 180; gaudichaudii 181; gaya- nus 181; gibberulosus 181; gossypinus ined] guadeloupensis Sits haskarlii 181; hasseltii 181; heteromorphus 181; hymenius 181; inquinatus 181; kickxi- anus 181; Korthalsii 181; lenis 181; lenziteus 181; lteucomelas 181; lindi- gii 181;longipes 182; macropus 182; mangiferae 182; manubriatus 182; mar- chionicus 182; mastoporus 182; mega- loma 182; melanaleucus 182; melaneus Iss finikelreevcelluis: iteV45 (iiliteleralketine: — iitsvae moritzianus 182; murinus 182; nephel- odes 182, 183; nordmannii 183; noto- pus 183; ostreatus 183; pala 183; pec- tunculus 183; perpusillus 183; phaeus 183; placopus 183; platypilus 183; plumbeus 183; rhodophaeus 183; rigi- dus 183; rudis 183; rugulosus 183; sanguinaria 178; sclerodermus’7 183; scleropodius 183; sericellus 183; seti- porus 180; sideroides 183, 184; sordi- dus 184; splendens 184; spurcus 184; stevenii 184; subflavus 184; swartzi- anus 184; tegularis 184; tenax 184; tenuissimus 184; testaceus 184; tracho- des 184; tricolor 184; trigonis 184; tristis 184; unguiformis 184; vulnera- tus 184; zollingerianus 184 Poria 107, 111; albipellucida 105, 108; cinerascens 105-108, 110; lindbladii 108; quercuum 109; rivulosa 105, 107- 110; subvermispora 105, 107-110 Porina see 86-101 Protoblastenia russula 92 Psathyra urticicola 388 Pseudephebe pubescens 86 Pseudocochliobolus 350, 357 Pseudocoelomomyces milkoi 494 Pseudocyphellaria see 86-101 Pseudoparmelia see 86-101 Pseudoplectania nigrella 8 Psilocybe 346, 347; cubensis 346; subcu- bensis 346 Psilopezia 367; mummularia 367, 368; nummularialis 367, 368, 459 Psora see 86-101; cerebriformis 85, 104 Doo Psoroma see 86-101, 433, 434, 439, 44], 445; bryantii 433-435, 448; cinnamome- um 433, 435, 439, 443, 445-448; foll- mannii 433, 434, 436, 448; hirsutulum 435; hypnorum 433-435, 445, 447, 448; paleaceum 435, 436, 447; rubromargi- natum 435; tenue 433-436, 437, 446, 448; v borealis 433, 434, 439, 441, 443, 445, 447, 448; v tenue 433, 436, 437, 439, 441, 443, 445, 447, 448 Psorotichia see 86-101 Psorula rufonigra 93 Ptychoverpa bohemica 8 Pulvinula globifera 9; niveo-alba 9 Pyrenophora 339, 350 Pyrenotrichum splitgerberi 97 ’ Pyrenula see 86-101 Pyrofomes 178, 179; albomarginatus 178; demidoffii 179 Pyxine pringlei 97 Ramalina see 86-101 Ramalodium succulentum 98 Reinkella see 86-101 Resinomycena 150, 151, 168, 169; acadi- GmsiS I5, Wai, Wl, WG2, We7s levine mescens 150, 151, 163, 165, 168; kala- Iannis ISA, WES, ley, IS, el, JO45 fuleliiete: WSO, We, W774, IE@=aiG7, l67eeirhododendriy iS 152. n 5S o>— lav, Sil, 162, ies Rhizocarpon macrosporum 98 Rhizoplaca see 86-101 Rhodosporidium 273; Rhodotorula 267, dacryoidum 273 275, 276; aurantiaca DZ See (CO Malactosa2 co Rigidoporus fusco-lineatus 183; micropo- rus 184 Rinodina see 86-101 Roccella see 86-101 Roccellaria see 86-101 Roccellina see 86-101 Ruhlandiella berolinensis 5, 9 Saccharomyces cerevisiae 275 Sarcogyne see 86-101 Sarcoleotia 365; globosa 6, 15 Sarcosoma 6; cyttarioides 5, 8 Scelophoromyces 412, 418; osorianus 412 Schismatomma cupressum 99 Schistophoron tenue 99 Schizopelte californica 99 Sclerotinia 405; bresadolae 405, 406; candolleana 405 Scutellinia 2, 10; erinaceus 10; pennsy!- vanica 10; scutellata 6, 10; Uumbro- rum 11 Scytinostromella 39; humifaciens 39 Septobasidium 207 Sepultaria 11; gregaria 6, 11 Setosphaeria 339, 350 Sinotermitomyces 171, 173; carnosus 172, USA CEBU TWA, WES Siphula see 86-101 Sistotrema 185; ochroleucum 185 534 Skelophoromyces 418 Speerschneidera euploca 99 Sphaeronaema spinella 370 Sphaerophorus see 86-101 Sporastatia testudinea 99 Sporormia 350 Squamarina lentigera: 92 Stachybotrys atra 370 Staurothele clopima 99 Stemphylium 411 Stereocaulon see 86-101, 289, 292, 307, 326; colensoi 326; fibula 312, 313; pi- leatum 307 Stereum australe 254 Sticta see 86-101 Stigmatomyces baeri 417 Stomiopeltis 350 Streptomyces 370 Striatosphaeria codinaeophora 451 Strigula elegans 99 Strongwellsea 192, 194, 196, 197, 202- 2, POS, Dl, Bit3, 222, db, ZO, 50254 006, 95061 5125 Castranse 202088216. Sis tiene WA, ZO, ils, BAS, Se Tabanomyces 192, 200; 204, 208, 210) oY. (Nee Hels ii likol 237, 48), Bes. 487, 491, 494 VaiieoiUin Ate ZIE2 zewskii 509, 511 Teloschistes see 86-101 Termitariopsis 207 Hetamiltomyceses | ClewmlkcZ Tnamnolia see 86-101 Thelephora mollis 125 Thelidea see 86-101 Theloma see 86-101 Thelotrema diminitum 100 Thrombium epigaeum 100 Thuemenidium 361, 363-365; um 364, 365 Thysanothecium 328; hookeri 100 Toninia see 86-101 Torula bantiana 127, 128, 130, 132 auxiliare 483; jac- atropurpure- 5 ES Trametes 185; acuta 185; cervina 179; crassa 185; cubensis 183; dermatodes 180, 185; hirsuta 179; incana- 185; ma- rianna 179, 181, 182; membranaceus 178; menziesii 178-182; modesta 178, 182-184; perrottetii 185; scabrosa 184; trichomallus 185; versicolor 183, 184; villosa 181, 184; vittata 186 Trapeliopsis wallrothii 93 Trichaptum 185; biformis 179, 181; bys- sogenus 184; perrottetii 175 Trichocladium 465; pavgii 465 Trichoderma viride 370 Trichoglossum 361, 366; hirsutum v_ hir- sutum 366 Trichophaea gregaria v intermedia 6, 11; f gregaria 5, 6, 11; f laevispora Be, 1 Trichophyton 259; mentagrophytes 259, 261, 265; raubitschekii 259, 260-262, 204 ZOD eu eU iin 259 a2 Ollie O49meZ OO Triplosporium 196, 198, 200, 208, 210 pein, (tH BikS.. ESP Trypethelium see 86-101 Tubulicrinis 120; angustus 123; cinctoi- ols A, Ale cliieimc 149, ile Gor ike) WAGs clidewlleswic 170, i177 25s hamatus 123; inornatus 123; ovalispo- PUS 12405 145 128 Tyromyces. 180; caesius 181; 175, 180; floriformis 180 o) dissectus Ulocladium 18 18 Umbilicaria see 86-101 Underwoodia beatonii 6, 12 Urnula craterium 12 Usnea see 86-101 , 20, 24; chlamydosporum Verrucaria laevata 101 Wynnea americana 12 Xanthoparmelia see 86-101 Xanthopsora texana 93 Xanthoria see 86-101 Xylographa see 86-101 /Agoyainiinera Wil, 4, IS, Wee, We. 197’. AAV =A08,. AVS, Olle 7A, All, BilZ, Bue. Js), PISA, (MA BOSS), SUG, Sil subg Erynia 213, 214; subg Zoophtho- ra 196, 208, 510; aphidis 514; bulla- ta 473; crassitunicata 201, 202, 212, 509; creatonoti 474; erinacea 213; exi- tialis 42, 512; ferruginea 512; forficu— lae 512; jaczewskii 510; lanceolata 212, 509; myrmecophaga 476, 477; occi- dentalis 41-47; orientalis 212, 509; oxsweimin AAilPas Yoyayauliieyitetess, CA, Yay (AP). S025 PACKERS 2/, AWA, ZI@, 22, 22). 506 Zygnemomyces 481, 486, 489, 492, 495, 498, 501, 502; echinulatus 482, 484- 490, 495, 496, 498, 503; pendulatus 484, 485, 488, 490, 495, 497, 498, 503 Zygorhynchus heterogamus 371 PUBLICATION DATES FOR MYCOTAXON Volume “12, "No. Z Volume 13, No. 1 Volume 13, No. 2 March= 17," 1981 May 11, 1981 USUPZ er AEN REVIEWERS The Co-Editors express their appreciation to the following individu- als who have reviewed one or more of the papers in this volume prior to acceptance for publication ee (Wedge) Rien OLEBERTSON T= Ae MOORE J. AMMIRATI Je. GINNS G. MORGAN-JONES R. J. BANDON! Ino Aig LEGO T. NASH Pee a OA TRA Joe heetiA LUNES DUE eGEER R. K. BENJAMIN A. HENSSEN Dz W.) ROBERTS eae GEN 7 Bey R. A. HUMBER Ora em ROGERS Hamed ee UROSALL SSRs J. W. KIMBROUGH M. A. SHERWOOD J. W. CARMICHAEL R. F. LANGDON Ro rey SLOP Sin Jee RAME FR G. A. LAURSEN W. J. SUNDBERG J. DREW-SMITH F. F. LOMBARD [Peel aA ARES F.-E. ECKBLAD D. M. MacLEOD J. Wo THOMSON J. ERILKSSON D. MALLOCH O. VERONA S. FRANCIS Dan Ee MiCC ADE R. WATLING W. GAMS M. R. McGINNIS Ne Mates ANE DIET ERRATA, VOLUME TEN Page 471, line 12: for 1094: La read 1094: La Palma. ERRATA, VOLUME TWELVE Page 280, line 8: for Rossman, J. Wy- read Rossman, L. J. Spielman, J. Wy- 536 Page 2; 182 203 216 217, 46] line Leg. 14, MA. ERRATA, VOLUME THIRTEEN > for gal read Gal : for (Remaudiére) Batko, read Remaud. & Henneb., : for form read from : for M. etcadina read Massospora cteadtna : for tmnttabults read tmmutabtlts : for tmmuttbtle read tmmutabtlis : for H. cauda- read Hymenoscyphus cauda- : for Hymenoscyphus fastidtosus read Helotium fasttdtosum : for sporen read spore tetnseererg. PEMLOCALL LY mabrd cade : for 4 read 5 7 tOr Poephi lipinens Usereadur aunt iippinensis : for brasilensis read brasiliensis “yor Po phil bipinens ts sready? philippinensis : for brasilensis read brasiliensis : for berkleyi read berkeleyi : for Phillipines read Philippines : for tenius read tenuis : for tabcinus read tabacinus : for supinus read supina : for Phillipines read Philippines : for Arendholtz read Arendholz : delete nuclei of : for EZ. read Entomophthora : for Z. read Erynta : for FE. caolintana and E. carpenttert read Erynta carolintana and Entomophthora carpentiert EO LOr ‘for :V for Tor - for 27for SEeOr for 27 for eo efor etTor 2 for SD idaye LOT POT. Sh op i for E. read Empusa E. read Entomophthora E. read Entomophthora E. read Entomophthora segregated read segregates E. read Entomophthora E. read Entomophthora C. read Contdtobolus Maraconesian read Macaronesian Fusareum read Fusartum Steptomyces read Streptomyces aspernum read asperum dengleyae read dingleyae P. read Psoroma P, read Psoroma C. read Codtnaea parkhalen- read pakhalen- C. read Codtnaea aquttilis read aquatilts , , ‘ i ? f lig bees a) 7 P » © i ie oan 7 in 7 Bs - ' 5 sad ye qc i of - - : Ac : 1 7 ie es. 4 ety , 7 on) SNe ae a a) j i my) p Liha ; ; p t :" ive «@ , we yr i et i, ‘ Ue - ' ’ ' + Cir ene | at ab Ta My ia i . ; 4 5 ' 4 ; i { ’ i ' F , : : : i iy | ti i i ' " 4 + 4 “4 ' : é 3. 1 ' = ae m ; : | Sf r, ' ' i a ¢ ; : ' = t i oy ( ts mT | =, ' , is th ' ' - = , 7 ' i =f = a) ! 4 ¥ ‘4 ‘ ! ' ? 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