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NO.2OCTUBRE2001 LANKESTERIANA LA REVISTA CIENTIFICA DEL JARDIN BOTANICO LANKESTER UNIVERSIDAD DE COSTA RICA Dora Emilia Mora de Retana, 1940-2001 CARLOSO.MORALES1 Dora Emilia Mora de Retana, a remembrance JOHNT.ATWOOD9 Stellilabium erratum , a comedy of blunders ROBERTL.DRESSLER11 Myoxanthus vittatus (Orchidaceae FRANCOPUPULINandMARIOA.BLANCO15 The genus Christensonia ERICA. CHRISTENSON19 Endophytes of Serapias parviflora Parlatore and Spiranthes spiralis (L.Orchidaceae description of endophytes of S. parviflora , and in vitro symbiosis development in S. parviflora and Spiranthes spiralis PIERLUIGIPACETTIandSABINERIESS23 ISSN 1409-3871 

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LANKESTERIANA LAREVISTACIENTIFICADELJARDINBOT ANICOLANKESTERUNIVERSIDADDECOSTARICA Editors CARLOSO. MORALESFRANCOPUPULIN Editorial Board MARTAALEIDADIAZERICHAGSATERJOHNT. ATWOODDORAEMILIAMORA-RETANAROBERTL. DRESSLERPHILIPSEATONJOAQUINGARCIA-CASTROJORGEWARNER Scientific Board J. ACKERMANNM.GRAYUMD.H. BENZINGB.E. HAMMELG. CARNEVALID.H. JANZENM.CHASEM. LOWMANT. CROATH. LUTHERP. CRIBBJ. MONGE-NAJERAG.GERLACHA. PRIDGEONL.D. GOMEZG. ROMEROJ. GOMEZ-LAURITON.H. WILLIAMS LANKESTERIANA is published at irregular intervals at prices listed below by the Jard’n Botnico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. POSTMASTER: Send address changes to Jard’n Botnico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. EDITORIAL OFFICE: Jard’n Botnico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. MANUSCRIPTS: Send to Editorial Office INFORMATION FORCONTRIBUTORs: Send request to Editorial Office. MEMBERSHIP OFFICE: Jard’n Botnico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. SUBSCRIPTIONRATEs: $ 4.00 per issue CR, $ 6.00 elsewhere. SUBSCRIPTIONTERM: Calendar year only, includind three copies.Differences in issue number will be compensated with next year subscription. REMITTANCES: All checks and money orders must be payable through a Costa Rica bank in U.S. dollars or Colones. Pay toJard’n Botnico Lankester, Fundevi. BACK ISSUES: Single issues are availbale for sale, single copies at $ 6.00 (CRelsewhere inquiries to Jard’n Botnico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. CHANGES OF ADDRESS: Send mailing label or complete old address and new address to Membership Office. CLAIMS: Copies lost in mails must be claimed at the Membership Office within 90 days from issue date. After 90 days, the single-issue price will be charged for replacement. REPRINTS: No free copies supplied. Price quotations and order blanks will be sent with galley proofs. Send orders, inquiries, and payment for reprints to Membership Office, Jard’n Botnico Lankester, Universidad deCosta Rica, P.O. Box 1031-7050 Cartago, Costa Rica, A.C. 

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INFORMACION PARA LOS CONTRIBUDORES LANKESTERIANA , la revista cient’fica del Jard’n Botnico Lankester, Universidad de Costa Rica, se dedica a la publicaci—n de todo tipo de art’culos de botni ca, con especial atenci—n a las ep’fitas y a la sistemtica, la ecolog’a, le evoluci—n y la fisiolog’a de las orqu’deas, as’ como a revisiones de libros y conferencias en estos temas. Se aceptan, adems, comunica-ciones breves y comentarios, que sern titulados como tales. La revista publica art’culos en espa–ol con resumen en ingls o en ingls con un extenso resumen en espa–ol. Los manuscritos sern evaluados cr’tica-mente por dos o ms revisores. Los autores son invita dos pero no obligados a cooperar econ—micamente para la reducci—n de los costos de impresi—n de sus art’culos. Los manuscritos deben inviarse a: Editores de Lankesteriana, Jard’n Botnico Lankester, Universidadde Costa Rica, P.O. Box 1031-7050 Cartago, CostaRica, A.C. Los textos deben ser todos a doble espacio, incluyendo nombre del autor, t’tulo, resumen, agradec-imientos, cuadros, leyendas de ilustraciones y bibli ografia citada. Las pginas debern ser numeradas con secutivamente, empezando con la pgina del t’tulo y terminando con la ltima pgina de las leyendas de ilustraciones. Los cuadros se imprimirn en hojas sepa-radas. Cada cuadro y figura deber tener referencia enel texto. Las leyendas cada ilustraci—n se imprimirn enpginas separadas. Los herbarios se citarn segn el Index Herbariorum , las publicaciones en serie se abreviarn segn el Botanico-Periodicum-Huntianum ( BPH ) y lo autores de nombres cient’ficos segn el Draft Index of Author Abbreviations . Las ilustraciones originales debern montarse en cartulina gruesa. Lasfiguras estarn numeradas consecutivamente de acuer do a su aparici—n en el texto. Enviar el original y dos copias a doble espacio. No enviar las ilustraciones originales hasta que se haya terminado el proceso de revisi—n editorial y cient’fica. El material que no se conforme al estilo indicado arribaser devuelto para modificaciones antes de su env’o a los revisores. Despus del proceso de revisi—n, deber enviarse tambin una copia electr—nica del manuscritoen un disquete, procesado en Word u otro programa compatible con Word, sea para sistema PC o Macintosh. INFORMATION FOR CONTRIBUTORS LANKESTERIANA , the Scientific Journal of Jard’n Botnico Lankester Universidad de Costa Rica, isdevoted to the publication of any article on botany, with a special emphasis on epiphytic botany and orchid systematics, ecology, evolution, physiology, as well asreviews of books and conferences on these topics. Short communications and commentaries are also welcomed, and should be titled as such. The journal publishes papers in Spanish language with an English summary or in English with Spanish extensive summary. Manuscripts wil be evaluated critically by two or more referees. Authors are encouraged but not required tohelp defray the costs of printing their papers. Submitted manuscripts should be sent to: Lankesteriana Editors, Jard’n Botnico Lankester, Universidad deCosta Rica, P.O. Box 1031-7050 Cartago, Costa Rica,A.C. Text should be doubled-spaced throughout,including author/s name, title, abstract, acknowledg ments, tables, figure captions, and Literature Cited. Pages should be numbered consecutively, beginningwith the title page and ending with the last page of fig ure captions. Tables should be typed on separate sheets. Every table and figure must be referenced in the text.Figure captions for each plate should be typed on sepa rate pages. Nomenclature material should follow Index Herbariorum for designation of herbaria, BotanicoPeriodicum-Huntianum ( BPH ) for abbreviations of ser ial publications, and the Draft Index of Author Abbreviations for authors of scienific names. Original of illustrations should be mounted on heavy cardboard.Figures must be numbered consecutively according totheir appearence in the text. Submit double-spaced manuscripts the original and two copies for reviewers. Retain original illustrationsuntil after the review process should any changes berecommended. Submissions not adhering to the styleabove will be returned for modification before being sent out for review. After the review process an electronic copy of the manuscript must be submitted on diskette, processed on Word or Word compatible programs, both for PC or Macintosh systems. 

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E STE FASCICULO DE L ANKESTERIANA SE DEDICA A LA MEMORIA DE D ORA E MILIA M ORA DE R ETANA EMINENTE BOTANICA E INVESTIGADORA POR MUCHOS AOS D IRECTORA DEL J ARDIN B OTANICO L ANKESTER EN RECONOCIMIENTO POR PARTE DE TODOS LOS COLEGAS DE LA GRAN HERENCIA CIENTIFICA QUE NOS HA DEJADO 

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N O . 2 O CTUBRE 2001 L A N K E S T E R I A N A ISSN 1409-3871 Dora Emilia Mora de Retana, 1940-2001 C ARLOS O. M ORALES 1 Dora Emilia Mora de Retana, a remembrance J OHN T. A TWOOD 9 Stellilabium erratum , a comedy of blunders R OBERT L. D RESSLER 11 Myoxanthus vittatus (Orchidaceae), a new species from Costa Rica F RANCO P UPULIN and M ARIO A. B LANCO 15 The genus Christensonia E RIC A. C HRISTENSON 19 Endophytes of Serapias parviflora Parl. and Spiranthes spiralis (L.) Chevall. (Orchidaceae): description of endophytes of S. parviflora , and in vitro symbiosis development in S. parviflora and Spiranthes spiralis P IER L UIGI P ACETTI and S ABINE R IESS 23 LA REVISTA CIENTFICA DEL JARDN BOTNICO LANKESTER UNIVERSIDAD DE COSTA RICA

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L A N K E S T E R I A N A LA REVISTA CIENTIFICA DEL JARDIN BOTANICO LANKESTER UNIVERSIDAD DE COSTA RICA Copyright 2001 Jard’n Botnico Lankester, Universidad de Costa Rica Fecha de publicaci—n vlida / Effective publication date: 29 de octubre del 2001 Diagramaci—n: Jard’n Botnico Lankester Digitalizaci—n de las imgenes: Carlos Ossenbach y Asociados Imprenta: Litograf’a Ediciones Sanabria S.A. Tiraje: 500 copias Impreso en Costa Rica / Printed in Costa Rica Lankesteriana / La revista cient’fica del Jard’n Botnico Lankester, Universidad de Costa Rica. N o . 1 (2001). San Jos, Costa Rica, Editorial Universidad de Costa Rica, 2001-v. ISSN 1409-3871 1.Botnica Publicaciones peri—dicas, 2. Publicaciones peri—dicas costarricenses R 

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D ORA E MILIA M ORA DE R ETANA (1940-2001)

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El 12 de julio de 2001 muri— Dora E. Mora de Retana, quien desde 1979 hasta febrero de 2000 fue directora del Jard’n Botnico Lankester, de la Universidad de Costa Rica (U.C.R.). Ella dedic— la mayor parte de su energ’a vital a estudiar las orqu’deas y a promover la conservaci—n de la flora tropical; esto ltimo desde su nombramiento como autoridad cient’fica en el Comit de Flora de la Convenci—n Internacional sobre Comercio de Especies Amenazadas (CITES). Despus de Rafael Lucas Rodr’guez (1915-1981), ella fue durante mucho tiempo la nica orquide—loga costarricense con formaci—n acadmica y cient’fica en el campo de la botnica. Dora Emilia naci— el 24 de agosto de 1940 en Cartago. Realiz— los estudios primarios en la Escuela Sagrado Coraz—n de Jess, los secundarios en el colegio hom—nimo. En la Escuela de Biolog’a (U.C.R.) obtuvo los t’tulos de Profesora en Ciencias, Bachiller en Biolog’a y Licenciada en Biolog’a. En 1969 empez— a laborar en la Ctedra de Fundamentos de Biolog’a, Escuela de Estudios Generales (U.C.R.). En 1973 inici— labores en la Ctedra de Biolog’a General de la Escuela de Biolog’a. En esta misma unidad de la U.C.R. empez— en 1978 con la Ctedra de Botnica General y a partir de 1984 ofreci— el curso de Orquideolog’a, entre otros. Los primeros estudios publicados por Dora Emilia (ver adelante) se refer’an a la anatom’a y al desarrollo de una especie muy conocida por los orquide—logos y aficionados a las orqu’deas en Costa Rica: A c n i s t u s arborescens (L.) Schltdl. (Solanaceae), un arbusto o rbol peque–o llamado gitite, a cuya corteza suave y corchosa se adhieren fcilmente las ra’ces de las orqu’deas ep’fitas. Con la direcci—n de Dora Emilia, lenta pero pro gresivamente el Jard’n Botnico Lankester dej— de ser una peque–a finca de 10,7 hectreas para con vertirse en un jard’n botnico de renombre mundial, especialmente por su extraordinaria colecci—n de orqu’deas. Gran parte del desarrollo y del progreso del Jard’n se deben al esfuerzo y a la energ’a de Dora Emilia y al empe–o de los buenos trabajadores que siempre ha tenido el Jard’n. La capaci dad de trabajo de Dora Emilia muchas veces me sorprendi—: pod’a empezar su d’a de trabajo a las 7 horas y estar activa sin descanso hasta las 20 horas, todav’a en los ltimos a–os cuando ya pod’a haberse pensionado y retirado. Sus actividades eran tan verstiles, que en un solo d’a pod’a visitar el Jard’n Lankester, en Cartago, para coordinar labores prioritarias, ir a San Jos a dar clases de botnica u orquideolog’a, viajar al Ministerio de Recursos Naturales (ahora, Ministerio del Ambiente) a atender algn asunto propio de CITES, ir a tocar las puertas de las autoridades uni versitarias para exigirles apoyo para el Jard’n Lankester y, no pocas veces, al final del d’a, visitar a Joaqu’n Garc’a 1 para identificar especies y, jun tos, preparar publicaciones sobre orqu’deas. Sin embargo, una vez Dora Emilia me revel— que se cansaba mucho trabajando en herbarios; fue durante una de las visitas de John T. Atwood (SEL) al Herbario de la Universidad de Costa Rica (USJ) y al Herbario Nacional de Costa Rica (CR). “l, en cambio, es incansable”, me dijo Dora Emilia en CR se–alando a Atwood, quien estaba de pie y muy concentrado revisando ejemplares de M a x i l l a r i a . Es obvio que Dora Emilia prefer’a estudiar sus queridas orqu’deas in vivo. Muchas personas recuerdan a Dora Emilia como una persona accesible, comunicativa, enrgica, dis LANKESTERIANA 2: 1-7. 2001. DORA EMILIA MORA DE RETANA, 1940-2001 C ARLOS O. M ORALES Jard’n Botnico Lankester, Apdo. 1031-7050 Cartago, Costa Rica, y Escuela de Biolog’a, Universidad de Costa Rica. 2060 San Jos, Costa Rica 1 El Dr. Joaqu’n B. Garc’a falleci— el 22 de agosto del 2001 y sus cenizas yacen desde el 2 de septiembre en el Jard’n Botnico Lankester. En dos meses consecutivos, Costa Rica perdi— a sus dos mejores orquide—logos nativos. 

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LANKESTERIANA puesta siempre a ayudar y a compartir ideas y conocimientos. As’ la recuerdo yo desde que fui su asistente en el Jard’n Botnico Lankester y cuando, por no haber otros estudiantes interesados, tom su curso de Orquideolog’a por tutor’a. De este modo el trato era mucho ms personal que en un grupo de estudiantes. Yo la segu’a por el Jard’n Lankester, anotando en mi cuaderno las caracter’sticas de gneros y especies que ella me mostraba. Recuerdo muy bien que jams perdi— la paciencia conmigo; todo lo contrario: parec’a muy satisfecha de que alguien ms se interesara por las orqu’deas y era evidente que le gustaba transmitir sus conocimien tos sobre estas plantas. Yo s muy bien que otros no piensan igual. El carcter generalmente sobrio y apacible de Dora Emilia se volv’a frreo e irreductible al defender sus ideas y sus acciones frente a las de otros; una virtud que muchos costarricenses no tienen, todav’a fuertemente influidos por preceptos (pseudo)reli-giosos que dan a entender que oponerse a los argumentos de otros s—lo genera ren cor. Esto no es nuevo; ya lo hab’an notado los antiguos romanos: “ obsequium amicos, veritas odium parit ”; es decir, la cortes’a hace amigos, mientras que la verdad genera odio. Sin embargo, especialmente en el mbito cient’fico y cuando se trata de hacer progresar a una sociedad que, en varios aspectos, est anquilosada, es indispensable la confrontaci—n de ideas y la discusi—n nudis verbis , para lograr el mejor acuerdo. La huella de Dora Emilia y sus contribuciones a la orquideolog’a perdurarn sobre todo en el Jard’n Botnico Lankester, en el Herbario de la Universidad de Costa Rica (USJ) y en sus publica ciones, donde se preserva su legado cient’fico y el fruto de su esfuerzo prctico. E SPECIES DESCRITAS POR D ORA E MILIA M ORA DE R ETANA , COMBINACIONES NUEVAS Y NOMBRES INDITOS En total, son 10 especies nuevas, 7 combinaciones nuevas, 4 txones inditos y 3 nombres de herbario. Todos los ejemplares de los herbarios USJ y CR, citados a continuaci—n, fueron recolecta dos en Costa Rica. Cyclopogon prasophylloides (Garay) Mora-Ret. & J. Garc’a, Brenesia 37: 80. 1992. Basi—nimo: Beadlea prasophylloides Garay, Bot. Mus. Leafl. 28: 301. 1982. Con excepci—n de C. olivaceus (Rolfe) Schltr., tanto en CR como en USJ los ejemplares de este gnero terrestre estn sin determinar, lo que parece reflejar la dificultad de estudiarlo. Encyclia ionocentra (Rchb.f.) Mora-Ret. & J. Garc’a, Brenesia 33: 124. 1990. Basi—nimo: Epidendrum ionocentrum R c h b . f . , Gard. Chron. N.s. 20: 8. 1883. Epidendrum prismatocarpum Rchb.f. var. iono c e n t r u m (Rchb.f.) Teuscher, Amer. Orch. Soc. Bull. 38: 398. 1969. Hormidium ionocentrum (Rchb.f.) Brieg, Orchidee (Hamburg) 3. Auf. 1: 576. 1977. Prosthechea ionocentra (Rchb. f.) W.E. Higgins, Phytologia 82(5): 378. 1997. La publicaci—n en Brenesia citada arriba incluye una comparaci—n descriptiva y dibujos de las flores de Encyclia ionocentra y E. prismatocarpa (Rchb.f.) Dressler; la primera ha sido confundida con la segunda. Siguiendo a Higgins en Phytologia, ambas especies se tratan ahora en el gnero Prosthechea , como: P r o s t h e c h e a ionocentra (vide supra), Puntarenas; Monteverde, Bullpen, ca. 1550 m. F. Pupulin, Dora E. Mora, G. Barbosa & curso de Orquideolog’a # 880 . 16/7/1998 (USJ-71146) y P r o s t h e c h e a prismatocarpa (Rchb.f.) W.E. Higgins, San Jos; Prez Zeled—n, Villa Mills, Carretera Interamericana, Cordillera de Talamanca, ca. 3000 m. Gustavo Vargas # 302 . 26/4/1987 (USJ-71202). " Encyclia tardiflora (sp. nova?)", nom. herb. Este nombre indito ha sido utilizado por R. L. Dressler y Dora E. Mora para un taxon del cual existen dos ejemplares en USJ: Alajuela; Palmares, orillas del R’o Rastrojos. "Florece en pseudobulbos viejos" [fide Dora E. Mora, de all’ el nombre propuesto "tardiflora"]. Wilber Barahona s.n. , en floraci—n el 26/II/1994 (USJ-51074). Alajuela; Palmares, a orillas del R’o Jess Mar’a. Dora E. Mora s.n. Marzo de 1994 (USJ-57851). 2 N 2

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La especie referida debe tratarse actualmente en el gnero Prosthechea . " Epidendrum boothii (S.) Mora & Garc’a", nom. herb. Este nombre, as’ anotado en un ejemplar de herbario: Heredia; Horquetas de Sarapiqu’, Quebrada El Tigre. 600 m. Tania Bermdez # 5 . 14/6/1997 (USJ-63874), es un error, porque existe Epidendrum boothii (Lindl.) L.O. Williams, Ann. Missouri Bot. Gard. 26(4): 282. 1939, cuyo basi—nimo es Maxillaria boothii Lindl., Edwards's Bot. Reg. 24: misc. 52-53. 1838. En todo caso, en mayo de 2001 ric Hgsater anot— en el ejemplar el nombre de la especie en uso actualmente: Nidema boothii (Lindl.) Schltr. Epidendrum crassus [sic] (Dressler) Mora-Ret. & J. Garc’a, Brenesia 33: 124. 1990. Basi—nimo: Epidanthus crassus D r e s s l e r , Orqu’dea (Mx.) 9(1):15. 1983. Sin embargo, ya exist’a Epidendrum crassum C. Schweinf., Bot. Mus. Leafl. 15: 141. 1952. Un principio bsico del C—digo Internacional de Nomenclatura Botnica es que dos especies diferentes no pueden tener el mismo nombre cient’fico. As’, Barringer propuso el nuevo nombre Epidendrum insolatum Barringer, Brittonia 42(4): 245. 1991 [1992] para el taxon descrito por Dressler como Epidanthus crassus . En Brenesia 37: 126. 1992, Mora-Retana & J. Garc’a mencionan “ Epidendrum incrassatum ” , como un nombre sugerido por Robert Dressler para Epidanthus crassus . Sin embargo, desafortunadamente el art’culo se public— incompleto (falt— una pgina!), de modo que ese nombre nunca se public— vlidamente. Cuando me di cuenta de este error en Brenesia, le suger’ a Dora Emilia que publicara la informaci—n faltante en el siguiente nmero de la revista; no obstante, ella ya no ten’a voluntad para hacerlo, porque ten’a sus energ’as concentradas en nuevas publicaciones y proyectos. De modo que el nombre asignado por Barringer a esta especie tiene prioridad. Cartago; Cordillera de Talamanca, km 37, Carretera Interamericana sur. Dora E. Mora & Rosa Sandoval s.n . Abril de 1990 (USJ-37187, 37188). Epidendrum epidendroides (Garay) Mora-Ret. & J. Garc’a, Brenesia 33: 124. 1990. Basi—nimo: Neowilliamsia epidendroides Garay, Orch. Dig. 41(1): 20-21. 1977. Cartago; Parque Nacional Tapant’, sendero Palmito, 1450 m. Dora E. Mora s.n. 22/10/1992 (USJ-45209). Epidendrum talamancanum (J.T. Atwood) MoraRet. & J. Garc’a, Brenesia 33: 124. 1990. Basi—nimo: Epidanthus talamancanus J . T . Atwood, Icon. Pl. Trop. 14, pars 1: pl. 1315. 1989. Cartago/San Jos; Cordillera de Talamanca, km 36, Carretera Interamericana sur, 1850 m. I n g e Mack # 15 (USJ-49873). Varios nombres genricos utilizados en dcadas pasadas se tratan actualmente como sin—nimos de Epidendrum L. Entre stos, tenemos en la flora de Costa Rica E p i d a n t h u s L.O. Williams, N a n o d e s Lindl., Neolehmannia Krnzl., Neowilliamsia Garay y P h y s i n g a Lindl. Los estudios epidendrol—gicos llevados a cabo por ric Hgsater (AMO) han sido cr’ticos para entender poco a poco la circunscripci—n del gnero E p i d e n d r u m y poder diferenciarlo de los gneros vecinos, como E n c y c l i a y Prosthechea . Fernandezia tica Mora-Ret. & J. Garc’a, Brenesia 39-40: 164. 1993. El 27 de octubre de 1992 fue sorprendente el hallazgo repentino de una sola planta florecida de esta especie en la zona de Vara Blanca, Heredia, a ca. 2000 m de elevaci—n. La nica evidencia anterior de esta especie era una acuarela pintada por Rafael Lucas Rodr’guez en agosto de 1977, de “entre El Empalme y Ojo de Agua”, Cordillera de Talamanca (vide Rodr’guez, R.L. et al. 1986. Gneros de orqu’deas de Costa Rica. San Jos, Edit. Univ. de Costa Rica. p. 253). En la acuarela se anot— Fernandezia costaricensis (Ames & C. Schweinf.) Garay & Dunsterv., un nombre que no debe usarse por choque de sinonimia con otra especie diferente: Pachyphyllum costaricense (Ames & Schweinf.) L.O. Williams. Por ello fue necesario tanto asignar un nuevo nombre como M ORALES Dora Emilia Mora de Retana Octubre 2001 3

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LANKESTERIANA describir y publicar la nueva especie. En 1997 fueron halladas otras plantas de esta especie. Heredia; camino a Vara Blanca, ca. 2000 m. John T. Atwood & scar Rodr’guez # 4181 . 27/10/1992 (Holotipo, USJ-45216). Heredia; Cerro Chompipe, 2200 m. Dora E. Mora & R. L. Dressler s.n. 27/8/1997 (USJ-67288). Kefersteinia excentrica Dressler & Mora-Ret., Orqu’dea (Mx.) 13(1-2): 261. 1993. Cartago; La Selva, camino a Taus, 1300-1400 m. R.L. Dressler & Dora E. Mora s.n . 9/11/1984 (Holotipo, USJ-45426, isotipo en alcohol, USJ). Alajuela; Cataratas de San Ram—n. Dora E. Mora s . n . Julio de 1990 (USJ-45427, indicado como paratipo, con un duplicado en alcohol). Macroclinium doderoi Mora-Ret. & Pupulin, Selbyana 18(1): 7. 1997. Cartago; Turrialba, Ca–o Seco. Torito, ca. 1500 m. Robert L. Dressler, Dora E. Mora & R. Dodero s.n. 19/7/1994 (Holotipo, USJ-57025). Macroclinium robustum Pupulin & Mora-Ret., Selbyana 18(1): 7 & 10. 1997. San Jos; Alajuelita, en lomas sobre Barrio La Mora, ca. 1850 m. Franco Pupulin & M. Flores # 315 . 17/7/1995 (Holotipo, USJ-57743, isotipo, USJ-68694). Maxillaria cedralensis J.T. Atwood & Mora-Ret., Selbyana 18(1): 31. 1997 San Jos; Cordillera de Talamanca, Dota, El Cedral. 2000 m. Dora E. Mora & curso de Orquideolog’a s.n. Abril de 1990. (Holotipo, USJ-31579). “ Maxillaria humilis Atwood & Mora-Retana, ined.”, Brenesia 37: 99. 1992. Nombre indito incluido en la lista de orqu’deas de Costa Rica. Maxillaria rodrigueziana J.T. Atwood & MoraRet., Icon. Pl. Trop. 14: pl. 1361. 1989. Esta especie fue dibujada por Rafael Lucas Rodr’guez en junio de 1974 (vide Rodr’guez, R.L. et al. 1986. Gneros de Orqu’deas de Costa Rica. San Jos, Edit. Univ. de Costa Rica. p. 215) y fue dedicada a l p—stumamente. Hab’a sido confundi da con Maxillaria speciosa Rchb.f. de Sudamrica y con M. arachnitiflora Ames & Schweinf. El tipo de M. rodrigueziana es de El Cope, Panam ( T . Antonio 2137 , MO, SEL), pero la especie se halla tambin en Costa Rica a alturas medias de la Cordillera de Talamanca. Es una de las especies ms atractivas del gnero en Costa Rica y sus flores (entre junio y julio) son muy fragantes. San Jos; La Paulina, San Pedro, Montes de Oca. 1160 m. Planta cultivada. Rafael Lucas Rodr’guez #1003A . 5/6/1965 (2 flores) (USJ10937). “ Oerstedella satanica Hgsater & Mora-Retana, ined.”, Brenesia 37: 101. 1992. As’ se menciona en la lista de orqu’deas, pero permanece indita. En USJ se us— provisionalmente un segundo nombre para el mismo taxon: " Oerstedella diabola , ined.". El Ing. ric Hgsater (AMO) me comunic— en julio de 2001 que todav’a no est seguro de la identidad de este taxon, por cuanto hay varias especies similares involucradas. El ep’teto satnica se refiere al callo del labelo "con dos proyecciones elevadas semejantes a cuernos", segn la descripci—n indita. San Jos; Cordillera de Talamanca, Prez Zeled—n, Villa Mills, ca. 3100 m. Dora E. Mora s.n . Marzo de 1990 (USJ-31992, 32890, 32889). Ornithocephalus grex-anserinus Dressler & MoraRet., in Dressler, Novon 7(2): 120. 1997. Esta extra–a y bella planta ha sido recolectada solamente una vez. El aspecto de aves en vuelo de las flores motiv— a los autores para asignar el nom bre latino grex anserinus (bandada de gansos). Alajuela; cant—n Alajuela, distrito Sarapiqu’, alrededor de la Laguna Mar’a Aguilar. Dora E. Mora, F. Pupulin & A. Herrera s.n. 5/5/1992, en floraci—n el 11/11/1993 en el Jard’n Botnico Lankester, cuando prepar el espcimen tipo (Holotipo: USJ-49975). Reichenbachanthus lankesteri (Ames) Mora-Ret. & J. Garc’a, Brenesia 37: 81. 1992. 4 N 2

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Basi—nimo: Hexisea lankesteri Ames, Sched. Orch. 9: 47. 1925. Sin embargo, existe un basi—nimo anterior y, por tanto, prioritario, que hab’a pasado desapercibido durante mucho tiempo; se trata de S c a p h y g l o t t i s s u b u l a t a Schltr., del cual Dressler hizo la combinaci—n vlida ahora para esta especie: Reichenbachanthus subulatus (Schltr.) Dressler. No existe hasta ahora material de esta especie en USJ. En CR s—lo un ejemplar, determinado hace muchos a–os por C. Schweinfurth como H e x i s e a l a n k e s t e r i : Alajuela; San Ram—n. La Palma. 1175 m. Alberto M. Brenes 2292 . 29/8/1924 (CR-33931). Scaphyglottis geminata Dressler & Mora-Ret., Orqu’dea (Mx.) 13(1-2): 192. 1993. Cartago; Turrialba, Pejibaye, Taus. Dora E. Mora & R.L. Dressler # 31 . 20/8/1984 (USJ-32348). [En vez de agosto, podr’a ser octubre, al comparar con otro espcimen de Dressler]. Trichocentrum costaricense Mora-Ret. & Pupulin, Selbyana 15(2): 94. 1994. Alajuela; San Ram—n, Los Lagos. 800 m. M a r i o Blanco 1813, Robbin Moran, E. Watkins, E. Vargas . 23 de enero de 2001 (USJ-76755). Trichocentrum dianthum Pupulin & Mora-Ret., Selbyana 15(2): 90 & 92. 1994. San Jos; Prez Zeled—n, Las Nubes de Quizarr, 1000 m. J. Cambronero s.n . 1988 (Erbario Franco Pupulin). Floreci— en cultivo en marzo de 1989 (Holotipo, USJ-38972). “ Trichopilia pendula J. Garc’a & Mora-Retana, ined.”, Brenesia 37: 113. 1992. Nombre anotado en la lista de orqu’deas, pero no se public—. “ T r i c h o p i l i a x r a m o n e n s i s J. Garc’a & MoraRetana, ined. (nat. hyb.)”, Brenesia 37: 113. 1992. Nombre incluido en la lista de orqu’deas para designar un h’brido natural: T. marginata Henfr. x T. suavis Lindl. En Die Orchidee 51(4): 443-445, Ehrenfried Lucke public— notas sobre reproducci—n y floraci—n de este h’brido. El naturalista alemn Clarence Klaus Horich (1930-1994) hab’a enviado semillas a Alemania en enero de 1993, que exitosa mente dieron origen a plantas que florecieron siete a–os despus. Lucke se–ala, de paso, que el nombre T r i c h o p i l i a x ramonensis no ha sido vlidamente publicado, lo cual es correcto. En USJ hay dos ejemplares determinados con este nombre: Alajuela, San Ram—n. 800 1000 m. Luis Acosta s.n. Febrero de 1987, floreci— en cultivo en marzo de 1990 (USJ-31985). Alajuela; Cataratas de San Ram—n. 800-1000 m. Luis Acosta s.n. 19/3/1992 (USJ-44878). " Trichopilia xanthina Mora-Retana & J. Garc’a", nom. herb. Alajuela; San Ram—n. Dora E. Mora s.n. Floreci— en el Jard’n Botnico Lankester, junio de 1990 (USJ-31981). Este nombre fue anotado por Dora Emilia en la etiqueta de herbario, pero nunca se public—. Mario A. Blanco determin— este espcimen en 2000 como T. galeottiana A. Rich. & Gal. Trichosalpinx navarrensis (Ames) Mora-Ret. & J. Garc’a, Brenesia 37: 125. 1992. Basi—nimo: Pleurothallis navarrensis A m e s , Sched. Orch. 9: 34-35, fig. 5. 1925. No existe hasta ahora material de esta especie en USJ. En CR s—lo hay un ejemplar: Puntarenas; Pen’nsula de Osa. Aguabuena de Rinc—n, Fila Casa Loma, siguiendo el antiguo camino a Rancho Quemado. 8 43’ 00’’ N, 83 32’ 00’’ O. 500 m. Gerardo Herrera 4440 . 2/10/1990 (CR-210573). M ORALES Dora Emilia Mora de Retana Octubre 2001 5 E SPECIES DEDICADAS A D ORA E MILIA M ORA Y A SU FAMILIA 1. Epidendrum mora-retanae Hgsater, Icon. Orchid. 2, pars 1: pl. 160. 1993. Alajuela; Lajas de Alfaro Ruiz. Dora E. Mora s.n . Julio de 1997 (USJ-67287). Esta especie habita desde Nicaragua hasta Panam entre 600 y 1800 m de altitud. El holotipo es de Panam; Cuesta de Piedra, Finca El Milagro, 800-1000 m. Leg. ric Hgsater 6499 & R.L. Dressler (AMO). 2. Kefersteinia retanae G. Gerlach ex C.O.

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LANKESTERIANA Morales, Brenesia 52: 75. 1999 [2000] Gnter Gerlach (M y Jard’n Botnico de Mnich, Alemania) public— esta especie en Brenesia 41-42:100, 1994, pero faltaba informaci—n necesaria para validar el nombre. En Brenesia 52: 75, 1999, se aporta esa informaci—n. San Jos, Prez Zeled—n, Pe–a Blanca. 600 m. Gnter Gerlach s.n . Floreci— en cultivo en el Jard’n Botnico de Mnich en junio de 1992 (Holotipo, USJ-69249). 3. Sobralia doremiliae Dressler, Novon 5(2): 142. 1995. Cartago; Para’so, Orosi, Parque Nacional Tapant’. Dora E. Mora & R.L. Dressler s.n. 1 2 / 1 1 / 1 9 9 3 (Paratipo, USJ-50001). En el art’culo, Dressler explica que prefiri— eliminar la a de Dora en doremiliae para evitar una pronunciaci—n err—nea. En lat’n clsico, doraemiliae se pronunciar’a "doraimiliai", mientras que en lat’n vulgar, que es ms utilizado por los botnicos, la pronunciaci—n de doremiliae e s "doremilie". 4. Stelis morae Luer, Lindleyana 11(2): 103, f. 32. 1996. El ejemplar tipo ( C. Luer 17375 ) fue recolectado por Dora E. Mora & R.L. Dressler en San Carlos de Alajuela en enero de 1990. En floraci—n en el Jard’n Botnico Lankester el 7/3/1995. La publicaci—n indica que el holotipo est en el Herbario Nacional de Costa Rica (CR), lo cual no es cierto. Un isotipo debiera hallarse en MO. Un solo ejemplar en USJ: San Jos: Prez Zeled—n, sin ms datos. Leg. W. Fonseca s.n. ( F. Pupulin 1710 ). Floreci— en culti vo en San Isidro del General. 21/1/1999. (USJ71130). 5. Telipogon retanarum Dodson & Escobar, Orquideolog’a 17(2): 114. 1987. Una de las muchas especies de orqu’deas del Parque Nacional Tapant’, en Cartago. Es endmica en Costa Rica y probablemente posee poblaciones muy peque–as, pues s—lo se conocen dos ejemplares tipo de 1984 y, de la misma localidad, dos ejemplares ms de 1985, como consta en Orquideolog’a 17(2): 119. 1987. Cartago; Parque Nacional Tapant’, 1630 m de alti tud, ca. a 6 km de la entrada, 200 m despus de una catarata grande. Rodrigo Escobar, D. Portillo & M. Retana 3628 . 8/8/1985 (USJ27440). Este ejemplar tiene una etiqueta manuscrita de Dora Emilia, con el # 3555 de R. Escobar et al. de 1984, que es el tipo de la especie. En marzo de 1992, tanto Dora Emilia como Rodrigo me indicaron que el ejemplar de USJ era un isotipo. Sin embargo, al revisar la publicaci—n en Orquideolog’a y una etiqueta de herbario ms una nota que envi— Escobar en julio de 2001, es claro que se trata de R. Escobar et al. # 3628, que no es un tipo. Aun as’, es un ejemplar tan valioso como un tipo, pues procede de la localidad t’pica y hasta ahora es el nico de esta especie en los herbarios de Costa Rica. P UBLICACIONES DE D ORA E MILIA M ORA DE R ETANA Y SUS COLEGAS (En orden cronol—gico) Mora, Dora E. 1971. Desarrollo y crecimiento del pice vegetativo en Acnistus arborescens . I. Estructura, cambios plastocr—nicos y primeros estadios en el desarrollo de la hoja. Rev. Biol. Trop. 19(1,2): 171-189. Mora, Dora E. 1975. Desarrollo y crecimiento del pice vegetativo en Acnistus arborescens . II. Desarrollo del procambio y los tejidos vasculares de la hoja y crecimiento de la lmina foliar. Rev. Biol. Trop. 23(2): 155-163. Mora-Retana, Dora E. (sin a–o, circa 1983). G o n g o r a . I n : Asociaci—n Costarricense de Orquideolog’a (ACO). Orqu’deas, su cultivo en Costa Rica. San Jos, Impresora Delta. p. 26-28. Rodr’guez, R.L., Mora, Dora E., Barahona, Mar’a E. & Williams, N.H. 1986. Gneros de orqu’deas de Costa Rica. San Jos, Edit. Universidad de Costa Rica. 334 p. Mora Monge, Dora E. & Valerio G., C. 1988. 6 N 2

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Polinizaci—n y producci—n de frutos en la guaria morada ( Cattleya skinneri , Orchidaceae) en Costa Rica. Orquideolog’a 17(3): 242-251. Mora-Retana, Dora E. & Garc’a-Castro, J. 1990. Nuevas combinaciones de Orchidaceae de Costa Rica. Brenesia 33: 123-126. Mora, Dora E. 1991. El gnero L o c k h a r t i a H o o k . I n : Orqu’deas Nativas de Colombia 54(2). Sociedad Colombiana de Orquideolog’a. Mora-Retana, Dora E. & Atwood, J.T. 1992. Orchids of Costa Rica, part 2. Ic. Pl. Trop. 15, pl. 1401-1500. Mora-Retana, Dora E. & Garc’a-Castro, J.B. 1992. Nuevas combinaciones en Orchidaceae de Costa Rica (II). Brenesia 37: 125-126. Mora-Retana, Dora E. & Garc’a-Castro, J.B. 1992. Lista actualizada de las orqu’deas de Costa Rica (Orchidaceae). Brenesia 37: 79-124. Dressler, R.L. & Mora-Retana, Dora E. 1993. Scaphyglottis geminata , an attractive new species from Costa Rica. Orqu’dea (Mx.) 13(1-2): 191194. Dressler, R.L. & Mora-Retana, Dora E. 1993. Kefersteinia excentrica , a distinctive new species from Costa Rica. Orqu’dea (Mx.) 13(1-2): 261264. Garc’a-Castro, J.B. Mora-Retana, Dora E. & Rivas, Marta E. 1993. Lista de orqu’deas comunes a Costa Rica y Panam. Brenesia 39-40: 93-107. Mora-Retana, Dora E. & Atwood, J.T. 1993. Orchids of Costa Rica, part 3. Ic. Pl. Trop. 16, pl. 1501-1600. Mora-Retana, Dora E. & Garc’a-Castro, J.B. 1993. Fernandezia tica Mora-Retana & J.B. Garc’a, una nueva especie orquidcea de Costa Rica. Brenesia 39-40: 163-165. Gonzlez, Mar’a I., Mora-Retana, Dora E. & Warner, J. 1994. Crecimiento de C a t t l e y a dowiana (Orchidaceae) en varias condiciones de cultivo. Rev. Biol. Trop., 42 (1/2):73-79. Pupulin, F. & Mora-Retana, Dora E. 1994. A revi sion of the Costa Rican species of Trichocentrum (Orchidaceae). Selbyana 15(2): 87-103. Mora-Retana, Dora E. & Warner, J. 1995. La con servaci—n de las orqu’deas en el Jard’n Botnico Lankester. I n : Asociaci—n Costarricense de Orquideolog’a (ACO). Orqu’deas de Costa Rica y su cultivo. Vol. I: Cattleya y gneros relacionados. San Jos, Litograf’a e Imprenta Lil. p. 54-56. Gonzlez L., Mar’a I. & Mora-Retana, Dora E. 1996. El color de las flores en la taxonom’a de cuatro especies de Stanhopea ( O r c h i d a c e a e ) . Rev. Biol. Trop. 44(2): 525-539. Mora-Retana, Dora E. & Gonzlez L., Mar’a I. 1996. Variabilidad floral de cuatro especies de Stanhopea (Orchidaceae). Rev. Biol. Trop. 44(2): 517-523. Atwood, J.T. & Mora, Dora E. 1997. M a x i l l a r i a cedralensis , a new species from the highlands of Costa Rica. Selbyana 18(1): 31-32. Mora, Dora E. & Warner, J. 1997. Orqu’deas de Costa Rica. Cartago, Jard’n Botnico Lankester, Universidad de Costa Rica. 32 p. Mora-Retana, Dora E. & Pupulin, F. 1997. Two new species of Macroclinium (Orchidaceae) from Costa Rica. Selbyana 18(1): 7-10. Atwood, J.T. & Mora-Retana, Dora E. 1999. Orchidaceae: Tribe Maxillarieae: Subtribes Maxillariinae and Oncidiinae. In: Burger, W. (ed.). Flora costaricensis. Fieldiana, Bot., n.s. 40: 1-182. Agradezco sinceramente la colaboraci—n de Robert L. Dressler (FLAS), ric Hgsater (AMO) y Jorge Warner (Jard’n Botnico Lankester) en la preparaci—n de este escrito. M ORALES Dora Emilia Mora de Retana Octubre 2001 7

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It was with severe regrets that I learned of the untimely death of Dora Emilia Mora de Retana. In 1985 while completing the Orchidaceae for Flora of La Selva Biological Station, I started a search for a Costa Rican with whom to collaborate on a larger project. After Robert Dressler suggested approaching Dora Emilia Mora, I visited the Universidad de Costa Rica where I found her work ing patiently with a student over herbarium specimens. I introduced myself forwardly proposing to collaborate on treatments of Costa Rican orchids, an approach that visibly overwhelmed this reserved Costa Rican professor. But I vowed to develop a project that would benefit her and Costa Rica as much as myself and Selby Gardens. We made a pactI would push her and she would push me. This led over several years to mutually beneficial interactions resulting in three hundred Costa Rican orchids illustrated under Icones Plantarum Tropicarum. This activity culminated in a volume with the Field Museum of Natural History featuring subtribes Maxillariinae and Oncidiinae with help from Franco Pupulin, German Carnevali, and Joaqu’n Garc’a. Dora Emilia invited me frequently to visit Lankester Gardens to share a garden visibly thriv ing despite dauntingly severe budget constraints. She kept a vision of Lankester Gardens not only as a tourist garden for generating revenues but as a garden dedicated to Costa Rican orchid research. The living collection of Costa Rican native orchids served as a basis for most of her descriptions and line drawings. In later years I learned that much of her success maintaining Lankester Gardens lay in creating a positive working atmosphere for fostering students, employees, and orchid enthusiasts. Garden Manager scar Rodr’guez would proudly show his inspired landscapesthe cactus garden, the water fall, even a Japanese garden; Jorge Warner his propagation lab; Carlos Quir—s the exquisitely curated orchid collection. Carlos, Jorge, and Oscar would soon visit Selby Gardens with Dora Emilia’s help. Their enthusiastic interaction and good cheer proved them to be the best of ambassadors. Dora Emilia also introduced Costa Rican orchid society member and collaborator Joaqu’n Garc’a to me, a man of prodigious memory. One evening he brought me a spidery flowered maxillaria to my hotel. I identified the maxillaria as M a x i l l a r i a arachnitiflora , but the plant was odd in that it was somewhat miniature for the species. I was telling him nothing new, but for that moment the plant would remain M. arachnitiflora . Dora Emilia made several visits to Selby Gardens finalizing treatments of Costa Rican orchids, once bringing her husband Marco who volunteered his time complet ing the organization of the spirit collection. Beyond her professional accomplishments, her greatest legacy is to rise above personal ambitions to foster efforts of those around her. Because of her, Lankester Gardens is blessed with a talented, imaginative, and altruistic staff continuing the tra dition of fostering others, a tradition that will carry this garden to a higher level of international recog nition. In a very real sense her spirit lives on. LANKESTERIANA 2: 9. 2001. DORA EMILIA MORA DE RETANA, A REMEMBRANCE J OHN T. A TWOOD Marie Selby Botanical Gardens Missouri Botanical Garden

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The collection of tiny orchids for botanical study is frustrating. If it is available, one gathers what seems to be abundant material but when the plants are dried, there is little left. The plant to be described here is large for a S t e l l i l a b i u m , being about 20 cm tall and branched. Even so, each plant may have few flowers at a given moment. I first saw the species along the R’o Pejibaye at La Selva de Cartago in 1984, where we were mistakenly calling the locality Taus, which I mistakenly spelled as Taos. Being reluctant to gather enough material for a good specimen, I convinced myself that it must be a named species. We photographed a flower, and a photograph was published in the Field Guide (Plate 30, fig. 4, Dressler 1993) as Stellilabium bullpenense Atwood, which it is not by any stretch of the imagination. A few minutes after photographing that Stellilabium , we crossed a pas ture and found another species at the edge of the forest. This tiny plant, with a single flower, was pressed and I later made a label for it, as number 310 of the plants collected by Dressler and Biolog’a 350 and sent the label to the University of Costa Rica Herbarium (USJ), where the first set of speci mens is deposited. When I looked at the Stellilabium specimens at USJ in 1999, I found my label number 310 mounted with a plant of the large plant from the gallery forest. Someone else had pressed a specimen, and the tiny plant for which the label was intended was simply lost among the newspapers. Later specimens were collected by Dressler and Mora in 1991, by Atwood and Mora in 1992, and by Pupulin, Spadari and the Orchidology Course in 2000. The best of these specimens originally had several flowers but they were destroyed by insects. The remaining specimens have only a few flowers, but the label of Atwood & Mora 4202 mentions two flowers in alcohol. I asked Franco Pupulin to search for the flowers in alcohol, and he found them to be the same as his own collection from the same locality. There is not much good material of this species, but it is more than is available of most other Central American species of Stellilabium and with Franco's excellent drawing it can finally receive a name. Stellilabium (Taeniorhachis) erratum Dressler, sp. nov. F IG . 1. T Y P E : C O S T A R I C A . Cartago: Cant—n Para’so, Dist. Orosi, Tapant’, Sendero Oropndola, en P. N. Tapant’, 1350 m, 2 nov. 1992, ep’fita, flores moradas, J. T. Atwood & D. E. Mora 4202 (holo type, USJ). Rhachis alata, labellum trilobatum, medio inflatum; columna trilobata, lobulis lateralibus setis stel latis praedita, lobulo medio hispido. Roots flattened, 1.5-2 mm wide; leaves 16-37 x 34 mm, elliptic-oblanceolate, acute; stems b a s a l l y terete, then sulcate; peduncular bracts 1.8-2.4 x 1.5 mm, acute; rachis flattened, 1-1.2 mm wide, up to 16 cm long; floral bracts 1-1.2 x 1.3 mm, triangular, acute; pedicel and ovary 6.5 mm; flowers d a r k wine-purple; dorsal sepal 3-3.5 x 2-2.2 mm, ovate, LANKESTERIANA 2: 11-13. 2001. STELLILABIUM ERRATUM , A COMEDY OF BLUNDERS R OBERT L. D RESSLER Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Florida Museum of Natural History; Marie Selby Botanical Gardens Mailing address: 21305 NW 86th Ave., Micanopy, Florida 32667, U.S.A. R ESUMEN . Se describe Stellilabium erratum , una especie de Tapant’ y el valle del R’o Pejibaye, de planta y flores relativamente grandes para el gnero. La columna y el labelo son trilobados y los l—bulos laterales de la columna llevan setas conspicuas. K EY W ORDS : Orchidaceae, Stellilabium sect. Taeniorhachis, Stellilabium erratum , Costa Rica

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LANKESTERIANA Figure 1. Stellilabium erratum Dressler. A Habit. B Flower. C Perianth parts flattened. D Column and lip, lateral view. E Bristles of lateral lobes of column. F. Bristles of midlobe of column. G. Pollinarium without and with anther cap. Based on Pupulin et al. 2266 . 12 N 2 A F E D C B G

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subacute; lateral sepals 3-4 x 1.6-2 mm, obliquely ovate, obtuse, apiculate; petals 3.3-3.5 x 0 . 9 1 . 1 mm, narrowly elliptic-oblong, subacute, margins ciliate; lip 3-3.5 x 3-3.3 mm, 3-lobed, lateral lobes 1.3-2.5 x 0.4-0.5 mm, lanceolate, fleshy, papillose, midlobe 2.5-3 x 1.5-2.5 mm, ovate-triangular, acuminate, thickened in middle, puberulent-papillose; column 3-lobed, lateral lobes 1.3-1.5 x 0.4-0.8 mm, oblong, flattened, the dorsal margins with dense band of stellate bristles ca. 0.7 mm long, midlobe rounded, ca. 1.5 mm, bristles simple, ca. 0.4 mm; capsule ovoid, ca. 7 x 3 mm, including beak ca. 1.5 mm, fruiting pedicel ca. 3 mm. Among Central American species of Stellilabium , S. erratum is distinguished by its large size (for a Stellilabium ), by the deeply 3-lobed lip and the deeply 3-lobed column. The 3-lobed column would suggest its placement in D i p t e r o s t e l e , according to Garay and Romero (1998), but none of the species described from South America has the flattened, tapeworm-like rachis of the Central American complex. The section Taeniorhachis has been proposed for the Central American group (Dressler 1999). The available samples of DNA are very few (one species each of Dipterostele in the strict sense, Stellilabium in the strict sense and Taeniorhachis ), but the analysis does suggest that the three may be distinct groups (N. H. Williams, pers. comm.) Further and better samples, though, may well show that all three are subgroups of Telipogon in the broad sense. The epithet erratum , or error, refers to the improbable series of errors that started when we first saw the plant by the R’o Pejibaye. P A R A T Y P E S : Same locality as type, 1200 m, 9 apr. 2000, F. Pupulin, L. Spadari y Curso de Orquideolog’a 2266 (USJ). "Taos," altura aprox. 1000 m, 11 dic. 1984, por el borde entre selva y potrero, R. L. Dressler y Biolog’a 350 no. 310 (USJ). La Selva. Valle del R’o Pejibaye, elev. aprox. 1000 m, 18 marzo 1991, ep’fita, cerca del r’o, flores morado-rojo, R. L. Dressler y D. E. Mora de Retana 6136 (USJ). A CKNOWLEDGEMENTS . I am very much indebted to the late Dora Emilia Mora, for helping me to reach this and many other interesting localities, and Franco Pupulin for the excellent drawing. C ITED L ITERATURE Dressler, R. L. 1993. Field guide to the orchids of Costa Rica and Panama. Ithaca, Cornell University Press. Dressler, R. L. 1999. A reconsideration of Stellilabium and Dipterostele . Harvard Pap. Bot. 4: 469-473. Garay, L. A., & G. A. Romero-Gonzlez. 1998. Schedulae Orchidum. Harvard Pap. Bot. 3: 53-62. D RESSLER Stellilabium erratum Octubre 2001 13

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The circumscription of genera within the largest tribes of the Orchidaceae has not yet reached a gen eral agreement among taxonomists. The subtribe Pleurothallidinae, restricted to the tropics and sub tropics of the New World, is perhaps the largest taxonomic orchid group, with 28 genera and an estimation of more than 3000 species (Dressler 1993). The genus Myoxanthus was first described by Endlicher on M. monophyllus (Poeppig & Endlicher 1835), based on a plant collected in the Peruvian department of Huanuco by Poeppig. In 1847 another species currently assigned to Myoxanthus was identified as a distinct genus and described as Duboisia by Karsten (later renamed D u b o i s R a y m o n d i a Karsten due to the priority of Duboisia R. Br. in the Solanaceae) (Karsten 1847, 1848). A third attempt to assign species of Myoxanthus to a new genus was made by Barbosa Rodrigues (1882), who described Chaetocephala to accommodate its former Restrepia lonchophylla . In the opinion of leading orchid taxonomists at that time, notably Lindley, Reichenbach, and Cogniaux, the criteria selected to define the new genera were insufficient to sustain the proposals, and Myoxanthus species were newly submerged into the conglomerate genus Pleurothallis . It was not until 1982 when the genus Myoxanthus was resurrected by Luer who recognized 34 species sharing a combination of LANKESTERIANA 2: 15-18. 2001. MYOXANTHUS VITTATUS (ORCHIDACEAE), A NEW SPECIES FROM COSTA RICA F RANCO P UPULIN 1 and M ARIO A. B LANCO 2 1 Jard’n Botnico Lankester, Universidad de Costa Rica Research Associate, Marie Selby Botanical Gardens, Sarasota, U.S.A. P.O. Box 1031-7050 Cartago, Costa Rica. fpupulin@cariari.ucr.ac.cr 2 Instituto Centroamericano de Investigaci—n Biol—gica y Conservaci—n P.O. Box 2398-250 San Pedro de Montes de Oca, San Jos, Costa Rica Present mailing address: Herbarium, Florida Museum of Natural History, University of Florida 385 Dickinson Hall, P.O. Box 117800 Gainesville, Florida 32611-7800, U.S.A. A BSTRACT . The taxonomic position of Myoxanthus Poepp. & Endl. (Orchidaceae: Pleurothallidinae) is discussed and Silenia Luer is maintained as a subgenus of M y o x a n t h u s . A new species of Myoxanthus is described and illustrated. Myoxanthus vittatus Pupulin & M.A. Blanco is apparently endemic to the low, tropical wet forests of central Pacific Costa Rica, where it has been collected in two different places. It differs from closely related species for the small size of the plant, the whitish flowers, longitudinally striped with purple, the densely pubescent ovary, and the ligulate, obtuse lip. The new combination Myoxanthus tomentosus (Luer) Pupulin & M.A. Blanco is proposed. R E S U M E N . Se discute la posici—n taxon—mica de Myoxanthus Poepp. & Endl. (Orchidaceae: Pleurothallidinae) y se mantiene a Silenia Luer como subgnero de Myoxanthus. Se describe y se ilustra una nueva especie de Myoxanthus . Myoxanthus vittatus Pupulin & M.A. Blanco es aparente mente endmico de los bosques tropicales de bajura en el Pac’fico Central de Costa Rica, donde ha sido recolectado en dos diferentes localidades. Difiere de las especies afines por el tama–o peque–o de las plantas, por sus flores blanquecinas con estr’as longitudinales prpura, por el ovario densa mente pubescente y por el labelo ligulado y obtuso. Se propone la nueva combinaci—n Myoxanthus tomentosus (Luer) Pupulin & M.A. Blanco. K E Y W O R D S : Orchidaceae, Pleurothallidinae, M y o x a n t h u s , Myoxanthus sect. S i l e n i a , M y o x a n t h u s vittatus , Costa Rica

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LANKESTERIANA characters sufficiently distinct to warrant generic recognition, among which the presence of tubular sheaths concealing the ramicaul, often bearing digitate trichomes, a leaf abscission layer without the “node” which may be found in racemose species of P l e u r o t h a l l i s , the fascicled inflorescence, and the often thickened apices of the petals, sometimes developed into osmophores (Luer 1982). The original circumscription of Myoxanthus was emended by Luer (1992) in his systematic monograph of the genus, with the description of subgenus Silenia to include the P . aspasicensis complex, encompassing 3 species with a fascicle of single, successive flowers borne terminally from the ramicaul, fleshy flowers exter nally shortpubescent, and ciliate anther cap. Eventually in 1995 Luer proposed to include Myoxanthus subgenera Satyria and Silenia i n Pleurothallis subgenus Acianthera in order to restore a circumscription of Myoxanthus sensu stricto (Luer 1995). The generic delimitation of Myoxanthus among other pleurothalloids is supported by a unique assemblage of morphological characteristics and anatomical features revised by Pridgeon and Stern (1982). The presence of two series of foliar veins and a medullated stele in the root is characteristic of Myoxanthus and may be found elsewhere in the Pleurothallidinae only in Octomeria and in the Pleurothallis aspasicensis complex. Spiral thicken ings in hypodermal cells and mesophyll idioblasts, which are present in most pleurothallids, are notably absent from all the tissues of M y o x a n t h u s and the P. aspasicensis complex (Pridgeon & Stern 1982). In view of the diagnostic anatomical characters which allow recognition of evolutionary affinities in the Pleurothallidinae and to preserve the natural ness of M y o x a n t h u s , we prefer to maintain the P . a s p a s i c e n s i s complex as Myoxanthus s u b g e n u s Silenia and we therefore propose the following Myoxanthus species new to science: Myoxanthus vittatus Pupulin & M.A. Blanco, s p . nov. F IG . 1 TYPE: C OSTA R ICA . San Jos: Prez Zeled—n, El Brujo, near R’o Divisi—n, 450 m, 30 Jan. 2000, flowered in cultivation at Jard’n Botnico Lankester, 3 June 2001, M.A. Blanco 1324 ( h o l o type, USJ!; isotype, SEL!). Species Myoxantho aspasicensi (Rchb.f.) Luer similis, sed planta floribusque omnine minoribus, ovario pubescente, sepalis purpureo vittatis, interne laevis, synsepalo lanceolato, apice petalorum dorsaliter tuberculato, labello ligulato apice laevi differt. Plant epiphytic, cespitose, up to 17 cm tall, roots coarse. Ramicauls stout, erect, 4-7 cm long, enclosed by 3-4 loose, tubular sheaths sometimes fragmented. Leaf erect, thickly coriaceous, narrow ly lanceolate-elliptic, minutely emarginate, 7-10 cm long, 1.3-1.5 cm wide, cuneate below into a condu plicate, sessile base. Inflorescence a fascicle of sin gle, successive flowers, at the apex of ramicaul, the peduncle sparsely pubescent, 8-13 mm long, subtended by a papyraceous spathe 1.4-1.6 cm long; pedicels 2-3 mm long; ovary less than 1 mm long, densely pubescent. Flowers small, whitish, longitudinally striped with purple, fleshy, densely short-pubescent externally, glabrous within. Dorsal s e p a l linear-oblanceolate, obtuse, 10-12 mm long, 3 mm wide, 5-veined. Lateral sepals connate into a lanceolate, shortly emarginate, concave synsepal, 10-13 mm long, 4-5 mm wide, each half 4-veined. Petals fleshy, linear-oblong, obtuse, minutely apic ulate, adaxially provided with a tuft of short papil lae near the apex, 4 mm long, 1 mm wide, 3veined. Lip 3-lobed, ligulate, subacute, 4 mm long, 2.5 mm wide between lateral lobes; the apical lobe smooth, slightly undulate along the margins; the lateral lobes erect, narrowly uncinate, antrorse; the disc with an erect, narrow, low, channeled callus above the base, extending in front into a low keel just to near the lip apex, laterally provided with a pair of low lamellae extending to the apex, hinged on the end. Column arcuate, semiterete, 1.6 mm long, provided with narrow wings above the middle, the foot less than 1 mm long. Anther cap glo bose, with long hairs on the upper margin, 2-celled. Pollinia 2, obovoid, flattened, on a short caudicle. E T Y M O L O G Y : from the Latin v i t t a t u s , “longitudinally striped”, in reference to the stripes on sepals and petals. 16 N 2

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P UPULIN and B LANCO Myoxanthus vittatus Figure 1. Myoxanthus vittatus Pupulin & M.A. Blanco. A Habit. B Flower. C Dissected flower ( petal to the left in abaxial view). D Column and lip, lateral view. E Lip, spread. F Pollinarium and anther cap. I LLUSTRATION VOUCH ER : M.A. Blanco 1324 (USJ). Drawn from the holotype. Octubre 2001 17 B A C E D F 5 cm 1 cm 3 mm 3 mm 1 mm 5 mm

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LANKESTERIANA D ISTRIBUTION AND E COLOGY : Known only from two specimens from the same area in central Pacific Costa Rica, epiphytic in tropical wet forest, premontane belt transition, at 450 m elevation. P HENOLOGY : Flowering occurs from March to June. P A R A T Y P E : C O S T A R I C A . San Jos: Prez Zeled—n, Viento Fresco del Brujo, 450 m, epiphytic on culti vated trees in the village, 21 Jan. 2001, flowered in cultivation at Gaia Botanical Garden, 13 June 2001, F. Pupulin, D. Castelfranco & L. Elizondo 2878 (USJ, Spirit Coll.!). Myoxanthus vittatus is closely related to M . aspasicensis, Pleurothallis tomentosa, M. sem p e r g e m m a t u s and M. uncinatus , but it is distinguished by the small size, the densely pubescent ovary, the whitish flower striped with purple, the sepals adaxially smooth, and the ligulate, obtuse lip. Pleurothallis tomentosa Luer was described from a single Costa Rican collection made by Endres around 1867 and kept at Reichenbach’s herbarium in Vienna, and it is supposed to be extinct (Luer 2000). This species may be easily dis tinguished from M. vittatus by the purple-black flowers with an oblong lip, rounded at apex. In order to avoid paraphyly in Myoxanthus as intend ed here, a new combination is required for Endres’ species: Myoxanthus tomentosus (Luer) Pupulin & M.A. Blanco, comb. nov. Bas.: Pleurothallis tomentosa Luer, Orquideolog’a 21(3): 337. 2000. C ITED L ITERATURE Barbosa Rodr’gues, J. 1882. C h a e t o c e p h a l a , Gen. Spec. Orch. Orchid. Nov. 2: 37. Dressler, R.L. 1993. Phylogeny and classification of the orchid family. Dioscorides Press, Portland. Karsten, H. 1847. Duboisia . Allg. Gartenz. 15: 394. _____. 1848. Dubois-Raymondia . Bot. Zeitung (Berlin) 6: 397. Luer, C.A. 1982. A reevaluation of the genus Myoxanthus (Orchidaceae). Selbyana 7(1): 34-54. _____. 1992. Icones Pleurothallidinarum IX. Systematics of M y o x a n t h u s . Addenda to Platystele, Pleurothallis subgenus Scopula, and Scaphosepalum . Monogr. Syst. Bot., Missouri Bot. Gard. 44: 1-128. _____. 1995. A re-evaluation of the Pleurothallid subgen era Satyria and Silenia (Orchidaceae). Monogr. Syst. Bot., Missouri Bot. Gard. 57: 146. _____. 2000. Miscellaneous new species in the Pleurothallidinae. Orquideolog’a 21(3): 318-340. Poeppig, E.F. & S.L. Endlicher. 1835. Myoxanthus mono phyllus . Nov. Gen. Sp. Pl. 1: 50, t. 88. Pridgeon, A.M. & W.L. Stern. 1982. Vegetative anatomy of Myoxanthus (Orchidaceae). Selbyana 7(1): 55-63. 18 N 2

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In 1993 Czech botanist Jiri (George) Haager kindly honored me by describing a new genus of Vietnamese orchids Christensonia (Haager 1993). In that account and later mention of the genus (Christenson 1996), space consideration did not allow publication of an excellent line drawing of the type plant by Haager. This article rectifies this situation. A member of the subtribe Aeridinae (synonym: Sarcanthinae), Christensonia exhibits a mosaic of characters found in the closely related genera Aerides Loureiro, Rhynchostylis Blume, and Vanda Jones. Christensonia differs from Aerides by having no column foot. And unlike most species of A e r i d e s , the lip of Christensonia is not flexibly hinged below the lateral lobes allowing deflexion of the lip by pollinators. Christensonia is similar to Rhynchostylis , especially in having a narrowly lin ear stipe to the pollinarium, but differs by having a clearly three-lobed lip. In addition, plants of Christensonia are miniature and few-flowered compared with Rhynchostylis . Christensonia is perhaps most similar to Vanda . In particular, Christensonia shares two characteris tics with Vanda . First, the spurs are straight and in the same plane as the midlobe of the lip. Second, the only flexible portion of the lip is the base of the midlobe. The green and white flowers of C h r i s t e n s o n i a are unlike the generally white and rose-purple pat terns seen in Aerides and Rhynchostylis. However, they are similat to some species of Vanda, especial ly those of Vanda section Cristatae Lindley which are sometimes segregated as the genus T r u d e l i a Garay (q.v., Christenson 1994). Nothing is known of the pollinaton biology of Christensonia nor the significance, if any, of the green and white floral coloration. Plants of Christensonia have entered horticulture and are readily available commercially in the United States. A plant cultivated in Sarasota, Florida, for several years has proven to be vigorous and trouble-free to grow. Its first flowering was on a plant of the similar size to the one shown in the drawing. Plants in the Czech Republic and the United States flower in july. Its flowering season in nature is unknown. Christensonia vietnamica J.R. Haager, Orchid Digest 57(1): 40. 1993. TYPE. Southern Vietnam, Prov. Khanh Hoa, Ninh Tay village, 100 m, April 1989, collected by O. Dubec and J. Jurak, flowered in cultivation in Prague, PR 11392 (holotype, PR). F IG . 1. Monopodial epiphytes. Stems to 30 cm long. Leaves strap-shaped, erose at apex, to 6 x 1 cm. LANKESTERIANA 2: 19-21. 2001. THE GENUS CHRISTENSONIA HAAGER E RIC A. C HRISTENSON 1646 Oak Street, Sarasota, Florida 34236, U.S.A. A BSTRACT . The monotypic Vietnamese genus Christensonia Haager is discussed and illustrated by a detailed line drawing. A warm-growing, lowland plant, C. vietnamica is an exciting addition to the tropical horticulture as well as a taxonomic bridge between the genera Aerides , Rhynchostylis , and Vanda . R ESUMEN . Se discute y se ilustra con un dibujo de plumilla detallado el gnero monot’pico vietnami ta Christensonia Haager. Una planta de clima clido, nativa de tierras bajas, C. vietnamica es un interesante aporte a la horticultura tropical, as’ como un puente taxon—mico entre los gneros Aerides , Rhynchostylis y Vanda . K EY W ORDS : Orchidaceae, Aeridinae, Christensonia , Christensonia vietnamica , Vietnam 

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LANKESTERIANA Figure 1. Christensonia vietnamica J.R. Haager. 1. Habit. 2. Leaf apices. 3. Flower from below. 4. Longitudinal sec tion of the lip. 5. Anther. 6. Pollinarium. 7. Pollinium. 8. Stipe. 9. Sepals and petal. 10. Fruit. 11. Cross-section of the ovary. Drawing of the type plant by Jiri Haager. 20 N 2 

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C HRISTENSON The genus Christensonia Inflorescences 1-3, erect, few-flowered, to 8 cm long. Flowers 3-5, resupinate, somewhat nodding, the sepals, petals and lateral lobes of the lip yellowgreen, the midlobe of the lip white. Sepals and petals subequal, to 1.6 x 0.7 cm, the dorsal sepal oblong-elliptic, obtuse, convex, the lateral sepals obliquely elliptic-obovate, obtuse, convex, the petals elliptic, obtuse, somewhat incurved. L i p three-lobed, to 3.4 cm long from the apex of the spur to the apex of the midlobe, the lateral lobes rectangular, obtuse, to 0.7 x 0.3 cm, the midlobe fan-shaped with a broad claw, with finely toothed and undulate margins, to 1.8 x 1.6 cm, the spur with small, scale-like, internal glandular processes below the apex. Column very short, to 0.3 cm long. Pedicel and ovary six-winged, twisted. A C K N O W L E D G E M E N T S . I graciously thank Jiri Haager for honoring me with this commemoration and for lending the accompanying illustration for this article. L ITERATURE CITED Christenson, E.A. 1994. Taxonomy of the Aeridinae with an infrageneric classification of Vanda Jones ex R. Br. Proc. 14th World Orch. Conf. 206-216. _____. 1996. Sarcanthinae genera 22: C h r i s t e n s o n i a . Orchids 65(3): 282-283. Haager, J.R. 1993. Some new taxa of orchids from Southern Vietnam. Orchid Digest 57(1): 39-44. Octubre 2001 21 

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Introduction . Orchids need endomycorrhizae to develop: green orchids need fungi for the early developmental stages, while orchids lacking chlorophyll are always completely dependent upon fungi. Endophytes of orchids are always Mitosporic Fungi. Three classes are recognized in this group: Hyphomycetes, Coelomycetes, and Agonomycetes. To the latter class belong sterile fungi that may pro duce chlamydospores or sclerotia and the genus R h i z o c t o n i a DC., whose members may also produce sclerotia. Most orchid endophytes are ascribed to this group. Only few traits were used to describe this genus (Curtis 1937) but unfortunately they have no taxonomic value since they do not allow discrimination between very similar R h i z o c t o n i a species with very different perfect states. Rambelli (1981) suggested the presence of dolipores or clamp connections as a character to assign a fungus to the basidiomycetes when reproductive structures are absent. Moreover, Riess LANKESTERIANA 2: 23-30. 2001. ENDOPHYTES OF SERAPIAS PARVIFLORA PARL. AND SPIRANTHES SPIRALIS (L.) CHEVALL. (ORCHIDACEAE): DESCRIPTION OF ENDOPHYTES OF S. PARVIFLORA , AND IN VITRO SYMBIOSIS DEVELOPMENT IN S. PARVIFLORA AND SPIRANTHES SPIRALIS P IER L UIGI P ACETTI and S ABINE R IESS Dipartimento di Scienze Ambientali, Universit della Tuscia, I-01100 Viterbo, Italy A B S T R A C T . Endophytes were isolated from Serapias parviflora (Orchidaceae) roots. They are described and partially classified under microscope, after growth on PDA. Two fungi had symbiotic characters: A-Sepa-1, an ascomycete, and B-Sepa-1, a basidiomycete. At the same time, many plants of S. parviflora and Spiranthes spiralis were asymbiotically obtained by sterilisation of seeds and sowing on modified Frosch medium. After transplantation to symbiotic medium (modified Basic Oats), they were inoculated with isolated fungi. Serapias parviflora was used as control and S. spi ralis was used to establish specificity between hosts and endophytes. The two fungi were able to induce symbiosis in S. parviflora roots in vitro , while only B-Sepa-1 induced symbiosis in S. spiralis roots in vitro . We conclude that specificity between these fungi and the two orchid species studied in vitro is different, possibly substantiating the hypothesis of a potential and ecological specificity. R ESUMEN . Hongos end—fitos fueron aislados de ra’ces de Serapias parviflora (Orchidaceae). Fueron descritos y clasificados parcialmente bajo el microscopio de luz luego de ser cultivados en PDA. Dos de ellos presentaron caracter’sticas de simbiontes: la Cepa-A-1, un ascomicete, y la Cepa-B-1, un basidiomicete. Al mismo tiempo muchas plantas de S. parviflora y Spiranthes spiralis f u e r o n obtenidas asimbi—ticamente por esterilizaci—n de semillas y cultivo en un medio Frosch modificado. Luego de transplantarlas a un medio simbi—tico (Oats modificado), las plantas fueron inoculadas con los hongos aislados. Serapias parviflora fue utilizada como control y S. spiralis lo fue para estable cer especificidad entre hospederos y end—fitos. Los dos hongos fueron capaces de inducir simbiosis in vitro en ra’ces de S. parviflora , mientras que s—lo la Cepa-B-1 indujo simbiosis in vitro en ra’ces de S. spiralis . Concluimos que la especificidad in vitro entre estos hongos y las dos especies de orqu’deas estudiadas es diferente, posiblemente apoyando la hip—tesis de una especificidad ecol—gi ca potencial. K E Y W O R D S : Orchidaceae, Serapias parviflora , Spiranthes spiralis , symbiotic fungi, endophytes, Italy.

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and Scrugli (1987) suggested to classify orchid endophytes in four morphometrical classes: A, B, C, and D. This classification has no taxonomic value, but because fungi are studied when they are into cortical cells, it gives information on fungal ecology. Three ascomycetes strains with R h i z o c t o n i a -like anamorphs were isolated from Pterostylis sp. but no seeds germinated when they were inoculated with these fungi (Warcup 1975). Some tropical orchids have ascomycetes as symbiont (Dreifuss & Petrini 1981 and 1984); formerly these fungi were described as A s c o r h i z o c t o n i a Chin S. Yang & Korf, but since they are saprobe fungi and do not produce sclerotia they can not be ascribed to the genus R h i z o c t o n i a DC. (Andersen 1986). Septum ultrastructure of Leptodontidium orchidicola Sigler & Currah (Mitosporic Fungi) isolated from tropical orchids reveals that this anamorph is related to ascomycetes in having Woronin bodies near its septum pore. One of the first questions about the relationship between orchids and endophytes deals with the specificity. Bernard (1909) hypothesised a high level of specificity but such close relationship was soon rejected. Burgeff (1936) proposed that specificity existed between some fungi and ecological host groups, and Curtis (1937) suggested a closer relationship between fungus and habitat. Hadley and Harvais (1967) questioned Curtis’ ecological specificity because not all fungi isolated from ripe plants were able to support host seed germination. Further works built up evidence in favour of the absence of specificity (Downie 1959, Hadley 1970). Riess and Scrugli (1987) observed that some orchid species ( Ophrys bombyliflora Link. and Ophrys tenthredinifera Willd.) had different endophytes when collected in different sites. They also observed, in the same work, that in Limodorum abortivum (L.) Sw. there were simul taneously two endophytes with different morphological characters. Similar results were obtained from Curtis (1937), Downie (1943), Talbot and Warcup (1967), and Harley (1969). Masuhara and Katsuya (1989, 1994), by studying S p i r a n t h e s s i n e n s i s (Persoon) Ames var. a m o e n a ( M . Bieberstein) Hara, suggested two kinds of specifici ty: 1) “ecological specificity”, i.e. when pelotons are into root cortical cells or into the protocorms in nature ( in situ ); and 2) “potential specificity”, i.e. associations between orchids and fungi in other conditions, both in vitro or ex vitro . Masuhara and others (1993) observed ecological specificity only in some fungi with potential specificity. For example, Microtis parviflora R. Br. (Orchidaceae) has a narrow ecological specificity in the field, while showing a broad potential specificity in vitro. The factors that contribute to ecological specificity could be fungal growth and survival in the soil, which are influenced by environmental factors, or fungal density in the field (Masuhara and others 1995). Milligan and Williams (1988; in Masuhara & Katsuya 1995) suggested that differences between ecological and potential specificity could be due to a succession of fungi in orchid tis sues, but further investigations are necessary to confirm this hypothesis. Our study was carried out in two steps: 1) isola tion and description of Serapias parviflora endomycorrhizal fungi; 2) description of associations, in vitro, between S. parviflora and fungi, and between Spiranthes spiralis and fungi. S. parviflora was used as control and S. spiralis was used to ver ify the existence of specificity between host and endophyte. Materials and methods . Serapias parviflora roots samples were collected at Allerona Scalo (Umbria, Italy) on 05/16/1999. The whole plant, together with a clump of soil, was collected, in order to prevent root damage. Samples were pre served in sterilised envelopes at 5 C until fungal isolation (two days after collection). To remove fungi and bacteria from external surfaces, roots were sterilised by immersion on H 2 O 2 (30%) for 4 minutes; then they were rinsed 4 times in sterile distilled water. Roots were cut, 1 cm segments were sowed on Petri dishes with PDA and strepto mycin (a broad-spectrum antibiotic) and then incu bated at 20 C. Fungi were observed under phase contrast microscope, scanning and transmission electron microscope and confocal laser microscope. For the latter there is no need of a particular prepa ration but fungi stained better when coloured with acid fuchsin. Fungi observed under TEM were LANKESTERIANA 24 N 2

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treated as follows: growth on liquid medium, fixa tion for 2 hours in para-formaldehyde/glutaraldehyde (2%/2.5% v/v) in phosphate buffer 0,025 M pH 7 at 25 C, rinsed 3 times in phosphate buffer 0,025 M, postfixation in OsO4 1% in 0,05 M phos phate buffer v/v for 12 hours at 4 C, dehydration in a graded ethyl alcohol series for 15 minutes each (10%, 30%, 50%, 75%) and for 1 hour in pure ethyl alcohol, embedding on Spurr’s epoxy resin and polymerisation for 8 hours at 70 C. Thin sections (70 nm), stained with uranil acetate, 10% v/v in 50% ethanol (v/v) for 7 minutes at 70 C and fresh ly prepared lead citrate (1% v/v on 50% ethanol v/v) for 12 minutes at 25 C were observed. Fungi were inoculated in S. parviflora plants to confirm symbiosis and in Spiranthes spiralis plants to establish specificity. All plants were asymbiotically grown from seeds (modified Frosch medium) and transplanted on medium for symbiot ic growth (modified Basic Oats medium, Riess and Pacetti, 2001) before inoculation. Results . Many fungi were isolated from roots of Serapias parviflora but only two strains had char acters of symbiotic fungi: mycelium with septa, monilioid cells, sclerotia, without asexual spores (grown on PDA until substrate exhaustion). Two fungi were able to form, in vitro, typical pelotons in S. parviflora and Spiranthes spiralis root cortex cells. They were called B-Sepa-1 and A-Sepa-1. B-Sepa-1 (PDA) produces floccose to velutinous colonies, without water-soluble pigments and substrate pigmentation. Mycelium superficial and from white to light grey. On PDA, monilioid cells (fig. 1) and sclerotia (fig. 2) are differentiated. Vegetative hyphae hyaline, septated, with constric tion on branched point and with smooth wall (fig. 3). Thin wall and hyaline monilioid cells, from ellipsoidal to spherical, 4,5 x 6,5 mm, organised in septated chains and branched or linear chains (fig. 4). Sclerotia torulose, 120 x 70 mm. Dolipores always present under TEM (fig. 5). A-Sepa-1 (PDA) consisting of floccose and from light yellow to yellow colonies with white to light grey micelium on peripheral area. Uncoloured exu date, from dark grey to black water-soluble pigments and monilioid cells are produced. Vegetative hyphae hyaline, septated, with smooth walls. Hyaline monilioid cells are differentiated. Spherical to irregular, 4,5-15 x 6,5-15 mm, organised in lin ear and never branched chains. Thick and irregular electron dense layer around hyphae (fig. 6), Woronin bodies near septum pore (fig. 7). Symbiosis between S. parviflora and B-Sepa-1 was observed under microscope. In these associations there is a massif fungal penetration of roots, preferentially through hairs (fig. 8) but in some case there is epidermal penetrations too. Subepidermal invasion is confined to the first two cortex layers. Pelotons and digested pelotons can be observed from the third to the eighth cellular layer. In some sections we can find both pelotons (fig. 9) and digested pelotons (fig. 10) in the same cell. We can also assume an infective cyclical pat tern by observing mycelium intercellular connec tions (fig. 11). P ACETTI and R IESS Endophytes of Serapias parviflora and Spiranthes spiralis Octubre 2001 25 Figure 1. B-Sepa-1 monilioid cells (SEM, x 2000). Figure 2. B-Sepa-1 sclerotium (SEM, x 1000). 

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In S. parviflora roots, inoculated with A-Sepa-1, there are few observations of epidermal penetrations (fig. 12). There is a slight subepidermal inva sion and there are pelotons (fig. 13) and digested pelotons in the inner layers of the cortex (from the third to the sixth). Pelotons occupy a smaller cyto plasm portion compared to B-Sepa-1 in the same orchid. As to S. spiralis inoculated with B-Sepa-1, it is possible to say that fungus preferentially penetrates through hairs (fig. 14), even if epidermal penetration is quite frequent (fig. 15). It is impossible to see subepidermal invasion because the first five layers are completely invaded by pelotons and digested pelotons (fig. 16 and 17). In this case, intercellular micelium connections are very clear. Beyond the fifth cellular layer of the cortex there is no fungus but there are many starch granules (fig. 18). In S. spiralis , inoculated with A-Sepa-1, there are only very few partially or completely digested pelotons. Epidermal invasion is the only means observed for fungal penetration; hairs root invasion is never observed. Fungal hyphae diameters are quite constant both LANKESTERIANA 26 N 2 Figure 3. Trasversal section of B-Sepa-1 hypha (TEM, x 22500). Figure 4. B-Sepa-1 monilioid cells (confocal laser microscope, x 80). Figure 5. B-Sepa-1 septal ultrastructure with dolipore (TEM, x 45000). Figure 6. Trasversal section of A-Sepa-1 hypha, with a thick and irregular electron-dense layer sorrounding the cellular wall (TEM, x 30000).

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in pure cultures on PDA and in the hosts: in BSepa-1 diameters are on average 2 mm, while in A-Sepa-1 diameters are on average 3,5 mm. Conclusion . Endophytes isolation from roots of Serapias parviflora reveals the presence of two fungi. Our technique allows us to determine only the division. A-Sepa-1 is an ascomycetes (presence of Woronin bodies), and B-Sepa-1 is a basidiomycetes (presence of dolipores). We can also recognise Riess and Scrugli morphometrical class es: A-Sepa-1 belongs to class B, with diameters of hyphae ranging from 2,6 to 5 mm, while B-Sepa-1 belongs to class A, having hyphae with diameters less than 2,5 mm. Molecular markers could certain ly provide diagnostic tools for lower taxonomical levels and taxon fingerprints. . We can not tell if S. parviflor a needs two fungi simultaneously or in succession for its development. Only observations of endophytes in samples living in the same site could maybe answer this question. A-Sepa-1 and B-Sepa-1 behave differently when inoculated in plants of S. parviflora and Spiranthes s p i r a l i s : B-Sepa-1, the basidiomycetes, is a good symbiont for both orchids, i.e. it is always possible to see all typical mycorrhizal phases, their cyclical pattern and the absence of fungi in orchids root vascular tissue. A-Sepa-1, the ascomycetes, is a good symbiont only for S. parviflora ; in this orchid we can observe fungal confinement in the first layer of root cortex, starch accumulation in the inner layers, cyclic infections and absence of fungus in vascular tissue. A-Sepa-1 is not tolerate by S. spi r a l i s : there is a massif epidermal penetration but P ACETTI and R IESS Endophytes of Serapias parviflora and Spiranthes spiralis Octubre 2001 27 Figure 7. A-Sepa-1 septal ultrastructure, with Woronin bodies (TEM, x 45000). Figure 8. S. parviflora hair massively invaded by BSepa-1 (confocal laser microscope, x 40). Figure 9. B-Sepa-1 pelotons in S. parviflora root cor tex, stained by acid fuchsin (phase contrast microscope, x 40 ). Figure 10. B-Sepa-1 partially digested pelotons in S . parviflora roots, stained by acid fuchsin (x 40 ).

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LANKESTERIANA 28 N 2 Figure 11. B-Sepa-1 pelotons with intercellular connections, stained by acid fuchsin (confocal laser microscope, x 24). Figure 12. A-Sepa-1 epidermal penetration in S. parvi flora (phase contrast microscope, x 40). Figure 13. A-Sepa-1 pelotons in S. parviflora , stained by acid fuchsin (phase contrast microscope, x 20). Figure 14. B-Sepa-1 penetration in S. spiralis root by hair (x 40). there is no penetration through hairs. The very few pelotons are digested immediately and mycelium can not invade the other cells: typical symbiotic alternation of infection and digestion seems to be shifted toward digestion. Symbiosis between A-Sepa-1 and S. parviflora and digestion of fungus in S. spiralis suggest a dif ferent potential specificity of the fungus for the two hosts: S. parviflora is micorrhizated by the two fungi and S. spiralis only by B-Sepa-1, in vitro . If it is true, inoculation of fungi in S. parviflora and S. spiralis and their infective patterns are an addition al confirmation that there is no species-specificity, but potential or ecological specificity between orchids and fungi, as suggested from Masuhara and Katsuya (1989). 

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The different pattern of infection of the two fungi in S. spiralis could also suggest that there is a fungal succession in this host, and that A-Sepa-1 can not induce symbiosis in S. spiralis in the phase of development studied. L ITERATURE C ITED Andersen, T. F. 1996. A comparative taxonomic study of Rhizoctonia sensu lato employing morphological, ultra structural and molecular methods. Mycol. Res. 100: 1117-1128. Bernard, N. 1909. L’evolution dans la symbiose. Ann. Sci. Nat. Bot. 9, ser. 9: 1-196. Burgeff, H. 1936. Samenkeimung der Orchideen. G. Fischer, Jena. Curtis, J.T. 1937. The relation of specificity of Orchid mycorrhizal fungi to the problem of symbiosis. Amer. J. Bot. 26: 390-398. Downie, D.G. 1943. Notes on the germination of Corallorhiza innata . Trans. Bot. Soc. Edinburgh 33: 380-392. _____. 1959. The mycorrhiza of Orchis purpurella . Trans. Bot. Soc. Edinburgh 38: 16-29. Dreyfuss, M. & O. Petrini. 1981. Endophytische Pilze in Epiphytischen Araceae, Bromeliaceae und Orchidaceae. Sydowia 34: 135-148. P ACETTI and R IESS Endophytes of Serapias parviflora and Spiranthes spiralis Octubre 2001 29 Figure 15. S. spiralis epidermal root cell penetrated by B-Sepa-1 (SEM, x 1,000). Figure 16. B-Sepa-1 pelotons in S. spiralis root cells (confocal laser microscope, x 40). Figure 17. B-Sepa-1 pelotons in a S. spiralis root sec tion (SEM, x 500). Figure 18. S. spiralis root section with B-Sepa-1 inva sion area (confocal laser microscope).

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LANKESTERIANA _____. 1984. Further investigations on the occurrence and distribution of endophytic fungi in tropical plants. Bot. Helvetica 94: 33-40. Hadley, G. 1970. Non specificity of symbiotic infection in orchid micorrhiza. New Phytol. 69: 1015-1023. Hadley G. & G. Harvais. 1967. The relation between host and endophyte in orchid micorrhiza. New Phytol. 66: 205-215. Harley, J.L. 1969. The biology of micorrhiza. 2nd ed., Leonard Hill Books, London. Masuhara, G. & K. Katsuya. 1989. Effects of mycorrhizal fungi on seed germination and early growth of three japanese terrestrial orchids. Sci. Hort. 37 : 331-337. Masuhara, G., K. Katsuya & K. Yamaguchi. 1993. Potential for symbiosis of Rhizoctonia solani and binu cleate Rhizoctonia with seeds of Spiranthes sinensis var. amoena (Orchidaceae) in vitro . Mycol. Res. 97: 746-752. Masuhara, G. & K. Katsuya. 1994. In situ and in vitro specificity between Rhizoctonia spp. and S p i r a n t h e s sinensis (Persoon) Ames. var. amoena (M. Bieberstein) Hara (Orchidaceae). New Phytol. 127: 711-718. Masuhara, G., P.A. McGee & A.J. Perkins. 1995. Specificity of the associations between Microtis parvi flora (Orchidaceae) and its mycorrhizal fungi. Austral. J. Bot. 43: 85-91. Rambelli A. 1981. Fondamenti di micologia. Edizioni Zanichelli, Bologna. Riess, S. & A. Scrugli. 1987. Associazioni micorriziche nelle orchidee spontanee della Sardegna. Micol. Ital. XVI (3): 21-28. Riess, S, A. Cogoni & A. Scrugli. 1992. Rilevamenti mor fometrici sugli endofiti di 38 Orchidaceae spontanee della Sardegna. Micol. Ital. XXI (2): 21-28. Riess, S. & P.L. Pacetti. 2001. Germinazione e sviluppo in vitro, simbiotico ed asimbiotico, di alcune specie di orchidee spontanee italiane. Caesiana 15: 25-32. Talbot, P.H.B. & J.H. Warcup. 1967. Perfect states of Rhizoctonias associated whith Orchids. New Phytol. 66: 631-641. Warcup, J.H. 1975. Factors affecting symbiotic germination of orchid seed. In Sanders F.E., Mosse B., Tinker, P.B (eds.). Endomycorrhiza, p. 85-104. London. 30 N 2