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Lankesteriana

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Title:
Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Physical Description:
v. : ill. (some col.) ; 25 cm.
Language:
English
Creator:
Jardín Botánico Lankester
Publisher:
Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Place of Publication:
Cartago Costa Rica
Cartago, Costa Rica
Publication Date:
Frequency:
three times a year[2002-]
irregular[ former 2001]
three times a year
regular

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Subjects / Keywords:
Botany -- Periodicals -- Costa Rica   ( lcsh )
Epiphytes -- Periodicals -- Costa Rica   ( lcsh )
Orchids -- Periodicals -- Costa Rica   ( lcsh )
Plantkunde   ( gtt )
Botanische tuinen   ( gtt )
Genre:
periodical   ( marcgt )
serial   ( sobekcm )
Spatial Coverage:
Costa Rica

Notes

Language:
In English and Spanish.
Dates or Sequential Designation:
No. 1 (mayo 2001)-
Numbering Peculiarities:
Issues for May 2001-Oct. 2003 designated no.1-8; issues for Apr. 2004- designated vol. 4, no. 1-
General Note:
Latest issue consulted: Vol. 4, no. 1 (abr. 2004).
General Note:
International journal on orchidology.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
All applicable rights reserved by the source institution and holding location.
Resource Identifier:
oclc - 48491453
lccn - 2001240973
issn - 1409-3871
System ID:
UF00098723:00030

MISSING IMAGE

Material Information

Title:
Lankesteriana la revista científica del Jardín Botánico Lankester, Universidad de Costa Rica
Physical Description:
v. : ill. (some col.) ; 25 cm.
Language:
English
Creator:
Jardín Botánico Lankester
Publisher:
Jardi´n Bota´nico Lankester, Universidad de Costa Rica
Jardín Botánico Lankester, Universidad de Costa Rica
Place of Publication:
Cartago Costa Rica
Cartago, Costa Rica
Publication Date:
Frequency:
three times a year[2002-]
irregular[ former 2001]
three times a year
regular

Subjects

Subjects / Keywords:
Botany -- Periodicals -- Costa Rica   ( lcsh )
Epiphytes -- Periodicals -- Costa Rica   ( lcsh )
Orchids -- Periodicals -- Costa Rica   ( lcsh )
Plantkunde   ( gtt )
Botanische tuinen   ( gtt )
Genre:
periodical   ( marcgt )
serial   ( sobekcm )
Spatial Coverage:
Costa Rica

Notes

Language:
In English and Spanish.
Dates or Sequential Designation:
No. 1 (mayo 2001)-
Numbering Peculiarities:
Issues for May 2001-Oct. 2003 designated no.1-8; issues for Apr. 2004- designated vol. 4, no. 1-
General Note:
Latest issue consulted: Vol. 4, no. 1 (abr. 2004).
General Note:
International journal on orchidology.

Record Information

Source Institution:
University of Florida
Holding Location:
University of Florida
Rights Management:
All applicable rights reserved by the source institution and holding location.
Resource Identifier:
oclc - 48491453
lccn - 2001240973
issn - 1409-3871
System ID:
UF00098723:00030


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INTERNATIONAL JOURNAL ON ORCHIDOLOGYISSN 1409-3871VOL. 12, No. 1 APRIL 2012 Encylia nizanburyi (Orchidaceae), un nuevo hbrido natural del istmo de Tehuantepec, Mxico EDUARDO A. PREZ-GARCA and ERIC HGSATER A new species of Campylocentrum (Orchidaceae: Angraecinae) from Colombia MARTA KOLANOWSKA, OSCAR ALEJANDRO PREZ ESCOBAR and EDICSON PARRA SNCHEZ On the relationship between bryophyte cover and the distribution of Lepanthes spp. BENJAMIN J. CRAIN New species and records of Orchidaceae from Costa Rica. II ADAM P. KARREMANS, DIEGO BOGARN, MELANIA FERNNDEZ, CHRISTINA M. SMITH and MARIO A. BLANCO New combinations in Odontoglossum (Orchidaceae: Oncidiinae) and a solution to a taxonomic conundrum STIG DALSTRM Morfoanatoma en Cranichideae (Orchidaceae) de la Estacin Loma Redonda del Parque Nacional Sierra Nevada, Mrida, Venezuela BLANCA A. DUGARTE CORREDOR & REBECA LUQUE ARIAS 1 9 13 19 53 61

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The Vice-Presidency of Research UNIVERSITY OF COSTA RICA is sincerely acknowledged for his support to the printing of this volume

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INTERNATIONAL JOURNAL ON ORCHIDOLOGY Copyright 2011 Lankester Botanical Garden, University of Costa Rica Effective publication date: April 26, 2012 Layout: Jardn Botnico Lankester. Cover: Acianthera oscitans (Ames) Pridgeon & M.W.Chase. Photograph by F. Pupulin. Printer: Masterlitho Printed copies: 500 Printed in Costa Rica / Impreso en Costa RicaR Lankesterian a / International Journal on Orchidology No. 1 (2001)-. -San Jos, Costa Rica: Editorial Universidad de Costa Rica, 2001-v. ISSN-1409-3871 1. Botnica Publicaciones peridicas, 2. Publicaciones peridicas costarricenses

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Visit the new webpage at www.lankesteriana.org Originally devoted to the publication of articles on general botany, with special attention to epiphytic plants and orchid systematics, ecology, evolution and physiology, along with book reviews and conferences on these subjects, since 2007 LAN KE ST E RIANA focused exclusively on scientific papers on orchidology. LANKESTERIANA is a peer-reviewed journal that publishes original works in English and occasionally in Spanish, and it is distributed to more than 350 libraries and institutions worldwide. In order to increase visibility of the articles published in LANKESTERIANA the journal maintains since 2009 a web page with downloadable contents. With November, 2011, we present to our readers a new interface of the journal at www.lankesteriana.org Please bookmark the new address of the webpage, which substitutes the previous address hosted at the internal server of ucr.ac.cr. Readers can now browse through all the past issues of LAN KE ST E RIANA including the currrent issue, and download them as complete fascicles or, via the Index to the single issues, only the articles of their interest. According to the Open Access policy promoted by the University of Costa Rica, all the publications supported by the University are licensed under the Creative Commons copyright. Downloading LANKESTERIANA is completely free. At the home page of LANKESTERIANA words or any other word which should appear in the text you are looking for. We take the opportunity to acknowledge our authors, reviewers and readers, who help us making a better The editors

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The Global Orchid Taxonomic Network at a click www.epidendra.orgNow with a new user interface, the online database on taxonomic information by Lankester Botanical Gar den includes more than 7,000 orchid names, completely cross-referenced and with evaluated synonymies. free, immediately downloadable protologues, type images, illustrations of the original materials, historical and modern illustrations, photographs, pertinent literature and, when available, digital images of species pollinaria. An index (under the button List of species) allows the users to search for any published name, independently if it is accepted or not by the taxonomic compilers. Synonyms are linked to their accepted name, where additional materials (including images) are available for download. Hundredscations and other materials relative to orchid systematics, distribution and history are added to the database on a monthly basis (new entries can be searched by clicking on the New records button). Since March, 2012, new pages are devoted to the orchid species recorded in the rich system of national parks Central American countries (Floras button) and to interesting aspects of orchid history. Under the button Collectable plates, the research staff at Lankester Botanical Garden makes available to the public the most detailed images of orchids from the collections at the Center, organized in a series of collectable plates that can be downloaded for free. New ones are added each week. Supported by the University of Costa Rica and the Darwin Initiative, EPIDENDRA The Global Orchid Taxonomic Network counts with the collaboration of respected taxonomists and leading botanical institutions worldwide.

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LANKESTERIANA 12(1): 1. 2012.ENCYCLIA NIZANBURYI (ORCHIDACEAE) UN NUEVO HBRIDO NATURAL DEL ISTMO DE TEHUANTEPEC, MXICO EDUARDO A. PREZ-GARCA 1,3 & ERIC HGSATER 21 Departamento de Ecologa y Recursos Naturales, Facultad de Ciencias, Universidad Nacional Autnoma de Mxico. Ciudad Universitaria, Del. Coyoacn, 04510, Mxico, D.F., Mxico 2 Herbario AMO, Montaas Calizas # 490. Lomas de Chapultepec, Del. Miguel Hidalgo, 11000, Mxico, D.F., Mxico3 Autor para la correspondencia: eduardo.perez-garcia@ciencias.unam.mx RESUMEN Se describe e ilustra Encyclia nizanburyi, una notoespecie nueva la cual slo se conoce de la discute el posible origen hbrido de este nuevo taxn y se le compara con sus progenitores hipotticos: Encyclia nizandensis y Encyclia hanburyi ABSTRACT. Encyclia nizanburyi watershed of the Tehuantepec isthmus, state Oaxaca, Mexico, is described and illustrated. The possible hybrid origin of the new taxon is discussed, and it is compared with its putative parents: Encyclia nizandensis and Encyclia hanburyi PALABRAS CLAVE / KEY WORDS : Encyclia x nizanburyi, hbrido natural, natural hybrid, Nizanda, Oaxaca, orqudea, orchidIntroduccin. Durante las exploraciones botnicas que se han realizado desde hace ms de 15 aos en la regin de Nizanda, Oaxaca (Fig. 1), se encontraron unas plantas que presentaban diferencias claras con las otras especies conocidas del gnero Encyclia (Orchidaceae), en la regin. Al realizar un estudio ms detallado se encontr que estas plantas no correspondan a ninguna especie conocida, y que al parecer eran el resultado de la hibridacin natural entre dos especies de ese gnero, E. nizandensis Prez-Garca & Hgsater y E. hanburyi (Lindl.) Schltr., por este motivo se propone este nuevo taxn. Encyclia nizanburyi Prez-Garca & Hgsater, hyb. nat. nov. (Fig. 2). Hybrida ex Encyclia hanburyi (Lindl.) Schltr. et E. nizandensi Prez-Garca & Hgsater exorta, ad illam ovario verrucoso, ad hanc pseudobulbis fusiformibus et foliis angustis longisque accedens. Hierba Races carnosas de 2 mm de grosor. Seudobulbos agrupados, cnicos-ovoides, 40 mm, envueltos por vainas escariosas. Hojas 2, apicales, linear-lanceoladas, acanaladas, arqueadas, obtusas, coriceas, de 110 9 mm. apical, erecta, racemosa o paniculada, 25 cm de largo, ramas cortas hasta 5 cm, brcteas del pednculo tipo vaina, envolventes de 9 mm de largo. Ovario subterete, dilatado hacia el pice, sulcado, ligeramente verrugoso, de 23.6 mm de largo 2 mm de ancho. triangulares, pequeas, < 4 mm. Flores 3 sucesivas, de 37 mm de dimetro natural, sin aroma perceptible. Flores con spalos y ptalos de color caf chocolate-rojizo, las venas ligeramente ms oscuras; labelo rosado con rayas magenta, el callo un poco ms claro de color, columna rosa-blanquecina con reas ms pigmentadas hacia el pice, antera blanco crema. Spalos extendidos, arqueados hacia el frente, oblanceolados, el dorsal es casi perpendicular a los laterales, estos ltimos se ubican a la altura del lbulo medio del labelo, pices agudos, de 19.0 6.2 mm, 5 nervados pero las Ptalos unguiculados, arqueados (doblados) hacia en frente, de 19 6 mm, 3 nervados, las dos venas de los lados se bifurcan y luego

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Labelo, unido en la base a la columna, trilobado, 21.5 15.0 mm; lbulos laterales oblongo obovados, redondeados (similares a E. hanburyi), 14.0 6.2 mm, lbulo medio reniforme, emarginado, margen ondulado, 10.7 13.4 mm, el istmo de 4 mm, callo carnoso, surcado, terminado en una quilla. Columna recta de 11.0 4.1 mm, sin alas, pice romo, con tres pequeos apendices que no sobrepasan a la antera. Antera reniforme, tetralocular, carnosa. Polinios 4 obovoides, lateralmente comprimidos, 1.5 mm de largo. Cpsula no vista. TIPO : Mxico, Oaxaca: Municipio de Ciudad Ixtepec, Distrito de Juchitn. Montculo de caliza a 3.65 km al noroeste (319) del poblado de Nizanda, Asuncin caliza. Colectada el 4 de febrero de 1996. Prensado de material cultivado el 26 de abril de 2010. Eduardo A. Prez-Garca #982, Marco A. Romero y Bartolino Reyes Ros (holotipo: AMO!; Illustration voucher). OTROS EJEMPLARES : MEXICO: Oaxaca: Nizanda, Municipio de Asuncin Ixtaltepec, Distrito Juchitn, E.A. Prez-Garca #981 y Bartolino Reyes (MEXU!). RECONOCIMIENTO Encyclia nizanburyi tiene caractersticas intermedias entre E. nizandensis y E. hanburyi. de E. nizanburyi se parecen a E. hanburyi (Fig. 3), mientras que las plantas (la parte vegetativa) se parecen ms a E. nizandensis E. nizanburyi son de color ligeramente ms claro y pequeas que las E. hanburyi tpicas (Fig. 4). Otra diferencia notable es que E. nizanburyi tiene el ovario verrucoso y no liso como E. hanburyi. En la regin de Nizanda, E. hanburyi tiene pseudobulbos muy redondeados (globosos) y hojas linear-lanceoladas, acanaladas, mientras que E. nizandensis tiene pseudobulbos fusiformes y sus hojas son muy variables en tamao y forma, pero lo ms comn es que sean hojas delgadas y largas (acintadas). de las principales diferencias entre E. nizandensis y E. nizanburyi, ya que en la primera los tpalos son caf pardo y el labelo es amarillo-crema, mientras que en la segunda los tpalos son caf chocolate claro y el labelo es rosado. Otra diferencia notoria son los lbulos laterales del labelo, ya que en E. nizanburyi son largos, anchos y envuelven a la columna, mientras que en E. nizandensis son cortos y se presentan de manera casi horizontal al lbulo medio. DISTRIBUCIN Y ECOLOGA Encyclia nizanburyi slo se conoce de la regin de Nizanda, Oaxaca, en la Vertiente Meave, 2004). Esta caliza presenta estratos gruesos y estn ligeramente plegadas. En esas calizas se han restos fsiles de rudistas y corales recristalizados, que sugieren una edad del Albiano-Cenomaniano (Cretcico superior; Prez-Gutirrez et al. 2009). Encyclia nizanburyi se encontr creciendo sobre la roca expuesta, en la cima de un cerro de baja altitud (entre 200 y 300 m s.n.m). La vegetacin presente es pues stos se encuentran inmersos en una matriz de selva baja caducifolia (Prez-Garca & Meave, 2004; PrezGarca et al. 2001, 2009). Las plantas dominantes en Agave ghiesbreghtii Lem. ex Jacobi, Bursera excelsa (Kunth) Engl., B. schlechtendalii Engl., Cephalocereus nizandensis (Bravo & T.MacDoug.) Buxb., Comocladia engleriana Loes., Neobuxbaumia scoparia (Poselg.) Backeb., LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.2 LANKESTERIANA FIGURA 1. Ubicacin de la localidad de Nizanda, en la porcin sur del Istmo de Tehuantepec, Oaxaca, Mxico.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. PREZ-GARCA & HGSATER Encyclia nizanburyi3 FIGURA 2. Ilustracin de Encyclia nizanburyi realizada por Rolando Jimnez Machorro (Herbario AMO, Mxico, D.F.), basada en el ejemplar tipo (EAPG #982). Se indica la escala en cada seccin del dibujo.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.4 LANKESTERIANAPlumeria rubra L. f. acutifolia (Poir.) Woodson, Pseudosmodingium multifolium Rose, acompaadas de algunas otras como Mammillaria albilanata Backeb., M. voburnensis Scheer var. collinsii (Britton & Rose) Repp., Hechtia caudata L.B.Sm. y H. rosea E.Morren, y adems estn presentes algunas orqudeas como Barkeria whartoniana (C.Schweinf.) Soto Arenas, Cohniella brachyphylla (Lindl.) Cetzal & Carnevali, Cyrtopodium macrobulbum (La Llave et Lex.) G.A.Romero & Carnevali, Mesadenus lucayanus (Britton) Schltr., Encyclia hanburyi (Lindl.) Schltr. y E. nizandensis E.A.Prez-Garca & Hgsater (PrezGarca et al. 2010). La temperatura promedio anual en la regin de Nizanda es relativamente alta (26 C) y la precipitacin pluvial es muy estacional, pues prcticamente toda la precipitacin anual (ca. de 1,000 mm) caen en los meses de verano (de mayo a octubre). Sin embargo, la regin es frecuentemente impactada por vientos fuertes, por un efecto de encaonamiento del los vientos alisios. Los vientos son particularmente fuertes en el otoo y durante el invierno. El posible efecto refrescante del viento puede ser un agente que permita que E. hanburyi se localice a una altitud relativamente baja para esa especie, de la misma manera que algunas otras especies lo hacen en la regin (Prez-Garca et al. E. nizanburyi es en privamera, en otoo. En la regin de Nizanda, tanto Encyclia hanburyi como E. nizandensis temporada, por lo cual es posible que hayan podido hibridar Hibridacin natural en Enyclia Uno de los aspectos ms interesantes que tiene este nuevo taxn es la posibilidad de que sea un hbrido natural. La hibridacin natural es frecuente en orqudeas (Nielsen, 2000, Cozzolino et al. 2005, Azevedo et al. 2006) y numerosos hbridos naturales han sido publicados (por ejemplo, Romero & Carnevali, 1989). Estos hbridos naturales pueden aparecer como casos aislados o llegar a formar grandes poblaciones. Para algunas orqudeas se ha propuesto que existen barreras genticas que evitan que los hbridos sean exitosos (Moccia et al. 2077), pero es frecuente que los hbridos sean frtiles y se puedan cruzar de nuevo con sus progenitores o entre ellos, generando abanicos de formas intermedias. Quiz el caso de hibridacin natural ms conocido por los cultivadores mexicanos es Guarianthe guatemalensis (T. Moore) W.E. Higgins [= G. skinerii (Bateman) Dressler & W.E. Higgins con G. aurantiaca (Bateman ex Lindl.) Dressler & W.E. Higgins] de Chiapas y Guatemala. Particularmente dentro del gnero Encyclia la hibridacin parece ser algo comn (Dressler y Pollard, 1974), e incluso en algunas localidades puede ser algo difcil encontrar una especie en estado puro, como ha sido mencionado para algunas especies Encyclia en Las Antillas (Sauleda et al., en lnea). Particularmente, para Encyclia phoenicea se ha encontrado una gran FIGURA algunas especies Encyclia para su comparacin. A Encyclia hanburyi (del estado de Oaxaca). B Encyclia nizandensis (Oaxaca). C Encyclia nizanburyi (Oaxaca). D (Oaxaca). E Encyclia adenocarpa (Guerrero).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. PREZ-GARCA & HGSATER Encyclia nizanburyi5variacin fenotpica (Llamacho & Larramendi, 2005), y se ha discutido su posible hibridacin frecuente con E. plicata (Sauleda et al., en lnea). Hasta la fecha, se naturales en Encyclia y uno intergenrico [ Encyclaelia = Encyclia Laelia (ahora Cattleya)], adems existen al menos cuatro hbridos naturales ms sin describir formalmente (Apndice I). Entre las particularidades de la hibridacin que pueden ser relevantes para E. nizanburyi se ha encontrado que E. hanburyi puede hibridar con E. selligera (Bateman ex Lindl.) Schltr., Encyclia cf. spatella (Rchb. f.) Schltr. (Soto, 2003) y con E. ceratistes (Lindl.) Schltr. (Dressler & Pollard, 1974). Por otro lado, se ha propuesto que E. nizandensis sea una especie de origen hbrido (Prez-Garca & Hgsater, 2003), o que al menos una parte importante de sus individuos tengan genes de dos especies. Encyclia nizandensis se ubica dentro del complejo de especies de E. adenocarpa (La Llave & Lex.) Schltr. En la regin sur-este del estado de Oaxaca no se han encontrado individuos de E. adenocarpa y la especie del grupo ms prxima a la regin de Nizanda, E. papillosa (Batem.) Aguirre-Olavarrieta, recolectada cerca de Salina Cruz en costa oaxaquea. No obstante, existen varias diferencias notables entre E. nizandensis y E. papillosa FIGURA 4. Fotocomposin donde se muestran las plantas de algunas especies Encyclia para su comparacin. A Encyclia hanburyi (del estado de Oaxaca). B Encyclia nizanburyi (detalle de la planta, Oaxaca). C Encyclia nizandensis (Oaxaca). D Encyclia nizanburyi (planta completa, Oaxaca). E (Oaxaca). F Encyclia adenocarpa (Guerrero).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.6 LANKESTERIANA APNDICE 1 Lista de algunos hbridos naturales del gnero Encylia Hook, obtenidos de diversas fuentes, entre ellas las bases de datos en lnea del Missouri Botanical Garden (www.tropicos.org) y del International Plant Names Index (www.ipni. org). Hbridos intragenricos descritos Encyclia alcardoi V.P.Castro & Chiron (2002) = E. argentinensis Richardiana 2(4): 154. Encyclia bajamarensis Sauleda & R.M.Adams (1981) = E. gracilis E. rufa, Brittonia 33(2): 189. Encyclia camagueyensis Rodr.Seijo, Gonz.Estv., Sauleda, Risco Vill. & Esperon (2010) = E. altissima E. phoenicea Orchid Digest 74(2): 58. Encyclia carbonitensis Campacci (2008) = Encyclia kundergraberi E. patens, Colet. Orqudeas Brasil. 6: 194. Encyclia fabianae B.P. Faria, A.D. Santana & Pres Junior (2007) = E. santanae, Orquidrio 21(4): 140. Encyclia guzinskii Sauleda & R.M.Adams (1990) = E. altissima E. plicata Brittonia 42(4): 288. Encyclia hillyerorum Sauleda & R.M.Adams (1990) = E. fehlingii E. fucata, Brittonia 42(4): 288 Encyclia knowlesii Sauleda & R.M.Adams (1990) = E. fehlingii E. plicata Brittonia 42(4): 288. Encyclia lleidae Sauleda & R.M.Adams, (1984) = E. gracilis E. plicata Brittonia 36(3): 257. Encyclia lucayana Sauleda & R.M.Adams (1981) = E. fahlingii E. gracilis Brittonia 33(2): 190. Encyclia perplexa (Ames, F.T. Hubb. & C. Schweinf.) Dressler & G.E. Pollard (1971) = E. bractescens E. candollei Phytologia 21(7): 437. Encyclia profusa (Rolfe) Dressler & G.E. Pollard (1971) = E. ambigua E. ceratistes Phytologia 21(7): 437. Encyclia raganii Sauleda & R.M.Adams (1984) = E. altissima E. gracilis Brittonia 36(3): 258. Encyclia verboonenii V.P.Castro & Campacci (2001) = E. argentinensis E. osmantha, Orquidrio 15(1): 18. Hbridos intragenricos aparentemente no descritos Encyclia hanburyi (Lindl.) Schltr. Encyclia cf. spatella (Rchb. f.) Schltr. (Dressler & Pollard 1974; Soto 2003). E. hanburyi (Lindl.) Schltr. E. selligera (Bateman ex Lindl.) Schltr. (Dressler & Pollard 1974) E. hanburyi (Lindl.) Schltr. E. ceratistes (Lindl.) Schltr. (Dressler & Pollard, 1974). E. phoenicea (Lindl.) Neumann E. plicata (Lindl.) Britton et Millsp. (Sauleda et al. en lnea), el hbrido no natural fue registrado como Encyclia Pinar Del Rio en 1966 (The International Orchid Register). Hbrido intergenrico Encylaelia intermedia Miranda (1991) = Encyclia seideli i x Laelia crispata Albertoa 3: 94. especie, en incluso se puede dudar que sta sea uno de sus posibles progenitores, tal como se haba propuesto originalmente (Prez-Garca & Hgsater, 2003). Encyclia nizandensis es una especie nica dentro del grupo de E. adenocarpa, pero es probable que haya tenido algunos eventos de hibridacin con E. Regel. En este sentido, existe la duda que E. nizandensis sea en verdad una especie de origen hbrido; esto se basa en que la evidencia molecular preliminar aparentemente no es congruente con la hiptesis de la hibridacin y porque no se consider la presencia de en la porcin sur del Istmo de Tehuantepec (G. Carnevali, com. pers.). Este ltimo punto es rebatible porque adems de los individuos de encontrados en la regin de Nizanda, desde antes de la publicacin de E. nizandensis en 2003 (Prez-Garca et al., 2001), recientemente hemos localizado varias plantas de en sur del Istmo de Tehuantepec, en los alrededores de Almoloya y Mazahua, ambas localidades ubicadas en el estado de Oaxaca y muy cercanas a Nizanda. Cabe mencionar que la posible presencia de genes de en las plantas del complejo de E. adenocarpa en la regin de Nizanda ya haba sido mencionada por Dressler y Pollard (1974). En la regin de Nizanda se ecuentra una gran variacin en cuanto a la morfologa vegetativa en E. nizandensis, desde plantas tpicas del complejo de E.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. PREZ-GARCA & HGSATER Encyclia nizanburyi7adenocarpa hasta plantas de bulbos ms esfricos y hojas anchas como en Por otro lado, las E. nizandensis no son muy consistentes en cuanto a su variacin con respecto al tipo de planta, la forma pura de la especie y cual puede tener alguna mezcla con En todo caso es necesario hacer estudios ms detallados, tanto moleculares como morfolgicos para poder llegar a una mejor conclusin sobre el posible papel de la presencia de genes de E. sobre E. nizandensis. De existir la introgresin gentica de en E. nizandensis, entonces E. nizanburyi podra tener involucradas tres especies en su genealoga (E. hanburyi, E. nizandensis y E. parvifolia). Hasta el momento no se han registrado hbridos que contengan a tres especies, pues en principio solo hay dos posibles padres originales. Pero algunos de los hbridos naturales podran en realidad tener ms de dos especies en su genealoga. Por ejemplo, en el caso de algunos de los hbridos de las Antillas se ha propuesto que existe un elevado grado de entrecruzamiento entre las especies, particularmente para los hbridos que contienen E. phoenicea o E. plicata en Cuba (Sauleda et al., en lnea). ESTADO DE CONSERVACIN Encyclia nizanburyi parece ser muy rara en la naturaleza, pues solamente se conocen dos clones diferentes, uno de ellos se ha conservado en cultivo durante varios aos. Los dos clones fueron colectados en la misma localidad, en el localizaron varias plantas de E. hanburyi. Cabe sealar que, a nivel ms general en la regin de Nizanda, E. hanburyi es relativamente escasa y de distribucin restringida, ya que slo se le encuentra en las partes ms altas de los cerros de caliza, sobre las peas donde la vegetacin es muy abierta. En contraste, E. nizandensis es relativamente comn en la zona, y aunque crece preferentemente en sitios con substrato calcreo puede encontrarse sobre esquistos. Esta sobre Comocladia engleriana, Beaucarnea recurvata y Neobuxbaumia scoparia. Dada la relativa frecuencia de ambas especies, y considerando que prcticamente son existe la posibilidad de que se encuentren ms clones o que esta hibridacin pueda volver a ocurrir. En general, no existe una gran presin de colecta sobre ninguna de las dos especies progenitoras, pero E. hanburyi puede ser ms vulnerable pues tiene cierto inters hortcola, sus nmeros poblaciones no son tan grandes, y su distribucin es ms restringida. Esta regin del Istmo de Tehuantepec se est transformando fuertemente por la instalacin de centrales de aerogeneradores, lo cual seguramente traer tambin un cambio en el patrn de uso de la tierra y en los patrones de consumo. De igual forma, el tendido de lneas elctricas ha generado facilitar el ingreso de factores de disturbio. Por otro lado, la industria cementera es una amenaza directa materia prima, de igual forma la extraccin de mrmol y de grava pueden poner en riesgo a este ecosistema. de la regin est actualmente bajo un rgimen de conservacin (Prez-Garca, 2006). ETIMOLOGA El nombre proviene de la combinacin de los nombres de los posibles progenitores de este hbrido natural: E. nizandensis y E. hanburyi. E. nizandensis localidad tipo de esta especie, Nizanda. Esta localidad es una pequea poblacin ubicada en la porcin sur del Istmo de Tehuantepec, Oaxaca. En lengua zapoteca, de por la presencia de manantiales de agua termales. AGRADECIMIENTOS R. Jimnez Machorro realiz la ilustracin botnica. El programa PAPIIT de la Universidad Nacional Autnoma de Mxico y el prospecciones botnicas en la regin de Nizanda. La gente de Nizanda, particularmente la familia Reyes-Manuel, han brindado numerosas facilidades durante el trabajo de campo. LITERATURA CITADA Azevedo, C.O., E.L. Borba & C. Van Den Berg. 2006. Evidence of natural hybridization and introgression in Bulbophyllum involutum Borba, Semir & F. Barros and B. weddellii (Lindl.) Rchb. f. (Orchidaceae) in the Chapada Diamantina, Brazil, by using allozyme markers. Rev. Brasil. Bot. 29: 415-421. Cozzolino, S., A.M. Nardella, S. Impagliazzo, A. Widmer & C. Lexer. 2005. Hybridization and conservation of

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.8 LANKESTERIANAMediterranean orchids: Should we protect the orchid hybrids or the orchid hybrid zones? Biol. Conserv. 129 (1): 14-23. Dressler R.L. & G.E. Pollard. 1974. El Gnero Encyclia en Mxico. Asociacin Mexicana de Orquideologa, A.C. Mxico, D.F. 158 pp. Llamacho J.A. & J.A. Larramendi. 2005. The orchids of Cuba (Las orqudeas de Cuba). Greta editores. Lleida, Espaa, 287 pp. Moccia, M.D., A. Widmer & S. Cozzolino. 2007. The strength of reproductive isolation in two hybridizing food-deceptive orchid species. Molec. Ecol. 16: 28552866 Nielsen, L.R. 2000. Natural hybridization between Vanilla claviculata (Wowright) Sw. and V. barbellata Rchb.f. (Orchidaceae): genetic, morphological, and pollination experimental data. Bot. J. Linn. Soc. 133: 285-302. Prez-Garca, E.A. 2006. Plan de Manejo de las reas Asuncin Ixtaltepec, Oaxaca. Comisin Nacional de reas Naturales Protegidas, Sociedad para el Estudio de los Recursos Biticos de Oaxaca, A.C., Universidad Nacional Autnoma de Mxico. Mxico, D.F. 71 pp. Indito. Prez-Garca, E.A. & E. Hgsater. 2003. Encyclia nizandensis Prez-Garca et Hgsater. Plate 564 in: E. Hgsater & M. Soto (Eds.). Icones Orchidacearum (Mexico) Fascicles 5 & 6, Orchids of Mexico, Parts 2 & 3. Herbario AMO, Mxico D.F. Prez-Garca, E.A. & J.A. Meave. 2004. Heterogeneity of xerophytic vegetation of limestone outcrops in a tropical deciduous forest region. Pl. Ecol. 175: 147-163. Prez-Garca, E.A., J. Meave y C. Gallardo. 2001. Tehuantepec, Oaxaca. Acta Bot. Mex. 56: 19-88. Prez-Garca, E.A., J.A. Meave, J.L. Villaseor, J.A. Gallardo-Cruz & E.E. Lebrija-Trejos. 2010. Vegetation of the heterogeneous landscape of Nizanda, Oaxaca, Mexico. Folia Geobot. 45: 143-161. Prez-Garca, E.A., A.C. Sevilha, J.A. Meave & A. Scariot. 2009. Floristic differentiation in limestone outcrops of southern Mexico and central Brazil: a beta diversity approach. Bol. Soc. Bot. Mx. 84: 45-58. PrezGutirrez, R., L.A. Solari, A. Gmez-Truena & V. A. Valencia. 2009. El terreno Cuicateco: cuenca ocenica en el sur de Mxico? Nuevos datos estructurales, geoqumicos y geocronolgicos. Rev. Mex. Cienc.Geol. 26: 222-242. Romero, G.A. & G. Carnevali, 1989. Novelties in the orchid 76: 454-461. Sauleda, R., P. Esperon & M. Ragan (en lnea). A reconsideration of Encyclia phoenicea and Encyclia plicata (abril 2011) http://rsauleda.tripod.com/ rpspapersphoe.html. Soto, M. 2003. Encyclia hanburyi (Lindl.) Schltr., Die Orchideen 209. 1914. Plate 559 in: E. Hgsater & M. Soto (Eds.). Icones Orchidacearum (Mexico) Fascicles 5 & 6, Orchids of Mexico Parts 2 & 3. Herbario AMO, Mxico D.F.

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LANKESTERIANA 12(1): 91. 2012.A NEW SPECIES OF CAMPYLOCENTRUM (ORCHIDACEAE: ANGRAECINAE) FROM COLOMBIA MARTA KOLANOWSKA 1,4 OSCAR ALEJANDRO PREZ ESCOBAR 2 & EDICSON PARRA SNCHEZ 31 2 Cll 2 sur No 9-11 Bugalagrande, Valle del Cauca, Colombia 3 Facultad de Ciencias Agropecuarias, Universidad Nacional de Colombia-Sede Palmira, Valle del Cauca, Colombia 4 Author for correspondence: martakolanowska@wp.pl ABSTRACT. Campylocentrum palominoi, a new species from the western Cordillera of the Andes, department of Valle del Cauca (Colombia), is described and illustrated. This species is similar to C. brenesii, from which it differs mainly by the subequal, ovate, obtuse lobes of the 3-veined lip, abscense of papillae at the middle lobe and the saccate spur which is widened towards the apex. RESUMEN Se describe e ilustra a Campylocentrum palominoi, una nueva especie de la Cordillera occidental de los Andes, departamento del Valle del Cauca (Colombia). Esta especie es similar a C. brenesii principalmente por los lbulos laterales del labelo obtusos, ovados, subiguales, lbulo medio sin papilas en la lmina y por el nectario sacciforme, angosto en la base y ancho en el pice. KEY WORDS : Orchidaceae, Angrecinae, Campylocentrum palominoi, Colombia, new species Campylocentrum includes about 60 species of epiphytic plants, ranging from Florida, Mexico, and the West Indies to Brazil (Dressler 1993, Bogarn & Pupulin 2009, 2010). Eleven species have been reported from Colombia (Ortiz V. & Uribe V. 2009). Species of the genus Campylocentrum are characterized by the absence of pseudobulbs, the lateral to the leaves or from the centers of abbreviated, with the sepals and petals subequeal and free. The lip is characterized by having a spur made up by labellar tissues and the column is very short, without a foot (Williams & Allen 1980). Species of Campylocentrum grow in the wet forests from near the sea level up to about 2000 m of elevation (Bogarn & Pupulin 2009). the area of Alto Dapa (department of Valle del Cauca, municipality of Yumbo), a distinctive species of Campylocentrum was found and it is described here as new: Campylocentum palominoi M. Kolanowska, O. Prez & E. Parra, sp. nov. TYPE: Colombia. Department of Valle del Cauca, Alto Dapa, about 2000 m, 27 Dec. 2010, M. Kolanowska & O. Prez 238 (holotype: UGDA). FIG 1. Haec species Campylocentrum brenesii Schltr. similis est sed labelli lobis subequalibus ovatis obtusis, medii lobi papillis absentibus et calcari versus apicem dilatato differt. Epiphytic, monopodial herbs up to 8 cm long. Roots developed from the internodes of the stem, Stems 4-8 cm long, foliaceous. Leaves 4-10 3-5 mm, distichous, coriaceous, articulated to the sheathing bases, the blade broadly-lanceolate to ovate, the apex obtuse, margins shortly ciliate. few, lateral, from the leaf axils, racemes minutely ciliate, 1.0-3.2 cm long, Flowers small, 3-4 mm in length, distichous, white to greenish cream, tubular. Floral bracts less than half long of the ovary, margin erose. Ovary 1.5-2 mm long, pubescent, hairs short, rounded

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FIGURE 1. 1. Campylocentum palominoi bracts. C Floral bract surface. D Flower (side view). E Column and ovary. F Dissected perianth. G Pollinia. Drawn by O. Prez & M. Kolanowska from the holotype. LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.10 LANKESTERIANA

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KOLANOWSKA et al. A new species of Campylocentrum from Colombia11at the apex. Sepals subequal, narrowly-ovate to broadly elliptic, the apex obtuse, 1-veined. Dorsal sepal 1.8-2.0 1.0-1.2 mm. Lateral sepals 1.3-1.6 mm long 0.80.9 mm wide. Petals 1.2-1.5 0.8-1.0 mm, narrowlyovate to broadly elliptic, slightly shorter than dorsal sepal, the apex obtuse, 1-veined. Lip 3-lobed; the lobes subequal, ovate, obtuse; the middle lobe about 0.35 0.5 mm, slightly larger than the lateral lobes, 3-veined, smooth, without a hairly callous; extended at the base into a curved, saccate, clavate, 3-lobuled spur, narrow at the base and wider at the apex; spur about 1.5 1-1.2 mm, obtuse. Column about 0.5 mm long, without a foot. Pollinia 2, ovoid. Capsule 4-5 mm long, ellipsoid. PARATYPES : Colombia. Valle del Cauca: Municipio de Yumbo. DAPA, 18C aprox. 2000 m, aprox. 29 Jun 2009, Oscar Prez, Edicson Parra & Jos Campo 201 (VALLE!); Alto Dapa, about 1980 m alt, 18 Dec 2010. O. Prez & M. Kolanowska 213 (UGDA!). DISTRIBUTION AND ECOLOGY: Known only from Western Cordillera in Colombia, in the municipality of Yumbo. Populations, each of 1-5 individuals are found in the cloud forest at about 1980-2000 m of elevation in open areas, on tree trunks and fences. Some individuals have been reported growing on trees of Guettarda sp. (Rubiaceae) in open, disturbed areas. EPONYMY: Named in honor of Carlos Fernando Palomino, a conservation enthusiast of the Sopo forests biodiversity (department of Cundinamarca, Colombia). Campylocentum palominoi is similar to C. brenesii Schltr. from which it differs by subequal, ovate, obtuse lobes of the lip, the absence of papillae at the middle lobe and a saccate spur which is widened towards the apex. C. brenesii is a variable species, especially within the size of the tepals, spur length and the shape of the lateral lobes of the lip (Bogarn and Pupulin 2010). However, the presence of hairly callous on the acute, triangular middle lobe and cylindric spur are the constant characters which differs it from C palominoi. AC K NOWL E D GME NTS The authors wish to thank the curators and staff of the herbaria AMO, COL, CUVC, JAUM, K, MO and VALLE for access to their collections. We are grateful to Dr. Guy Chiron for translating the diagnosis of the new species. The research described here has been supported by the grant from the Polish Ministry of Science and Higher Education (project no. N N304 043939). LITERATURE CITED Bogarn D. & Pupulin F. 2009. The genus Campylocentrum (Orchidaceae: Angraecinae) in Costa Rica: some critical on Andean Orchids. Universidad Tcnica Particular de Loja, Loja, Ecuador: 32-45. Bogarn D & Pupulin F. 2010. The genus Campylocentrum (Orchidaceae: Angraecinae) in Costa Rica: a revision. Harv. Pap. Bot. 15(2): 353-414. Dressler R. 1993. Field Guide to the Orchids of Costa Rica and Panama. Cornell University Press, New York: 292-294. Ortiz Valdivieso, P. & C. Uribe Vlez. 2009. Galera de Orqudeas de Colombia (CD edition). Asociacin Bogotana de Orquideologa, Bogot. Williams L. O. & Allen P. H. 1980. Orchids of Panama. Monogr. Syst. Bot. Missouri Bot. Gard. 4: 578-579.

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LANKESTERIANA

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Introduction. Global climate change is adversely affecting ecosystems worldwide and empirical models indicate that over time, these changes will have an increasing impact on the earths environment (Markham 1996, Hulme & Viner 1998, Williams et al. 2003, Parmesean 2006). Although there is climate change models, there are several noteworthy global trends and projections. Foremost, expectations are that average temperatures will increase in many regions. In addition, most models suggest that global rainfall patterns will change considerably, and in turn, many areas (see Markham 1996, Pounds et al. 1999). Consequently, species that are adapted to cool, moist environments will likely be threatened by climate change. A large body of research shows that high elevation ecosystems, particularly those in the tropics, are especially susceptible to climate change for several LANKESTERIANA 12(1): 13. 2012.ON THE RELATIONSHIP BETWEEN BRYOPHYTE COVER AND THE DISTRIBUTION OF LEP ANTHES SPP. BENJAMIN J. CRAIN Department Department of Biology, University of Puerto RicoRio Piedras, P.O. Box 23360 San Juan, PR 00931-3360, U.S.A. bcrainium@gmail.com ABSTRACT Epiphytic plant communities in tropical montane ecosystems are particularly vulnerable to climate change. Lepanthes Generally, these orchids are very small and often occur in highly exposed habitats, thus leading to the theory that they are dependent on bryophyte cover to prevent desiccation. Accordingly, the aim of this project was to determine if the distribution of Lepanthes is limited by the distribution of moss cover on trees. To accomplish this, bryophyte cover was measured on phorophytes hosting Lepanthes spp. as well as on other unoccupied trees in the surrounding areas. The results suggest that Lepanthes spp. are correlated with high levels of moss cover and there is often substantially more moss cover at locations where Lepanthes occur than at locations where they do not occur. Accordingly, bryophytes appear to form a commensalism with this diverse group of orchids. The results of this study are in agreement with similar studies that suggest moss cover is important for orchid growth and survival. Consequently, if climate change adversely affects moss coverage on trees, Lepanthes orchids may suffer concomitantly. RESUMEN. vulnerables al cambio climtico. Lepanthes para prevenir la desecacin. El objetivo de este proyecto fue determinar si la distribucin de Lepanthes est limitada por la distribucin de la cobertura de musgos en los rboles. Para lograr esto, se midi la cobertura de Lepanthes localizados en las reas circundantes. Los resultados sugieren que Lepanthes spp. esta correlacionada con altos niveles de cobertura de musgos, adems hubo substancialmente ms cobertura de musgos en los lugares donde Lepanthes comensalismo con este grupo diverso de orqudeas. Los resultados de este estudio concuerdan con estudios similares que sugieren que la cobertura de musgos es importante para el crecimiento y supervivencia de las orqudeas. Por consiguiente, si el cambio climtico tiene un impacto negativo en la cobertura de musgos, las orqudeas Lepanthes sufriran simultneamente. KEY WORDS: bryophyte cover, commensalism, host trees, Lepanthes, microhabitat, orchid distribution

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reasons (Foster 2001, Williams et al. 2003, Parmesean 2006). In these systems, changes in temperature and moisture availability can lead to a lifting of orographic cloud cover, which can subsequently lead to other changes in microclimate and community dynamics (Markham 1996, Pounds et al. 1999, Foster 2001, Williams et al. 2003). Many plant species may be unable to emigrate away from these types of environmental pressures because of their sessile nature (Foster 2001, Kelly & Goulden 2008). Over extended periods, unfavorable upslope vegetation shifts can occur that lead to a compression of important habitats (Foster 2001, Parmesean 2006). For many high elevation biotic communities, these changes can lead directly to fragmentation and mountaintop extinctions (Foster 2001, Williams et al. 2003). Research consistently suggests that epiphytes will respond particularly unfavorably to climate change, resulting in range shifts and extirpations (Benzing 1998, Hietz 1999, Foster 2001, Kelly & Goulden 2008, Zotz & Bader 2009). Most epiphytes in high elevation ecosystems show high sensitivities to climatic conditions due to their direct interface with the atmospheric environment. For example, many species are dependent upon the relatively permanent fog cover in cloud forests (Benzing 1998, Hietz 1999, Zotz & Bader 2009). Even minor changes in local conditions can alter growth, phenology, reproduction, and biotic partnerships, all of which are factors contributing to reductions in species survival (Johnson & Bond 1992, Parmesean 2006, Colwell et al. 2008, Liu et al. 2010). Accordingly, epiphytic plant communities in tropical montane systems warrant immediate attention from conservation biologists. Lepanthes Swartz (Orchidaceae) is a large and diverse genus of primarily epiphytic orchids. These orchids are distributed throughout the Neotropics with a high number of species occurring in the montane cloud forests of Costa Rica (Dressler 1993, Luer 2003). Previous research suggests that the distribution of Lepanthes orchids may be highly sensitive to the presence of bryophyte cover, but this relationship has only been demonstrated on a single species from Puerto Rico (Tremblay et al. 1998). Nevertheless, the complex associations between bryophytes and other plant species are well established, and a variety of positive and negative interactions exist (During & Van Tooren 1990). Unfortunately, however, bryophytes may also be highly susceptible to changes in climatic conditions availability (Benzing 1998, Raven et al. 2005, Raabe et al. 2010), and thus, the fate of Lepanthes spp. may be jeopardized directly as a result of climate change, but also indirectly via impacts to potential orchid/ bryophyte commensalism. Therefore, the overall goal of this study was to determine if the local distribution of Lepanthes spp. is correlated with bryophyte cover. Results generated by this analysis should help to determine the potential effects that loss of bryophyte cover would have on the distribution of this diverse group of orchids. Previously, it has been predicted that bryophytes provide an important substrate for the recruitment, growth, and survival of orchids, and therefore Lepanthes spp. should occur on trees with substantial amounts of moss cover (Tremblay et al. 1998, Tremblay & Velazquez-Castro 2009). Accordingly, the hypothesis tested here is that the sections of phorophytes occupied by Lepanthes equivalent sections of un-colonized trees. Methods. To determine if the local distribution of Lepanthes orchids is limited by the distribution of bryophyte cover, moss coverage was measured on phorophytes hosting Lepanthes spp. as well as on unoccupied trees in the immediate surrounding areas. Between 15 June and 25 July 2011, surveys were conducted in several forests among various regions of Costa Rica in an attempt to locate individuals of a variety of different species of Lepanthes two forests were surveyed in Monteverde, Puntarenas Province, two forests in Heredia Province including one in La Selva Biological Station and another at the Bijagual Reserve, a single forest at the Cueric Field Station, San Jose Province, and two forests in Punterenas Province including one at the Las Cruces Field Station and another at the Las Alturas Lepanthes have either been previously collected or reported as being present at each of these sites (Luer 2003), populations were only located at half of the survey sites. Plants were Lepanthes genus based LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.14 LANKESTERIANA

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unavailable (see Dressler 1993, Luer 2003). At each site where a Lepanthes orchid was located, plots centered on the host tree were established. Bryophyte cover was then measured on the host tree at the position occupied by the orchid. This was accomplished by using a clear acetate sheet with a 20 x 20 cm metric grid printed on it. This grid size was chosen because it was large enough to cover the entire area surrounding each orchid, but small enough to limit sampling to the area directly interacting with it. In no instance was the plant larger than 20 cm and roots never extended outside of the grid area. To estimate moss cover on the section occupied by the orchid, the center of the grid was placed directly at the base of the orchid, and the number of 1 cm cells occupied by moss was counted. The overall number was then converted into a percentage of the maximum grid area (400 cm). This conversion was done mainly to accommodate different sizes of trees on which orchids were found. In cases where the circumference of the phorophyte was less than 20 cm, the entire grid could not be used. Therefore, the subset of the grid that did not overlap onto itself when placed over the orchid was used as the maximum potential grid area. In such cases, the percent cover was calculated in the same manner, but using the overall area of the usable subset of the grid as the denominator. Depth of moss cover was initially considered as a variable, but in all instances, the depth of the mat (when present) was approximately equal more than one individual of Lepanthes was found on a given tree, particularly on trees with branching trunks. To avoid effects of pseudo-replication, the sampling process was only repeated for other orchids that were located on separate trunks of the original host tree or for other orchids found on the adjacent trees within the plot. Lastly, the bearing and vertical position of the orchid on the central phorophyte was measured to the nearest cm. After data was collected at each host tree, the moss cover was measured on four non-host trees within each plot to establish a mean background moss cover. Nonhost trees were selected by choosing the individual trees nearest the central host tree within each of four quadrants delineated by the cardinal directions. Only trees that were greater than 2 cm diameter at breast height were selected for sampling, as no orchids were discovered on trees smaller than this size. Moss cover on each non-host tree was measured at the same height and bearing that was recorded for the position of the orchid at the plot center. Percent cover was calculated in the same manner as for host tree sites. Finally, the data from each site were analyzed with non-parametric Wilcoxon rank sum tests with continuity corrections to look for differences in median values of bryophyte cover between the sampled sections of host trees and non-host trees (i.e. baseline cover) at each individual site. These tests were conducted on data from each site to determine if the relationship between orchids and bryophytes was the same at each location. The Wilcoxon rank sum test was chosen mainly for its ability to handle the relatively small sample sizes that are often obtained during orchid surveys at a given site due to their patchy distributions. All statistical tests and analyses were conducted with the R version 2.13.0 computer statistical program (R Development Core Team 2011). Results. During the course of the surveys, Lepanthes populations were discovered at several sites within three areas. In the Monterverde area, Lepanthes host trees were located at three sites within one of the forests surveyed. In Cueric, two individual sites contained multiple host trees. Lastly, at Las Alturas, host trees were found at two additional sites. In Monteverde and Las Alturas, Lepanthes populations were found on at least two tree species; in Cueric, the populations were located on a single tree species. Non-host trees at each site included individuals of the same species as the host trees, as well as other species. The results of the surveys were consistent with expected results, as Lepanthes orchids are often patchily distributed among several tree species and are most common in middle to high elevation sites (Tremblay 1997, Riofro et al. 2007). Considering all the trees surveyed, moss cover on sampled sections ranged from 0-100% (Fig. 1) with an average cover of 65%. Considering only the host trees, moss was always present and overall cover on sampled sections ranged from 28-100%. The mean moss cover on the sampled sections of host trees was approximately 81%. Considering only non-host trees, moss cover on the sampled sections ranged from LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. CRAIN Lepanthes and bryophyte cover15

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0-100%, with a mean coverage of only 48% (Fig. 1). The Wilcoxon rank sum tests conducted for trees from individual sites showed that there was a difference in median moss cover on sampled sections of host trees versus non-host trees at Monteverde (W = 125, n = 12, P = 0.002373) and Cueric (W = 62, n = 8, P = 0.001717), but not at Las Alturas (W = 35, n = 8, P = 0.7909, Fig. 1). This was likely the result of the higher mean background moss cover on non-host trees in Las Alturas (60%) than on non-host trees at the other sites (Monteverde47%, Cueric36%). In all cases, however, orchids occupied locations with substantial amounts of moss cover (Fig. 1). Discussion. The results of this study are in agreement with and expand upon the findings of previous research on moss cover and the presence of Lepanthes spp. (Tremblay et al. 1998, Tremblay & Velasquez-Castro 2009) and other orchids (Gowland et al. 2007, Watthana & Pedersen 2008, OMalley 2009, Scheffknecht et al. 2010, Gowland et al. 2011). In short, it appears that a large number of Lepanthes spp. from various locations and environments form strong relationships with epiphytic bryophytes. Several theories as to why this is the case seem plausible, and indeed all may be acting in unison. First, bryophytes could provide a moist heterogeneous substrate that may be beneficial to Lepanthes orchids because they provide a suitable landing place for dust-like orchid seeds in an otherwise potentially smooth, dry, bare bark environment (Tremblay et al. 1998, OMalley 2009, Scheffknecht et al. 2010). Secondly, moss beds could also support the growth of mycorrhizal fungi necessary for seed germination and transfer of nutrients that the mosses themselves may provide (Tremblay et al. 1998, Cornelissen et al. 2007, Osorio-Gil et al. 2008, Tremblay & VelasquezCastro 2009). Third, moss beds could shield orchid roots from exposure to wind and sun and thus help buffer plants from desiccation during dry periods (Benzing 1998, Venturieri & Mendoza de Arbieto 2011). Finally, many moss species leach nutrients and other chemicals into water that may stimulate orchid growth (Coxson et al. 1992, Carlton & Read 1991, Benzing 1998, Clark et al. 1998) or act as fungicides (Frahm 2004), which could perhaps prevent fungal parasitism of Lepanthes. Certainly, these are only a subset of the potential benefits that bryophytes could provide to epiphytic Lepanthes and additional manipulative studies are necessary to test the theories as to why these orchids are associated with bryophyte cover. Accordingly, it is evident that in many cases, conservation of Lepanthes spp. will require us to focus our attention on epiphyte communities as a whole, as opposed to only on isolated species. Scheffknecht et al. (2010) conclude that bryophytes are highly representative of microclimate characteristics and could thus be useful indicators of habitat quality for orchids. Even in the absence of a direct causal relationship between the distribution of orchids and bryophytes (e.g. Gowland et al. 2011), their dependency on similar conditions is an indication that these plants could be susceptible to the same threats. A positive corollary of the general sensitivity of many tropical montane bryophytes is that they may act as important environmental indicators and may be useful for foreshadowing oncoming threats to many orchids before their LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.16 LANKESTERIANA FIGURE 1. Box-plot showing differences in the percent of moss cover between sampled sections of host trees (H) and random non-host trees, i.e., baseline moss cover (B), at three sites in Costa Rica. Numbers in parentheses next to site names are the mean baseline moss coverage estimates for sampled sections on random non-host quartiles, horizontal lines within the boxes are the median values, and lines extending from the boxes represent the overall range of values. The vertical lines between the paired box plots for each site represent the difference in median moss cover on sampled sections of host trees and median baseline coverage. Asterisk (*) indicates an outlier.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. CRAIN Lepanthes and bryophyte cover17effects are realized (Gignac 2001). Recent studies of bryophyte distributions have already indicated that global warming trends are driving species upslope and are causing extinctions at lower elevations (Bergamini et al. 2009). If orchids are reliant upon the distribution bryophytes, we can anticipate similar trends in their distributions. From a conservation perspective, the results presented here demonstrate that climate change could have a double impact on Lepanthes spp. in Costa Rica and elsewhere. A large number of species from this genus are distributed in highly susceptible montane cloud-forest environments (Hammel et al. 2003). No doubt, these species are adapted to, and dependant upon, the cool temperatures and the moist cloud cover that persists in these forests (Benzing 1998, Zotz & Bader 2009). Similarly, the bryophytes that these orchids associate with are equally dependent of these cool and moist conditions (Hallingbck & Tan 2010). Changes leading to hotter and dryer conditions in montane cloud forests such as Monteverde, Cueric, and Las Alturas could directly and indirectly eliminate the conditions necessary for the survival of numerous species. If Lepanthes spp. can adapt to these changing conditions, or perhaps migrate away from them (see Hietz 1999, Foster 2001), they may still be in jeopardy if their bryophyte partners cannot persist. Recent analyses indicate that loss of bryophytes could eliminate many important resources required by Lepanthes spp. (Tremblay et al. 1998, Tremblay & Velazquez-Castro 2009, Gowland et al. 2011). Consequently, the ability of these orchids to adapt to a changing climate may be a secondary concern if the bryophytes they depend on cannot. ACKNOWLEDGEMENTS. I would like to thank E. Kuprewicz, D. McClearn, K. McManus, N. MartnezCampos, and A. Sanchez-Cuervo for assisting with this project. I also thank K. Gerow for helping with the statistical analysis in this project. I greatly appreciate the generosity of the staff at the Lankester Botanical Garden in Cartago, Costa Rica. Special various stations at which this research was carried out. Finally, I would like to thank R. Tremblay for inspiring me to work with Lepanthes and providing me with the opportunity to do so. LITERATURE CITED Benzing, D. 1998. Vulnerabilities of tropical forests to Clim. Change 39 : 519-540. Bergamini, A., S. Ungricht, H. Hofmann. 2009. An elevational shift of cryophilous bryophytes in the last centuryan effect of climate warming? Divers. Distrib. 15: 871-879. Carlton, T., D. Read. 1991. Ectomycorrhizas and nutrient transfer in coniferfeather moss ecosystems. Can. J. Bot. 69: 778-785. Clark, K., N. Nadkarni, H. Gholz. 1998. Growth, net production, litter decomposition, and net nitrogen accumulation by epiphytic bryophytes in a tropical montane forest. Biotropica 30: 12-23. Cornelissen, J., S. Lang, N. Soudzilovskaia, H. During. 2007. Comparative cryptogram ecology: a review of bryophyte and lichen traits that drive biogeochemistry. Ann. Bot. 99: 987-1001. Coxson, D., D. McIntyre, H. Vogel. 1992. Pulse release of sugars and polyols from canopy bryophytes in tropical montane rain forest (Guadaloupe West Indies). Biotropica 24: 121-133. Dressler, R. 1993. Field Guide to the Orchids of Costa Rica and Panama. Cornell University Press, Ithaca, NY. During, H., B. Van Tooren. 1990. Bryophyte interactions with other plants. Bot. J. Linn. Soc. 104: 79-98. Foster, P. 2001. The potential negative impacts of global climate change on tropical montane cloud forests. Earth Sci. Rev. 55: 73-106. Frahm, J. 2004. Recent developments of commercial products from bryophytes. Bryologist 107: 277-283. Gignac, L. 2001. Bryophytes as indicators of climate change. Bryologist 104: 410-420. Gowland K., U. Mathesius, M. Clements, A. Nicotra. 2007. Understanding the distribution of three species of epiphytic orchids in temperate Australian rainforest by investigation of their host and fungal associates. Lankesteriana 7: 44-46. Gowland, K., J. Wood, M. Clements, A. Nicotra. 2011. Bot. 98: 197-206. Hallingbck, T., B. Tan. 2010. Past and present activities and future strategy of bryophyte conservation. Phytotaxa 9: 266-274. Hietz, P. 1999. Diversity and conservation of epiphytes in a changing environment. Proceedings of the International Conference on Biodiversity and Bioresources: Conservation and Utilization. 23 November 1997, Phucket, Thailand. 1-11.

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Hulme, M., D. Viner. 1998. A climate change scenario for the tropics. Clim. Change 39: 145-176. Johnson, S., W. Bond. 1992. Habitat dependent pollination success in a cape orchid. Oecologia 91: 455-456. Kelly, A., M. Goulden. 2008. Rapid shifts in plant distribution with recent climate change. P. Natl. Acad. Sci. U.S.A. 105: 118231826. Liu, H., Y. Luo, H. Liu. 2010. Studies of mychorrizal fungi of Chinese orchids and their role in orchid conservation in Chinaa review. Bot. Rev. 76: 241-262. Luer, C., 2003. Lepanthes. Pp. 216-255 in: Hammel, B. M. Grayum, C. Herrera, N. Zamora (eds.), Manual de Plantas de Costa Rica, Volumen II Gimnospermas y Monocotiledoneas (Agavaceae-Musaceae). Missouri Botanical Garden Press, St. Louis, MO. Markham, A. 1996. Potential impacts of climate change on ecosystems: a review of implications for policymakers and conservation biologists. Climate Res. 6: 179-191. OMalley, K. 2009. Patterns of abundance and diversity in epiphytic orchids on Parashorea malaanonan trees in Danum Valley, Sabah. Ply. Stud. Sci. 2: 38-58. Osorio-Gil, E., J. Forero-Montaa, T. Otero. 2008. Variation in mycorrhizal infection of the epiphytic orchid Ionopsis utriculariodes (Orchidiaceae) on different substrata. Caribbean J. Sci. 44: 130-132. Parmesan, C. 2006. Ecological and evolutionary responses to recent climate change. Ann. Rev. Ecol. Evo. Syst. 37: 637-669. Pounds, J., M. Fogden, J. Campbell. 1999. Biological response to climate change on a tropical mountain. Nature 398: 611-615. R Development Core Team. 2011. R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org. Raabea, S., J. Mller, M. Manthey, O. Drhammer, U. Teuber, A. Gttlein, B. Frster, R. Brandl, C. Bssler. 2010. Drivers of bryophyte diversity allow implications for forest management with a focus on climate change. Forest Ecol. and Manag. 260: 1956-1964. Raven, P., R. Evert, S. Eichhorn. 2005. The Biology of Plants. W.H. Freeman and Company, New York, NY. Riofro, L., C. Naranjo, J. Iriondo, E. Torres. 2007. Spatial structure of Pleurothallis Mazdevallia Lepanthes, and Epidendrum epiphytic orchids in a fragment of montane cloud forest in south Ecuador. Lankesteriana 7: 102-106. Scheffknecht, S., M Winkler, K. Hlber, M. Mata-Rosas, P. Hietz. 2010. Seedling establishment of epiphytic orchids in forests and coffee plantations in central Veracruz, Mexico. J. Trop. Biol. 26: 93-102. Tremblay, R. 1997. Distribution and dispersion patterns of individuals in nine species of Lepanthes. Biotropica 29: 38-45.LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.18 LANKESTERIANA

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. This article series is part of an ongoing study of et al. 2008). of a living collection of native orchids has been an invaluable tool for the inventory efforts that have resulted in a rapid increase in the knowledge of the be much more effective than studying only herbarium material. Since the latest treatment of the Orchidaceae in the Manual de Plantas de Costa Rica (referred to as the Manual throughout this paper; Dressler 2003), many species were reported or described from Costa Rica. A few hundred novelties (both nomenclatural and taxonomic) in some 50 genera were published since the Manual was printed. Most are actually just new names for known species and therefore do not account for a net increase in the number of species, however increase in Costa Rican orchid diversity. A summary of all articles that have appeared since the publication of the Manual and include species descriptions or nomenclatural changes affecting Costa Rican orchid taxa is presented in Table 1. Novelties were described from remote areas and from relatively well-botanized sites, and frequently citing few herbarium specimens. For example, only two plants of Acianthera cabiriae Pupulin, G.A.Rojas & J.D.Zuiga were ever found at the well botanized site of CATIE, Turrialba (Pupulin et al. 2007). On the other hand, several plants of Pleurothallis adventurae Karremans & Bogarn were found growing LANKESTERIANA 12(1): 19. 2012.NEW SPECIES AND RECORDS OF ORCHIDACEAE FROM COSTA RICA. II ADAM P. KARREMANS 1,2,3,6 DIEGO BOGARN 1,2,4 MELANIA FERNNDEZ 1,4, CHRISTINA M. SMITH 1 & MARIO A. BLANCO 1,51 Jardn Botnico Lankester, Universidad de Costa Rica, P. O. Box 302-7050 Cartago, Costa Rica 2 Centro de Investigacin en Orqudeas de los Andes ngel Andreetta, Universidad Alfredo Prez Guerrero, Ecuador 3 NCB Naturalis NHN Universiteit Leiden, The Netherlands4 Herbario UCH, Universidad Autnoma de Chiriqu, David, Panama 5 Escuela de Biologa, Universidad de Costa Rica 6 Corresponding author: adam.karremans@ucr.ac.cr ABSTRACT After the publication of the most recent and comprehensive treatment of the Orchidaceae in the Manual de Plantas de Costa Rica, new species continue being discovered in the country on a regular basis. Novelties in Acianthera Epidendrum, Lepanthes, Masdevallia Pleurothallis, and Specklinia are discussed. We present eight new records including the reconsideration of Epidendrum concavilabium (as different from E. circinatum Epistephium (E. ellipticum) for the country. Three new taxa, Epidendrum alieniferum, Epidendrum sandiorum (a putative natural hybrid between E. oerstedii and E. ciliare) and Lepanthes kabebatae are described. RESUMEN. Despus de la publicacin del tratamiento ms reciente y completo de Orchidaceae en el Manual de Plantas de Costa Rica, nuevas especies siguen descubrindose con regularidad en el pas. Se discuten novedades en Acianthera, Epidendrum, Lepanthes, Masdevallia Pleurothallis y Specklinia Presentamos ocho nuevos registros incluyendo la reconsideracin de Epidendrum concavilabium (como distinto de E. circinatum) y el primer registro del gnero Epistephium (E. ellipticum ) en el pas. Se describen tres nuevos taxones, Epidendrum alieniferum, Epidendrum sandiorum (un hbrido natural putativo entre E. oerstedii y E. ciliare) y Lepanthes kabebatae KEY WORDS: Epidendrum alieniferum, Epidendrum sandiorum, Lepanthes kabebatae, Epistephium, Orchidaceae, new records, new species, Costa Rica

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.20 LANKESTERIANA TABLE 1. List of orchid genera cited in the Manual de Plantas de Costa Rica for which species have been added and/or Record; NS= New Species; Y= Increase in net species number; N= No increase in net species number.Genus Reference Category Acianthera Pupulin, Rojas & Ziga 2007; Bogarn et al. 2008 NS-Y Anathallis Luer 2006 NR-Y Aspidogyne Ormerod 2007; Ormerod 2009 NS-Y Barbosella Bogarn et al. 2008 NR-Y Brassia Christenson 2003b; Pupulin & Bogarn 2005a NC/NS-Y Campylocentrum Pupulin & Bogarn 2010b NC/NS/NR-Y Chondroscaphe Pupulin 2005a; Pupulin et al. 2009. NC/NS-N Christenson 2003a; Dressler & Dalstrm 2004 NC/NS-N Coccineorchis Rutkowski et al. 2004 NS-Y Coryanthes Gerlach & Dressler 2003; Gerlach & Romero-Gonzlez 2008 NC/NS-N Crossoglossa Pupulin & Karremans 2010a NS-Y Cyrtopodium Romero-Gonzlez & Carnevali 1999 NC-N Dichaea Pupulin 2005a; Pupulin 2005b; Pupulin 2005c; Dressler, Pupulin & Folsom 2006; Pupulin 2007 NC/NS-N Dracontia Karremans in press; Karremans & Smith in press NC/NS-N Dryadella Luer 2005 NC-N Brenesia Bogarn et al. 2008 NR-Y Elleanthus Dressler & Bogarn 2007; Dressler & Bogarn 2010 NS-Y Encyclia Pupulin 2006 NS-Y Epidendrum Bogarn et al. 2008; Hgsater 2003; 2004; 2006; 2007; 2008; 2009; Karremans & Hgsater 2010; Pupulin & Karremans 2010b NC/NS/NR-Y Galeandra Pupulin 2005a NS-Y Gongora Jenny 2007 NS-Y Habenaria Batista et al. 2011 Kefersteinia Pupulin & Merino 2008 NS-Y Kraenzlinella Luer 2011 NS-Y Kreodanthus Ormerod 2008 NS-Y Lepanthes Blanco 2003; Bogarn & Fernndez 2010; Bogarn & Pupulin 2010a; 2011; Pupulin 2003a; Pupulin & Bogarn 2004; 2010a; 2011a; 2011b; 2012; Pupulin, Bogarn & Jimnez 2009; Pupulin, Medina & Bogarn 2010; Pupulin, Bogarn & Smith 2010 NS/NR-Y Lockhartia Bogarn et al. 2008 NR-Y Lycaste Bogarn 2007; Oakeley 2008 NC/NS-Y Masdevallia Smith & Pupulin in prep. NS-Y Maxillaria Pupulin 2003b; Bogarn et al. 2008 NS/NR-Y Microchilus Ormerod 2004; 2005; 2007 NC/NS-Y Mormolyca Bogarn & Pupulin 2010 NS-Y Myoxanthus Bogarn et al. 2008; Pupulin, Bogarn & Fernndez 2010b NS/NR-Y Octomeria Luer 2010 NR-Y Ornithidium Blanco et al. 2008a NR-Y Ornithocephalus Pupulin 2002a NS-Y Palmorchis Bainbridge & Aguilar 2008 NR-Y Panmorphia Luer 2006 NR-Y Phragmipedium Christenson 2006a; Pupulin & Dressler 2011 NC-N

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica21 TABLE 1. Continues.on a tree just outside of a shelter cabin close to the Panamanian border in the Cordillera de Talamanca, in a less intensely explored region. No material of either species was found in any of the main Costa Rican herbaria (CR, INB, JBL, USJ), which indicates that some species might be both conspicuous and relatively frequent but grow in relatively unexplored regions, while others might occur in more frequented sites but be scarce and/or inconspicuous. In addition to novelties, important nomenclatural changes were proposed for many orchid taxa present in Costa Rica in the last decade. Proposals to split Masdevallia Ruiz & Pav., Maxillaria Ruiz & Pav., Pleurothallis R.Br. and Prosthechea Knowles & Westc. are the ones that account for most of the new combinations. A summary of these proposed generic segregates is presented in Table 2. On the other hand proposals to reduce several genera to the synonymy of others were also published; most notable are the inclusion of Oerstedella Rchb.f. in Epidendrum L., Sigmatostalix Rchb.f. in Oncidium Sw. and Stellilabium Schltr. in Telipogon Kunth. A summary of those proposed generic lumpings is presented in Table 3. All in all, close to half of the orchid genera accepted in the Manual have either gained and/or lost species due to generic transfers, species descriptions and/or reduction to synonymy. Given the steady discovery of novelties and phylogenetic reconstructions based on DNA, any published orchid inventory for Costa Rica becomes outdated in a matter of a few years. Bogarn (2011) listed ca. 200 new names published for the that tendency is maintained, which is a likely scenario, in a couple of decades the number of known orchid species in the country will exceed 2000. As part of routine botanical exploration, documenBotanical Garden, more novelties continue to accumulate. On that note, we reveal the following new records from Costa Rica. Most of these were discovered from the direct study of live material; in all cases additional specimens were sought in the main Costa Rican herbaria (CR, INB, JBL, USJ) and, if found, are cited in the corresponding accounts presented below.Genus Reference Category Platystele Bogarn & Karremans 2010 NS-Y Platythelys Ormerod 2007 NS-Y Pleurothallis Karremans & Bogarn 2011; Karremans & Muoz 2011; Luer 2002; Pupulin & Ziga 2007; Pupulin, Bogarn & Fernndez 2010a; NS-Y Polycycnis Gerlach 2004 NC/NS-N Polystachya Mytnik-Ejsmont 2011 NR-Y Prosthechea Pupulin 2004 NS-Y Restrepiella Pupulin & Bogarn 2007 NS-Y Scaphyglottis Dressler 2004a; Bogarn et al. 2008 NS/NR-Y Scelochilus Pupulin & Bogarn 2005b NC-N Sigmatostalix Pupulin & Rojas 2006 NS-Y Sobralia Bogarn et al. 2008; Dressler 2002; 2007; Dressler & Pupulin 2008; Dressler & Bogarn 2010; 2011 NC/NS/NR-Y Specklinia Luer 2006 NR-Y Stanhopea Gerlach & Beeche 2004 NC-N Stenorrhynchos Christenson 2005 NC/NS-Y Stellilabium Pupulin 2003b; Pupulin 2003c; Pupulin & Blanco 2002 NS-Y Trichopilia Dressler & Pupulin 2005; 2006; Dressler & Bogarn 2009 NC/NS-Y Trichosalpinx Fernndez 2011; Fernndez & Bogarn 2012 NS-Y Trigonidium Christenson 2002 NC-N Vanilla Soto Arenas & Dressler 2010 NC/NS-Y Warmingia Bogarn et al. 2008 NC-N

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.22 LANKESTERIANA TABLE 2. List of orchid genera cited in the Manual de Plantas de Costa Rica which have been split by recent authors. Segregate genera are given for each generic name accepted in the Manual.Genus Segregate Genera Reference Cattleya Guarianthe Dressler & W.E.Higgins Dressler & Higgins 2003Chondrorhyncha Benzingia Dodson Romero-Gonzlez & Dodson 2010 Daiotyla Dressler Whitten et al. 2005 Stenotyla Dressler Whitten et al. 2005Elleanthus Adeneleuterophora Barb. Rodr. Dudek & Szlachetko 2010 Epylina Schltr. Dudek & Szlachetko 2010 Evelyna Poepp. & Endl. Dudek & Szlachetko 2010Eltroplectris Callistanthos Szlach. Szlachetko & Rutkowski 2008Epidendrum Coilostylis Raf. Whitner & Harding 2004Erythrodes Aspidogyne Garay Ormerod 2007; Ormerod 2009 Kreodanthus Garay Ormerod 2008 Microchilus C.Presl Ormerod 2002 Platythelys Garay Ormerod 2007Galeottiella Funkiella Schltr. Solano et al. 2011Habenaria Bertauxia Szlach. Szlachetko 2004a Habenella Small Szlachetko & Kras 2006 Platantheroides Szlach. Szlachetko 2004bKefersteinia Senghasia Szlachetko Szlachetko 2003; Szlachetko, Kulak & Romowicz 2006.Lycaste Selbyana Archila Archila 2010Malaxis Microstylis Masdevallia Acinopetala Luer Luer 2006 Alaticaulia Luer Luer 2006 Buccella Luer Luer 2006 Diodonopsis Pridgeon & M.W.Chase Pridgeon & Chase 2001 Fissia (Luer) Luer Luer 2006 Reichantha Luer Luer 2006 Spilotantha Luer Luer 2006 Zahleria (Luer) Luer 2006Maxillaria Adamanthus Camaridium Lindl. Blanco et al. 2007 Christensonella Heterotaxis Lindl. Ojeda, Carnevali & Romero 2005 Inti M.A.Blanco Blanco et al. 2007 Mapinguari Carnevali & R.B.Singer Blanco et al. 2007 Maxillariella M.A.Blanco & Carnevali Blanco et al. 2007 Mormolyca Fenzl Blanco et al. 2007 Nitidobulbon Ojeda, Carnevali & G.A.Romero Ojeda, Carnevali & Romero 2009 Ornithidium Salisb. ex R. Br. Blanco et al. 2007 Rhetinantha M.A.Blanco Blanco et al. 2007 Sauvetrea Szlach. Blanco et al. 2007Oncidium Brevilongium Christenson Christenson 2006b Chelyorchis Dressler & N.H.Williams Dressler & Williams 2000; Carnevali et al. 2009 Heteranthocidium Szlach., Mytnik & Romowicz Szlachetko, Mytnik & Romowicz 2006 Otoglossum (Schltr.) Garay & Dunst. Williams et al. 2001

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.23Genus Segregate Genera Reference Oncidium Rossioglossum (Schltr.) Garay & G.C.Kenn. Chase et al. 2008 Stacyella Szlach. Szachetko 2006 Trichocentrum Poepp. & Endl. Williams et al. 2001 Vitekorchis Romowicz & Szlach. Romowicz & Szachetko 2006. Pleurothallis Aberrantia Luer Luer 2005 Acronia C.Presl Luer 2005 Acianthera Scheidw Pridgeon & Chase 2001; Solano et al. 2011 Anathallis Barb.Rod. Pridgeon & Chase 2001; Hgsater & Soto 2003 Ancipitia (Luer) Luer Luer 2004 Apoda-prorepentia (Luer) Luer Luer 2004 Crocodeilanthe Rchb.f. & Warsz. Luer 2004 Didactylus Luer Luer 2005 Dracontia (Luer) Luer Luer 2004 Echinella Pridgeon & M.W.Chase Pridgeon & Chase 2001 Echinosepala Pridgeon & M.W.Chase Pridgeon & Chase 2002 Effusiella Luer Luer 2007 Elongatia (Luer) Luer Luer 2004 Empusella (Luer) Luer Luer 2004 Gerardoa Luer Luer 2006 Kraenzlinella Kuntze Luer 2004; Hgsater & Soto 2003 Loddigesia Luer Luer 2006 Lomax Luer Luer 2006 Muscarella Luer Luer 2006 Niphantha Luer Luer 2010 Pabstiella Brieger & Senghas Luer 2007 Panmorphia Luer Luer 2006 Phloeophila Hoehne & Schltr. Pridgeon & Chase 2001; Luer 2006 Rhynchopera Ronaldella Luer Luer 2006 Sarcinula Luer Luer 2006 Specklinia Lindl. Pridgeon & Chase 2001; Hgsater & Soto 2003 Stelis Sw. Pridgeon & Chase 2001 Tribulago Luer Luer 2004; Luer 2006 Sylphia Luer Luer 2006 Unciferia (Luer) Luer Luer 2004 Unguella (Luer) Luer Luer 2005 Zosterophyllanthos Prosthechea Anacheilium Rchb.f. ex Hoffmanns Whitner & Harding 2004 Hormidium (Lindl.) Heynh. Whitner & Harding 2004 Panarica Withner & P. A.Harding Whitner & Harding 2004 Pollardia Withner & P. A.Harding Whitner & Harding 2004 Pseudencyclia Chiron & V.P.Castro Chiron & Castro-Neto 2003Schiedeella Funkiella Schltr. Solano et al. 2011Stanhopea Stanhopeastrum Rchb.f Szlachetko 2007Trichosalpinx Tubella (Luer) Archila Archila 2000 KARREMANS et al. New species and records of Orchidaceae from Costa Rica TABLE 2. Continues.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.24 LANKESTERIANA1. Acianthera oscitans (Ames) Pridgeon & M.W.Chase, Lindleyana 16: 245. 2001; Pleurothallis oscitans Didactylus oscitans (Ames) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 257. 2004. TYPE : Honduras, Corts, Santa Cruz de Yojoa. Epiphyte in open mountain forest at 2000 feet altitude. Flowers dark purple. August 26, 1933. J.B. Edwards 515 (holotype, AMES). DISTRIBUTION : Honduras, Costa Rica and Panama. ETYMOLOGY : Not indicated in the protologue. Luer (2004) suggested the name probably refers to the HABITAT IN COSTA RICA : This species has been found growing epiphytically at around 700 m in the transition from very humid tropical lowland forest to premontane forest of the Braulio Carrillo National Park, on the Atlantic watershed of the Central Volcanic chain. PHENOLOGY : Flowering around August under cultivation. COSTA RICAN MATERIAL STUDIED : San Jos: Vzquez de Coronado, Jess, Parque Nacional Braulio Carrillo, Sendero La Botella, 10.9N 83.8W, 702 m, bosque muy hmedo tropical transicin a 2011, D. Bogarn 7621, M. Fernndez & A.P. Karremans Horquetas, Colonia Cubujuqu, hacia las orillas del Braulio Carrillo. 10N 84 W, ca. 300 m. A orillas de un riachuelo. 25 de febrero 2012, A.P. Karremans 5175 (CR!). Acianthera oscitans is most similar to Acianthera butcheri (L.O.Williams) Pridgeon & M.W.Chase, from which it can be distinguished by the ellipticovate leaves, the flower with the tip of the dorsal sepal connate to the synsepal (reminiscent of the flowers of some Zootrophion spp.), the synsepal with revolute margins and the lip narrowly ovate and ciliate. Luer (2005) placed A. oscitans, A. butcheri (also known from Costa Rica) and the Ecuadorian Pleurothallis paradoxa Luer & Dalstrm and Pleurothallis thysana Luer & J.Portilla together in the genus Didactylus, distinguished from Acianthera by the rostellum with two narrow, curved, lateral lobes. TABLE 3. List of orchid genera cited in the Manual de Plantas de Costa Rica which have been lumped into other genera by recent authors. The absorbing genus is given for each generic name accepted in the Manual. Genus Reduced under Reference Acostaea Specklinia Pridgeon & Chase 2001 Ada Brassia Chase & Whitten 2011 Amparoa Rhynchostele Hgsater & Soto 2003 Chelyorchis Rossioglossum Chase et al. 2008 Goniochilus Leochilus Chase et al. 2008 Hybochilus Leochilus Chase et al. 2008 Leucohyle Trichopilia Dressler 2004b Mesospinidium Brassia Chase & Whitten 2011 Oerstedella Epidendrum Hgsater & Soto 2005a Osmoglossum Cuitlauzina Dressler & Williams 2003 Pachyphyllum Fernandezia Chase & Whitten 2011 Pleurothallis Stelis Pridgeon & Chase 2001; Hgsater & Soto 2003 Psygmorchis Erycina Williams et al. 2001 Restrepiopsis Pleurothallopsis Pridgeon & Chase 2001 Salpistele Stelis Pridgeon & Chase 2001 Scelochilus Comparettia Chase et al. 2008 Sigmatostalix Oncidium Chase et al. 2008 Stellilabium Telipogon Williams et al. 2005 Ticoglossum Rossioglossum Chase et al. 2008

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.25 KARREMANS et al. New species and records of Orchidaceae from Costa Rica FIGURE 1. Acianthera oscitans (Ames) Pridgeon & M.W.Chase. A Habit; B Flower; C Dissected perianth; D Column and lip, lateral view; E Anther and pollinaria. Drawn by M. Fernndez from D. Bogarn 7621 (JBLSpirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.26 LANKESTERIANA FIGURE 2. Epidendrum alieniferum Karremans & Bogarn. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Column and lip, longitudinal section. F Pollinarium and anther cap. G Clinandrium lacerations. Drawn by A.P. Karremans, based on the plant used as type (JBL-Spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica272. Epidendrum alieniferum Karremans & Bogarn, sp. nov. TYPE : Costa Rica. Puntarenas: Coto Brus, Sabalito, Mellizas, entre Finca Gemelas y Finca Mellizas, 8.53 N 82.83 W, 1412 m, bosque muy cafetales pascuis supra Acnistus prope opidum Mellizas, 18 abril 2011, A. P. Karremans 3970, D. Bogarn & D. Jimnez 2, 14B). Species Epidendro lagenocolumnae Hgsater et Snchez profunde lacerato lacerationibus radialibus differt. Epiphytic, caespitose, erect herb, up to 25 cm tall. Stems terete, somewhat Leaves 4-8(9), distributed along the stem, especially close to the apex; sheath tubular, rugose; blades elliptic, obtuse, variable in size, the largest up to 6-10 2.5-3.2 cm. apical, subumbellate; peduncle reduced. Floral bracts acute, 0.5 cm long. Flowers 4-10, greenish. Ovary pedicellate, terete, smooth. Sepals partly spreading, slightly bent backwards, elliptic, obtuse, 7-veined, margin entire, revolute; dorsal sepal 21-22 11-12 mm, lateral sepals oblique, 20-22 10-11 mm. Petals spreading, porrect, narrowly-elliptic, obtuse, 5-veined, margin entire, revolute, 20-22 6-7 mm. Lip trilobed, basal lobes widely ovate-elliptic, mid-lobe sub-quadrate, emarginate, bicallose, with a low central keel, margins crenulate, folded, appearing cross-like in natural position, 22 22-23 mm. Column strongly arching downwards, thickened basally, sub-terete, 13-15 mm long, with a pair of arm-like wings; clinandriumhood erect, prominent, margins radially lacerate; rostellum near the apex of the column, slit. Anther cap transverse-elliptic, 4-celled, 3 mm wide. Pollinia 4, obovoid, laterally compressed. Nectary short, barely penetrating the ovary, smooth. Fruit ellipsoid. PARATYPES : Costa Rica. Same locality and date as the holotype, A. P. Karremans 3971, D. Bogarn & D. Jimnez (JBL-Spirit!). Puntarenas: Coto Brus. Z.P. Las Tablas. Cuenca Trraba-Sierpe. Estacin Progreso, S. 1440 m, 24 mayo 1999, M. Alfaro 183 (CR!, INB!). DISTRIBUTION : Known only from Costa Rica, but most probably occurring also in Panama, as the three known specimens were collected close to the Panamanian border. ETYMOLOGY : From the Latin alienus, alien, stranger, and fero, bearing. In allusion to the frontal view of the folded lip and column apex in natural position, strange beings. HABITAT IN COSTA RICA : Epiphytic in primary and secondary humid premontane forest, at around 1400-1450 m elevation. It is known only from the Cordillera de Talamanca. PHENOLOGY and in May under cultivation. Epidendrum alieniferum is most similar to E. lagenocolumna Hgsater & Snchez (1993), but it 22 vs. 13-18 mm long and 10-12 vs. 4-6 mm wide, respectively), the shape of petals (elliptic vs. linear), the lip crenulate (vs. entire), the base of the column than twice the width), and the prominent clinandrium, with radial lacerations (vs. obsolete). It also grows at lower elevations than E. lagenocolumna, which in Costa Rica is normally found above 1800 m, and watershed of the Talamanca range, whereas the former is known throughout the country. The variable E. Rchb.f. is also similar; however it is smaller in that is not thickened at the base and does not have such a prominent, erect, lacerate clinandrium (Snchez & Hgsater 2007). 3. Epidendrum concavilabium C.Schweinf., Bot. TYPE : Costa Rica. Colinas de San Pedro de San Ramn, Nov. 1927, A.M. Brenes (119) 1660 (holotype, AMES; photo of type at AMES!). DISTRIBUTION : Costa Rica.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.28 LANKESTERIANA FIGURE 3. Epidendrum concavilabium C.Schweinf. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Column, ventral view. F Pollinarium and anther cap. Drawn by A.P. Karremans, based on D. Bogarn 4848 (JBL-Spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica29 FIGURE 4. Epidendrum circinatum Ames. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Column, ventral view. Drawn by A.P. Karremans, based on Karremans 84 (JBL-Spirit).

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LANKESTERIANALANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.30 ETYMOLOGY : Referring to the concave shape of the lip. HABITAT IN COSTA RICA : Epiphytic in tropical wet forest along the Central Valley at about 700-800 m elevation. ADDITIONAL COSTA RICAN MATERIAL STUDIED : Cartago: Lmite entre Turrialba y Jimnez, La Suiza, Pejivalle, camino a Esperanza, en lomas cerca de la Quebrada Puente, 9.0N 83.0 W, 738 m, bosque secundario a la orilla de caaverales, 1 mayo 2008, D. Bogarn 4847, A.P. Karremans, Y. Kisel & R. Phillips (JBL-Spirit!). Same locality and date, D. Bogarn 4848, A.P. Karremans, Y. Kisel & R. Phillips (JBL-Spirit!; between Ro Taus and Quebrada Azul, elev. 760 m. Epiphyte in large clumps. Flowers green. 28 May 1972, R.W. Lent 2553, (CR!, F, NY). Heredia: Sacramento, marzo 1983. Floreci en cultivo en el Jardn Lankester en julio de 1990, Mora s.n., (USJ!); San Jos de la Montaa, 15 marzo 1981, Mora s.n., (USJ!). Without collecting data, Jardn Botnico Lankester, received 18 March 1982, pressed cult. 7 April 1983, Hgsater 6731 (AMO), idem. pressed cult. 4 September 1984 (AMO; INB!), idem. pressed cult. 18 September 1986 (AMO), idem. pressed cult. 23 November 1992 (AMO), idem. pressed cult. 18 September 1996 (AMO). Epidendrum concavilabium was traditionally considered a synonym of E. circinatum Ames (Hgsater et al. 2003; Jimnez & Hgsater 2008). Although their habits are similar, plants of E. concavilabium are much more robust, more compact and thicker, have wider leaves (length:width ratio 2-3:1 vs. 4-5:1) that are darker (even in herbarium material). Both have a concave, pandurate lip, but that of E. concavilabium is much more deeply concave and shorter (19-23 mm vs. 30-32 mm). The two species have completely retrorse sepals and petals, which are similar in length (lateral sepals 16-19 mm, petals 14-15 mm); however, those of E. concavilabium are wider (lateral sepals 10-11 mm vs. 7-8 mm and petals 6-7 mm vs. 3.5-4 mm), and generally more obtuse (vs. acute). Additionally, the column of E. concavilabium is shorter, almost as wide as long, whereas that of E. circinatum is clearly longer than wide. Aside from morphological characters, E. circinatum is typically found in the whereas E. concavilabium grows at higher elevations in the mountainous areas around the Central Valley. NOTE : Since E. concavilabium was relegated to the synonymy of E. circinatum by Hgsater et al. (2003) and Jimnez & Hgsater (2008), we also list the studied specimens of E. circinatum En los manglares frente a la costa, 0 m de altura. 18 febrero 2004, A.P. Karremans 84 4 & 16-D). Costa Rica. Limn: Cantn de Talamanca, Dapari y Ro Lari, bajando hasta el cauce del mismo, corola verde-blanco, columna verde. 3 marzo 1992, G. Herrera 5159 (CR, AMO, MO, INB!). Alajuela: canopy, dense understory, broken lava blocks covered w/duff. April 22, 1990. V.A. Funk 10809, F.O. Smith, G.S. McKee and others data, JBL-s.n. (digital photograph, 14C). We have not been able to see material from Hgsater 6810 (AMO) and Lankester 844 (AMES), cited by Jimnez & Hgsater (2008) under the Costa Rican specimens of E. circinatum, but based on their locality they probably are E. concavilabium. 4. Epidendrum cystosum 2(9):105. 1934. TYPE : Honduras. Yoro: Bajo Grande, 3000 ft, 14 March 1934, J. B. Edwards 675 (holotype: AMES; photo of type at AMES!; illustration of type!). DISTRIBUTION : Mexico, Belize, Honduras, Nicaragua, Costa Rica and Colombia. ETYMOLOGY reference to the prominent ventral vesicle behind the perianth. HABITAT IN COSTA RICA : the only known specimen was found growing as an epiphyte on a solitary tree in open pastures, in tropical wet forest, just above sea level around Drake Bay in the Pennsula de Osa. COSTA RICAN M AT E RIAL STUDI E D : Puntarenas: Osa, Sierpe. Baha Drake, frente a Finca Maresa, en rboles de potrero, bosque muy hmedo tropical, 8.9 N 83.5 W, 85 m, 19 marzo 2011, A.P. Karremans 3744 & M. Contreras

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica31 FI G UR E 5. Epidendrum cystosum below and spread to the right. D Column and lip, lateral view. E Column, lateral view, front view, and longitudinal section (from left to right). F Pollinarium and anther cap. Drawn by C. M. Smith based on Karremans 3744 (JBL-Spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.32 LANKESTERIANA This unique species is distinguished by the thin than the apical leaf, with partly spreading petals and sepals, the apices of the sepals recurved, the elliptical to suborbicular lip with evident laminar keels, and the prominent clinandrium-hood with the margin erose. Epidendrum cystosum is most similar to E. macroclinium Hgsater, but the latter has succulent, with a single low keel, and a prominent clinandriumbe related to E. physodes Rchb.f., which can be distinguished by the wider, lanceolate, short-acuminate lip transversely elliptic, with a single low keel, and an obsolete clinandrium-hood with a thick and crenate margin (Santiago & Hgsater 2010). 5. Epidendrum sandiorum Hgsater, Karremans & L.Snchez, nothosp. nov. TYPE : Costa Rica. Puntarenas: Coto Brus, Sabalito, Zona Protectora Las Tablas, 10 km al noreste de Lucha, Sitio Coto Brus, camino a la Finca de Miguel Sand, 8.4 N 82.9 W, 1862 m, bosque muy 2011, A. P. Karremans 2781 & D. Bogarn (holotype, Planta inter Epidendrum ciliare L. et E. oerstedii Rchb.f. quasi intermedia et verisimiliter ex hybridatione harum specierum orta, cum Epidendro ciliare sed lobulo apicali labelli ad medium dilatato, lanceolato, clavato, manifeste acuminato, margine vix eroso, cum Epidendro oerstedii sed lobulis lateralibus apicali labelli longiore et aliis characteribus inter parentes mediis. Epiphytic, sympodial, caespitose herb, up to 20 cm tall. Roots Stems thickened into a sub-spherical to ovoid homoblastic pseudobulb, 4.0-6.0 1.5-3.0 cm. Leaves 1 at the apex of the pseudobulb, coriaceous; blade ovate-elliptic, bilobed, 7.5-12.0 3.5-6.0 cm. Spathaceous bract lacking. apical, racemose, born from peduncle laterally compressed, 2.5 cm long; covered by triangular, obtuse bracts. Floral bract about half the length of the ovary, triangular, acuminate, 3.5 cm. Ovary length of the sepals, 6.5 cm. Flowers simultaneous, resupinate, sepals and petals yellowish green, lip white, column white turning green close to the base, age. Sepals spreading, narrowly elliptic-lanceolate, acuminate, 5-7 veined, margin entire, revolute, 5.5 0.9 cm. Petals incurved, embracing the column and lip, linear-lanceolate, acuminate, 5-veined, margin entire, 5.2 0.8 cm. Lip basally united to the column, 3-lobed, base truncate; bicallose, the calli laminar, prominent, narrowly elliptic, 5 mm; disc with numerous evident veins; lateral lobes obliquely oblong, inner margin 20 6 mm; mid-lobe separate from the lateral lobes by deep sinuses, lanceolate-clavate, widened beyond the middle and that portion trullate, acuminate, margin shallowly erose, 45 7 mm. Column straight, dilated towards the apex, 1.6 cm long; clinandriumhood apical, cleft, forming a slit-like aperture anther ovoid, 4-celled. Pollinia 4, obovoid, the inner margin straight, laterally compressed. DISTRIBUTION : Known only from Costa Rica; however, as it was found a few km from the border it could also occur in Panama. ETYMOLOGY : The name honors Miguel Sand and his family; the plant that served as the type was collected on the road leading to their property. HABITAT IN COSTA RICA : This natural hybrid is only known from the very humid lower montane forests of an elevation of around 1900 m. PHENOLOGY : Flowering at least in August and September in cultivation. Several species of Epidendrum (e.g., E. ciliare L. E. falcatum Lindl., E. nocturnum L., E. oerstedii Rchb.f., and E. parkinsonianum Hook.) have starwhite lip. They were traditionally considered close

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica33relatives, but DNA studies (Hgsater & Soto 2005b) Epidendrum, and that probably their pollination by nocturnal sphyngid features. The fact that they may all be pollinated by the same type of moth has led to occasional natural hybrids, such as E. parkinsonianum E. falcatum (Hgsater 1990) and Epidendrum dorotheae P.H. Allen (Hgsater & Snchez 2008a). The latter case is interesting because the putative parent species belong to different groups, E. nocturnum being in the Nocturnum group and E. ciliare in the Coilostylis Group. Both putative parents of Epidendrum sandiorum, E. oerstedii and E. ciliare, are members of the Coilostylis Group that is characterized by the sympodial, caespitose plants, the stems forming a fusiform pseudobulb, with an apical, racemose, the above-mentioned morphology. The hybrid is recognized by the sub-spherical to ovoid pseudobulbs produced from the immature stem. The outer margins the mid-lobe trullate beyond the middle, 45 mm long, apically long-acuminate, and the margin erose. Epidendrum oerstedii ranges from Honduras to immature, short pseudobulb. The margin of the lip is entire, and the mid-lobe shorter (25-33 mm long), widened beyond the middle. Epidendrum ciliare is widely distributed from western Mexico (Nayarit) south to Peru and Brazil and the Antilles, also elongate pseudobulb, but the outer margins of the lip widened in the middle (Snchez & Hgsater 2008b; 2010). The putative parents have not been recorded yet at the same location were the hybrid was found. 6. Epistephium ellipticum R.O.Williams & Summerh., Bull. Misc. Inform. Kew 1928(4): 145. 1928. TYPE : Trinidad: Valencia Road, Mora forest end, Sept. 1926, Freeman, William & Cheesman s.n. (holotype, TRIN no. 11324; isotype, K). DISTRIBUTION : Belize, Costa Rica, Venezuela, Trinidad, Guyana and Peru. Likely occurs (but not yet collected) in Colombia, Ecuador, Panama, and other Central American countries. ETYMOLOGY : From the Latin ellipticus, elliptic, in reference to the elliptic leaf shape of the type specimen. HABITAT IN COSTA RICA : Known from a single collection in the coastal lowlands of the Caribbean close to the Panamanian border. The specimen label does not describe the habitat, but species of Epistephium grow terrestrially, typically in open, grassy areas (Cameron 2003). COSTA RICAN MATERIAL STUDIED : Limn. Talamanca, Sixaola, Gandoca, El Llano entre Filas Manzanillo y Rio Mile Creek. 09 N, 82 W, 50-100 m. 27 Mar. 1995. Terrestre. Margen de la hoja liliceos. Flor lila morado. G. Herrera 7605 & E. Sandoval McCarthy Heretofore, E. ellipticum had been recorded for Belize, Venezuela, Trinidad, Guyana and Peru; it had not been recorded from anywhere in Central America outside of Belize. The Costa Rican record here reported was collected a few kilometers from the Panamanian border, and the species likely occurs in Panama as well. The duplicate specimen at MO had E. ellipticum by Robert L. Dressler in 2007. Epistephium ellipticum is a short-statured herb and so the plants are likely overlooked and undercollected. In addition, because of their reticulate leaf venation, herbarium collections are likely to be erroneously assigned to other plant families (e.g., Convallariaceae, or when the perianth has been damaged or lost. It is possible that E. ellipticum is more widespread along the Caribbean lowlands of Central America than what the few available collections suggest. Epistephium ellipticum is also known from Peru from a single collection made in the Amazonian lowlands of the department of Loreto (Beltrn & Foster 567, F). No collections of the Amazonian lowlands of Colombia and Ecuador are known to us. The size variation of the three plants included in the herbarium sheet at CR (all of them only 8 cm tall, whereas the third plant is 23 cm tall.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.34 LANKESTERIANA FI G UR E 6. Herbarium specimen of Epistephium ellipticum R.O.Williams & Summerh. (Herrera 7605 & Sandoval McCarthy, CR).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica357. Lepanthes kabebatae Bogarn, Karremans & Mel. Fernndez, sp. nov. Type: Costa Rica. Cartago: Turrialba, La Suiza, Llanos del Quetzal, ca. 1 km sobre el camino detrs de la Escuela de Kabebata (Alto Quetzal), 9.6N secundario, bosque muy hmedo premontano, supra arbores in nemoribus Llanos del Quetzal ad Turrialba in Cartago, 17 junio 2011, D. Bogarn 8873, M. Fernndez & A.P. Karremans (holotype, CR; isotype, A Lepanthes eleganti Luer petalis latioribus luteis basaliter scarlatis, lobo superno petalorum oblongovato, lobo infero petalorum angusto-ovato, labello scarlato minute ciliato et appendice longiore statim dignoscenda. Epiphytic, caespitose, erect herb, up to 20 cm tall. Roots Ramicaul erect to suberect, up to 12.5 cm, enclosed by 8-11 minutely ciliate, blackish, lepanthiform sheaths, lightly ciliate especially on new growth, the ostia dilated, ciliate, ovate, acute, adpressed. Leaves subcoriaceous, adaxially green, abaxially purple, elliptic to narrowly ovate, acute to acuminate, with an apiculus, 4.3-7.2 1.5-3.6 cm, the cuneate base narrowing into a petiole than the leaves, up to 4.5 cm, peduncle 2 cm long, rachis 2.5 cm. Floral bracts 1 mm long, ciliate, ovate, acuminate, conduplicate, membranaceous, muriculate. Pedicel 5 mm long, persistent. Ovary up to 3 mm long, glabrous. Flowers with the sepals light yellow, basally light red, petals yellow, basally scarlet, the lip scarlet. Dorsal sepal ovate, acute, connate to the lateral sepal for about 1 mm, 4.7 5.1 mm. Lateral sepals ovate to elliptic, acute, connate for about 2 mm, 4.3-4.9 mm 1.9-2.3 mm. Petals minutely pubescent with the margins minutely ciliate transversely bilobed, 1.8-2 5.6-8 mm, the upper lobe oblong-ovate, obtuse, the apex folded, the lower lobe, narrowly ovate, acute, slightly smaller than the upper lobe. Lip bilobate, adnate to the column, 1.8-2 mm 4.2-5 mm, the blades oblong, minutely pubescent, with rounded ends, falcate, the connectives terete, up to 1 mm long, the body oblong, connate to the base of the column, the appendix oblong, pubescent. Column cylindrical, up to 2 mm long, the anther apical, the stigma ventral; anther cap cucullate. Pollinia two, ovoid. DISTRIBUTION : Known only from the type locality in Costa Rica. ETYMOLOGY : After the Cabcar kabe, quetzal, and bata, slope, referring to the locality of the Alto Quetzal (Kabebata), where the type specimen was collected. The Cabcar is one of the six native languages of the Chibcha language family spoken in Costa Rica. HABITAT IN COSTA RICA : Epiphytic in secondary and primary vegetation in premontane wet forest on the Caribbean slopes of the Cordillera de Talamanca, northwestern Fila de Matama at around 1500 m elevation. PHENOLOGY in cultivation. PARATYPES : Costa Rica. Same locality and date as the holotype, D. Bogarn 8875, M. Fernndez & A.P. Karremans and date, A.P. Karremans 4278, D. Bogarn & M. Fernndez (JBL-spirit!; CR!). Same locality and date, A.P. Karremans 4279, D. Bogarn & M. Fernndez (USJ!). Lepanthes kabebatae is one of the largest species of the genus in Costa Rica, with plant up to 20 cm tall. L. kabebatae are similar to those of L. elegans, but the former can be recognized by the wider petals 5.6-8 mm (vs. up to 5 mm) with the upper lobe oblong-ovate (vs. widely ovate) and the lower lobe narrowly ovate (vs. obliquely triangular), the microscopically ciliate lip (vs. long ciliate lip towards the apex) and the large appendix (vs. reduced to a tuft L. kabebatae the petals are yellow, basally stained with scarlet (vs. yellow with a red stain in the middle of the upper lobe and along the external margins) and the lip is scarlet (vs. yellow). The size and habit of L. kabebatae are similar to those of L. atrata Endrs ex Luer, L. barbosae Luer, L. daniel-jimenezii Bogarn & Pupulin, L. ferrelliae Luer and L. guardiana Endrs ex Luer. However, the new species lacks the thick, protuberant body of the lip present in all those species.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.36 LANKESTERIANA FI G UR E 7. Lepanthes kabebatae Bogarn, Karremans & Mel. Fernndez. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Lip, spread. F. Pollinarium and anther cap. Drawing by D. Bogarn and A.P. Karremans, based on the plant used as type (JBL-Spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica379. Lepanthes psyche Luer, Phytologia 55: 192. 1984. Type: Panama. Chiriqu, epiphytic in small trees near Volcn, La Cordillera, alt. 1300 m, 9 Dec. 1983, C. Luer, J. Luer, A. Maduro & H. Butcher 9317 (holotype, SEL). Lepanthes setos Luer, Phytologia 55(3): 193-194. 1984. Type: Panama. Prov. of Chiriqu: epiphytic in scrubby trees near Volcn, alt. 1350 m, 7 Dec. 1983, C. Luer, J. Luer & H. Butcher 9279 (holotype, SEL). DISTRIBUTION : Costa Rica and Panama. ETYMOLOGY : From the Greek psyche reference to the large petals. HABITAT IN COSTA RICA : Epiphytic in secondary premontane wet and transition to wet forests in Talamanca at around 1500 m elevation. COSTA RICAN MATERIAL STUDIED : Puntarenas: Coto Brus, Sabalito, Cotoncito, Sitio Cotn, 8.07 N 82.55 W, 1545 m, en rboles a orillas de una en el Jardn Botnico Lankester, 7 abril 2011, D. Jimnez sub D. Bogarn 8538 14H). San Jos: Dota, San Lorenzo de Dota, camino que comunica esa localidad con el proyecto Pirrs, parte ms alta del trayecto, rboles de Ciprs en cerca de potreros, 9.72N 84.88W, recolectada por D. Jimnez, M. Fernndez 334 (JBL-spirit!). This species is closely related to Ames. Vegetatively they are similar, both having thick, heavy, concave, coppery leaves with pendent stems. species. However, L. psyche is distinguished from L. and the petals with the upper lobe suborbicular to widely obovate (vs. ovate). Lepanthes setos Luer from Panama is considered a synonym of (Luer 2003). 10. Lepanthes regularis Luer, Lindleyana 2: 210. 1987. Type: Panama. Prov. of Chiriqu: Cerro Punta, Las Nubes, collected by A. Maduro 6-B, C. Luer 11631 (holotype, MO). DISTRIBUTION : Costa Rica and Panama. ETYMOLOGY : From the Latin regularis, regular, standard, referring to the average morphology of this species. HABITAT IN COSTA RICA : Epiphytic in disturbed forest in pastures in lower montane rain forests in the southern around 1800 m elevation. COSTA RICAN MATERIAL STUDIED : Puntarenas: Coto Brus, Sabalito, Zona Protectora Las Tablas, 13 km al noreste de Lucha, Sitio Coto Brus, entre Ro Sur y Quebrada Sut, Finca de Miguel Sand, 8.1 N 82.9 W, 1778 m, bosque pluvial montano bajo, D. Bogarn 7767 & A.P. Karremans Puntarenas: Coto Brus, Sabalito, Zona Protectora Las Sand El Capricho, 8.1 N 82.9 W, 1778 m, epiphytic, mostly on Quercus sp. in pastures and along the river Sut, wet premontane forest, 6 October 2010, F. Pupulin 7928, D. Bogarn, R.L. Dressler & M. Fernndez (JBL-spirit!; CR!). According to Luer (1987) the morphological features of this species are standard for the genus and no one character is particularly distinctive, it is actually the combination of standard characters that make L. regularis unique. A population of this species was found near the border between Costa Rica and Panama in the region of Las Tablas (15 km from the type locality of Las Nubes in Panama). The features are consistent with those given in the protologue (Luer 1987), especially in the ovate, acute lateral sepals, the red oblong petals, the minutely pubescent appendix plant. 11. Masdevallia nicaraguae Luer, Selbyana 5(2): 148. 1979. Type: Nicaragua. Granada: epiphytic in cloud forest on Mombacho Volcano, J.T. Atwood s.n., cultivated by J. & L. Orchids, C. Luer 2118 (holotype, SEL). DISTRIBUTION : Nicaragua and Costa Rica. ETYMOLOGY : Named after Nicaragua, the country of origin of the type specimen. HABITAT IN COSTA RICA : The only known specimen of

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.38 LANKESTERIANA FI G UR E 8. Lepanthes psyche Luer. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Lip, spread. F Pollinarium and anther cap. Drawing by D. Bogarn and C. M. Smith based on Bogarn 8538 (JBLspirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica39 FI G UR E 9. Lepanthes regularis Luer. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Lip, spread. F Pollinarium and anther cap. Drawing by D. Bogarn based on Bogarn 7767 (JBL-spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.40 LANKESTERIANA FI G UR E 10. Masdevallia nicaraguae Luer. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Petals and lip. F Column and bases of ovary and lip. G Pollinarium and anther cap. Drawn by C. M. Smith based on Acosta s.n. (JBL-spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica41M. nicaraguae was found growing in premontane wet forest, around Ro Costa Rica in Gupiles at 600 m elevation. It is likely that this species would turn up in the northern Caribbean lowlands of Costa Rica, which are continuous with the Nicaraguan lowlands where plants of this species are known to grow.. COSTA RICAN MATERIAL STUDIED : [Costa Rica: Limn: Pococ, Gupiles] Ro Costa Rica, 600 m, L. Acosta s.n. 14J). This species is similar to Masdevallia Lindl., from which it is distinguished by the thick sepaline tails, particularly those of the shorter, triangular, lateral sepals, white, faintly suffused with rose within toward the bases (vs. dull brownish yellow, spotted). Masdevallia nicaraguae is also similar to M. Ames, but is distinguished by its larger, pure white sepals, the erect tail of the dorsal sepal, the broadly dilated lateral sepals and the entire apex of the column (Luer 2001). 12. Pleurothallis instar Luer, Selbyana 3: 320, f. 260. 1977. Type: Panama. Chiriqu: epiphytic in cloud forest on Cerro Hornito, alt. ca. 1700 m, 15 Dec 1976, C. Luer 1389, A. Luer, R.L. Dressler, N.H. Williams & F.L. Stevenson (holotype, SEL; illustration of type!). DISTRIBUTION : Panama and Costa Rica. ETYMOLOGY : From the Latin instar, image, likeness, in reference to the similarity of the plant to Pleurothallis eumecocaulon Schltr. HABITAT IN COSTA RICA : Both plants of Pleurothallis instar were found growing epiphytically in the premontane rain forests of the Fila Cruces, in the COSTA RICAN M AT E RIAL STUDI E D : Puntarenas: Coto Brus, Limoncito, Fila Cruces, camino al Cerro Paraguas, ca. 10 km al oeste del Jardn Botnico R. & C. Wilson sobre el camino a Ro Claro de Golfito, 8.4 N 82.1 W, 1367 m, bosque pluvial premontano, epfitas en bosque secundario a orillas del camino, 5 junio 2010, A.P. Karremans 2776 & D. Bogarn (JBL-Spirit!; figure 11, 14K). Same locality, 20 abril 2011, D. Bogarn 8742, D. Jimnez & A.P. Karremans (JBL-Spirit!). As indicated in the protologue, Pleurothallis instar resembles mostly closely P. eumecocaulon Schltr., but it can be recognized by the ovate leaves and wider sepals and petals. Also, P. eumecocaulon bears a papillose disc located below the middle of the lip, whereas the disc in P. instar is concave and located in the basal half. Luer (2004) moved P. eumecocaulon to Ancipitia Luer but seems to have forgotten P. instar. 13. Specklinia duplooyi (Luer & Sayers) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 260. 2004. Pleurothallis duplooyi Luer & Sayers, Revista Soc. Boliv. Bot. 3(1/2): 48-50. 2001. Panmorphia duplooyi (Luer & Sayers) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 105: 153. 2006. Type: Belize. Toledo District, Little Quartz Ridge Camp, alt. 740 m, 1 August 2000, B. Sayers 997 (holotype, DBN; isotype, MO). DISTRIBUTION : Belize and Costa Rica. ETYMOLOGY : Named in honor of the late Ken DuPlooy, former director of the Belize Botanical Garden, who HABITAT IN COSTA RICA : Epiphytic on twigs of abandoned trees of Macadamia integrifolia (Proteaceae) in premontane rain forest, on the Caribbean watershed of the Cordillera de Talamanca at around 700 m elevation. COSTA RICAN MATERIAL STUDIED : Cartago: Jimnez, Pejibaye, Taus, Ro Pejibaye, 1 km despus de la escuela de Taus, 9.7N 83.4W, 707 m, bosque a orillas del ro, 30 abril 2009, D. Bogarn 6955, M. Fernndez, R. Gmez, Y. Kisel, F. Pupulin, P. Renshaw & R. Trejos (JBL-Spirit). Same locality, 16 octubre 2009, D. Bogarn 7382 & A.P. Karremans (JBLSpirit!). Same locality and date, D. Bogarn 7384 & A.P. Karremans locality, 29 de enero 2012, A.P. Karremans 4887, R.J.C.M. & I.V. Ferreira Lok (JBL-Spirit!). The species was previously known only from Belize. It is closely related to the widely distributed Specklinia barbulata (Lindl.) Luer, from which it is the leaves (vs. as long as or shorter than the leaves),

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.42 LANKESTERIANA FI G UR E 11. Pleurothallis instar Luer. A Habit; B Flower; C Dissected perianth; D Column and lip, oblique view; E Lip, ventral view; F Detail of lip apical margin, G Pollinaria. Drawn by M. Fernndez based on Karremans 2776 (JBL-spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica43 FI G UR E 12. Specklinia duplooyi (Luer & Sayers) Luer. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Column, front view. F Lip, spread. G Pollinarium and anther cap. Drawing by D. Bogarn based on Bogarn 7382 (JBL-spirit).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.44 LANKESTERIANA FI G UR E 13. Epidendrum sandiorum Hgsater, Karremans & L.Snchez. A Habit. B Flower. C Dissected perianth. D Column and lip, lateral view. E Lip, spread. F Column, lateral and ventral views. G Anther cap and pollinarium. Photographs by A.P. Karremans and F. Pupulin based on A.P. Karremans 2781.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica45 FI G UR E 14. A Acianthera oscitans ( Bogarn 7621). B Epidendrum alieniferum ( Karremans 3970). C Epidendrum circinatum ( JBL s.n.). D Epidendrum concavilabium ( Bogarn 4848). E Epidendrum cystosum ( Karremans 3744). F Lepanthes kabebatae ( Bogarn 8873). G Lepanthes kabebatae ( Bogarn 8875). H Lepanthes psyche ( Bogarn 8538). I Lepanthes regularis ( Bogarn 7767). J Masdevallia nicaraguae ( Acosta s.n.). K Pleurothallis instar ( Karremans 2776). L Specklinia duployii ( Bogarn 7382). Photographs by D. Bogarn (F., G., H., I., J. & L.), A.P. Karremans (B, D, E, K), and F. Pupulin (A, C).

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the prostrate leaves (vs. erect) and the larger a synsepal). Also, the petals are sharply acute and microscopically ciliate (vs. acute, ciliate). The lip is narrowly elliptical, long-ciliate, and longitudinally channeled (Luer 2006). The Costa Rican plants were found growing on twigs of abandoned Macadamia trees in exposed condition, an unusual habitat for plants of Specklinia. The morphology of the Costa Rican specimens is consistent with the protologue (Luer 2006). ACKNOWLEDGEMENTS services of Costa Rican Ministry of Environment, Energy and Telecommunications (MINAET) and its National System of Conservation Areas (SINAC) for issuing the in this study were collected. Franco Pupulin kindly revised the manuscript and allowed us to use some of his photographs. We thank the Vice-Presidency of Research of the University of Costa Rica for providing support under the projects 814-A7-015 Inventario y taxonoma 052, Flora Costaricensis: Taxonoma y Filogenia de la subtribu Pleurothallidinae (Orchidaceae) en Costa Rica and 814-B1-239 and Filogenia molecular de las especies de Orchidaceae endmicas de Costa Rica. We are also grateful to the personnel at CR, INB, JBL and USJ for granting full access to their collections. Archila, F. 2000. Estudio taxonmico morfolgico y delimitacin de tres gneros de la subtribu Pleurothallidinae (Orchidaceae). Revista Guatemal. 3(1): 33. Archila, F. 2010. Selbyana Archila un nuevo gnero en la Lycastinae Schltr. Revista Guatemal. 13(1): s.n. Bainbridge, C. V. & R. Aguilar. 2008. A new addition to Palmorchis nitida (Orchidaceae: Neottieae) is documented from the Osa Peninsula. Lankesteriana 8: 1. Batista, J. A. N., L. B. Bianchetti, R. Gonzlez-Tamayo, X. M. C. Figueroa & P. J. Cribb. 2011. A synopsis of New World Habenaria (Orchidaceae) I. Harvard Pap. Bot. 16: 1. Blanco, M. A. 2003. Lepanthes gerardensis (Orchidaceae), a new species from Costa Rica. Lankesteriana 8: 19 22. Blanco, M. A., G. Carnevali Fernndez-Concha, D. Bogarn & R. B. Singer. 2008. Further disentangling of a taxonomic puzzle: Maxillaria ramosa, Ornithidium pendulum, and a new species, O. elianae (Orchidaceae). Harvard Pap. Bot. 13(1): 137. Blanco, M. A., G. Carnevali Fernndez-Concha, W. M. Whitten, R. B. Singer, S. Koehler, N. H. Williams, I. Ojeda, K.M. Neubig & L. Endara. 2007. Generic realignments in Maxillariinae (Orchidaceae). Lankesteriana 7: 515. Blanco, M. A., G. Carnevali Fernndez-Concha, W. M. Whitten, R. B. Singer, S. Koehler, N. H. Williams, I.Ojeda, K. M. Neubig & L. Endara. 2008. Generic realignments in Maxillariinae (Orchidaceae): corrigenda et addenda. Lankesteriana 8: 15. Bogarn, D. 2007. A new Lycaste (Orchidaceae: Maxillarieae) from Costa Rica. Lankesteriana 7(3): 543. Bogarn, D. & M. Fernndez, 2010. Lepanthes arenasiana (Pleurothallidinae: Orchidaceae), a new species from Costa Rica. Lankesteriana 9(3): 487. Bogarn, D. & A. P. Karremans. 2010. Un nuevo Platystele (Orchidaceae: Pleurothallidinae) de la regin central de Costa Rica/A new Platystele (Orchidaceae: Pleutothallidinae) from central Costa Rica. Orquideologa 27: 208. Bogarn, D., A. P. Karremans & F. Pupulin. 2008. New records and species of Orchidaceae from Costa Rica. Lankesteriana 8(2): 53. Bogarn, D. & F. Pupulin. 2010. Two new species of Mormolyca from Costa Rica and Panama. Orchid Digest 74(1): 43. Bogarn, D. & F. Pupulin. 2011. Lepanthes daniel-jimenezii Bogarn & Pupulin, spec. nov. (Pleurothallidinae: Orchidaceae), a new species from Costa Rica close to Lepanthes guardiana. Orchidee (Hamburg) 62(2): 11115. Cameron, K. M. 2003. Epistephium A. M. Pridgeon, P. J. Cribb, M. W. Chase & F. N. Rasmussen (eds.). Genera Orchidacearum volume 3: Orchidoideae (Part Two), Vanilloideae. Oxford University Press, Oxford, U.K. Carnevali Fernndez-Concha, G., R. Duno de Stefano, G. A. Romero-Gonzlez, R. Balam, W. Cetzal Ix, J. L. Tapia-Muoz & I. M. Ramrez. 2009. A reappraisal of the turtle-orchids, genus Chelyorchis (Oncidiinae: Orchidaceae): molecular, phylogenetic, and morphometric approaches. J. Torrey Bot. Soc. 136: 164. LIT E RATUR E CIT E D LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.46 LANKESTERIANA

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Chase, M. W. & W. M. Whitten. 2011. Further taxonomic transfers in Oncidiinae (Orchidaceae). Phytotaxa 20: 26. Chase, M. W., N. H. Williams, K. M. Neubig & W. M. Whitten. 2008. Taxonomic transfers in Oncidiinae to accord with Genera Orchidacearum, Volume 5. Lindleyana 21(3) in Orchids 77: 20. Chiron, G. R. & V. P. Castro Neto. 2003. Rvision du genre Prosthechea Knowles & Wescott [sic] et nouveau genre dans la tribu Laeliinae (Orchidaceae). Richardiana 4: 9. Christenson, E. A. 2002. On the identity of Trigonidium seemannii Rchb.f. (Orchidaceae). Kew Bull. 57: 503 504. Christenson, E.A. 2003a. The genus ; a charming group of miniature Oncidium relatives. Orchids 72: 120. Christenson, E.A. 2003b. Brassia brachiata; a showy Central American species usually confused with Brassia verrucosa. Orchids 72: 296. Christenson, E. A. 2005. Drei neue Stenorrhynchos-Arten/ Three new species of Stenorrhynchos. J. Orchideenfr. 12: 5. Christenson, E. A. 2006a. Das frhere Phragmipedium caudatum aus Panama/The former Phragmipedium from Panama. J. Orchideenfr. 13: 141. Christenson, E. A. 2006b. Brevilongium un nouveau genre notropical (Orchidaceae: Oncidiinae). Richardiana 6: 45. Dressler, R. L. 1993. Field Guide to the Orchids of Costa Rica and Panama. Comstock Publishing Associates, Ithaca, New York. Dressler, R. L. 2002. The major sections or groups within Sobralia, with four new species from Panama and Costa Rica, S. crispissima, S. gloriana, S. mariannae and S. nutans. Lankesteriana 5: 9. Dressler, R. L. 2003. Orchidaceae. Pages 1 in: B.E. Hammel, M.H. Grayum, C. Herrera & N. Zamora (eds.). Manual de Plantas de Costa Rica, Volumen Monogr. Syst. Bot. Missouri Bot. Gard. 93. Dressler, R. L. 2004a. Der Scaphyglottis pulchella Komplex/ The Scaphyglottis pulchella complex. J. Orchideenfr. 11: 305. Dressler, R. L. 2004b. Una clave preliminar para las Trichopilia centroamericanas/A preliminary key to the species of Trichopilia in Mesoamerica. Orquideologa 23: 35. Dressler, R. L. 2005. Guarianthe A.M. Pridgeon, P.J. Cribb, M.W. Chase & F.N. Rasmussen (eds.). Genera Orchidacearum, Volume 4: Epidendroideae (Part One). Oxford University Press, Oxford, UK. Dressler, R. L. 2007. Sobralia macrophylla : y otras Sobralias de brcteas tubulares, con dos especies nuevas, S. rarae-avis y S. sororcula. Orquideologa 25(1): 31. Dressler, R. L. & D. Bogarn. 2007. Elleanthus ligularis, a name for a relatively common new species of Elleanthus Sect. Chloidelyna Lankesteriana 7(3): 539. Dressler, R. L. & D. Bogarn. 2009. Der Trichopilia tortilis complex (Orchidaceae: Oncidiinae) mit einer schwierig zu bestimmenden neuen art. The Trichopilia tortilis complex (Orchidaceae: Oncidiinae) with an elusive new species. Orchideen J. 2: 56. Dressler, R. L. & D. Bogarn, 2010. Some new Sobralieae from Costa Rica and Panama. Lankesteriana 9(3): 475. Dressler, R. L. & D. Bogarn. 2011. Sobralia sanctorum and Sobralia purpurella. Orchids (West Palm Beach) 80: 307. Dressler, R. L. & S. Dalstrm. 2004. A synopsis of (Orchidaceae). Selbyana 25: 1. Dressler, R. L. & W. E. Higgins. 2003. Guarianthe, a generic name for the Cattleya skinneri complex. Lankesteriana 7: 37. Dressler, R. L. & F. Pupulin. 2005. Some problems in Trichopilia (Orchidaceae: Oncidiinae), with two new 89. Dressler, R. L. & F. Pupulin. 2006. The carmine trichopilias of Central America: few but badly confused. Orchids (West Palm Beach) 75: 212. Dressler, R. L. & F. Pupulin. 2008. La identidad de Sobralia leucoxantha y una ms lejana. The identity of Sobralia leucoxantha with three new species, two closely allied and one more distant. Orquideologa 25(2): 134. Dressler, R. L. & F. Pupulin 2011. Phragmipedium Section Phragmipedium. The long-petaled tropical American ladys-slipper orchids. Lindleyana (in Orchids) 80(10): 626. Dressler, R. L., F. Pupulin & J. P. Folsom. 2006. Three new Mesoamerican taxa of Dichaea with caducous leaves. Novon 16: 336. Dressler, R. L. & N. H. Williams. 2000. Chelyorchis. In Romero, G. A. and G. Carnevali Fernndez-Concha. Orchids of Venezuela, ed. 2: 1130. Dressler, R. L. & N. H. Williams. 2003. New combinations in Mesoamerican Oncidiinae (Orchidaceae). Selbyana 24: 44. Dudek, M. & D. L. Szlachetko. 2010. New combinations in subtribe Elleanthinae (Orchidaceae, Epidendroideae). Richardiana 11: 1. Fernndez, M. 2011. A reconsideration of Trichosalpinx LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica47

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minutipetala (Orchidaceae: Pleurothallidinae). Orquideologa 28: 119-134. Fernndez, M. & D. Bogarn. 2011. A new species of Trichosalpinx (Orchidaceae: Pleurothallidinae) from 38: 41-48. Garay, L.A. 1961. Notes on the genus Epistephium Amer. Orchid Soc. Bull. 30: 496. Gerlach, G. 2004. Stanhopeinae mesoamericanae II (Orchidaceae). Dos especies nuevas: Polycycnis blancoi y Coryanthes maduroana. Lankesteriana 4: 67. Gerlach, G. & J. Beeche. 2004. Stanhopeinae mesoamericanae (Orchidaceae). III. Reestablecimiento de Stanhopea ruckeri y una especie nueva: Stanhopea confusa. Lankesteriana 4: 213. Gerlach, G. & R. L. Dressler. 2003. Stanhopeinae mesoamericanae I. Lankesteriana 8: 23. Gerlach, G. & A. Romero-Gonzlez. 2008. Stanhopeinae mesoamericanae IV: las Coryanthes de Charles W. Powell. Lankesteriana 8: 33. Hgsater, E. 1990, Epidendrum parkinsonianum Hook. in E. Hgsater and G. A. Salazar, (eds.) Icon. Orchid. 1: pl. 37. Hgsater, E. 2003. Epidendrum tolimense Lindl. (Orchidaceae), una especie sudamericana encontrada en Costa Rica. Lankesteriana 8: 41. Hgsater, E. & L. Snchez. 1993. Epidendrum lagenocolumna in E. Hgsater and G. A. Salazar, (eds.) Icon. Orchid. 2: pl. 150. Hgsater, E. & L. Snchez (eds.). 2004. The Genus Epidendrum. Part 4. A fourth century of new species in Epidendrum. Icon. Orchid. 7: pl. 701. Hgsater, E. & L. Snchez (eds.). 2006. The genus Epidendrum. Part 5. Species new & old in Epidendrum. Icon. Orchid. 8: pl. 801. Hgsater, E. & L. Snchez (eds.). 2007. The genus Epidendrum. Part 6. Species new & old in Epidendrum. Icon. Orchid. 9: pl. 901. Hgsater, E. & L. Snchez (eds.). 2008. The genus Epidendrum. Part 7. Species new & old in Epidendrum. Icon. Orchid. 11: pl. 1101. Hgsater, E. & L. Snchez (eds.). 2009. The genus Epidendrum. Part 8. Species new & old in Epidendrum. Icon. Orchid. 12: pl. 1201. Hgsater, E. & M. Soto (eds.). 2003. Orchids of Mexico. Parts 2 & 3. Icon. Orchid. 5-6: pls. 501. Hgsater, E. & M. A. Soto. 2005a. Transfers to Epidendrum L. from Oerstedella Rchb. f. Lankesteriana 5: 73. Hgsater, E. & M. A. Soto. 2005b. Epidendrum. In Pridgeon, A. M., Cribb, P. J., Chase, M. W. and F. N. Rasmussen (eds). Genera Orchidacearum, Volume 4. Epidendroideae (Part One): 236. Oxford University Press. Hamer, F. 2001. Orchidaceae Juss. In: W.D. Stevens, C. Ulloa, A. Pool & M. Montiel (eds.). Flora de Nicaragua, Tomo II. Monogr. Syst. Bot. Missouri Bot. Gard. 85: 1612. Gongora boracayanensis. Selbyana 28: 99. Karremans, A.P. In press. Illustrations and Studies in Neotropical Orchidaceae. 3. On the identity of Dracontia ramonensis (Pleurothallidinae), with a new species. Ann. Naturhist. Mus. Wien. B Karremans, A.P. & D. Bogarn. 2011. Pleurothallis adventurae (Orchidaceae: Pleurothallidinae), eine neue art aus einer unerforschten region in Costa Rica. Orchideen J. 18(3): 11114. Karremans, A.P. & E. Hgsater, 2010. Confusion in Epidendrum brenesii Schltr., and a new Costa Rican species: Epidendrum sotoanum (Orchidaceae: Laeliinae). Lankesteriana 9(3): 403. Karremans, A.P. & M. Muoz. 2011. Pleurothallis silvaepacis (Orchidaceae: Pleurothallidinae): a new species found at the Bosque de Paz Biological Reserve in Costa Rica. Orchid Rev. 121: 155. Karremans, A. P. & C. M. Smith. In press. A note on genus Dracontia (Orchidaceae: Pleurothallidinae), with a new species. Harvard Pap. Bot. Luer, C. A. 1987. New Lepanthes species from Costa Rica and Panama. Lindleyana 2: 185. Luer, C. A. 2001. Icones Pleurothallidinarum XXI. Systematics of Masdevallia part three. Monographs in Systematic Botany from the Missouri Botanical Garden. 86: 519. Luer, C. A. 2002. Miscellaneous new species in the Pleurothallidinae (Orchidaceae). Selbyana 23: 1. Luer, C. A. 2003. Lepanthes. Pp. 216 in: B. E. Hammel, M. H. Grayum, C. Herrera & N. Zamora (eds.). Manual de plantas de Costa Rica Volumen III, monocotiledneas (Orchidaceae-Zingiberaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 93. Luer, C.A. 2004. Icones Pleurothallidinarum XXVI: Pleurothallis subgenus Acianthera and three allied subgenera; A Second Century of New Species of Stelis of Ecuador; Epibator, Ophidion, Zootrophion. Monogr. Syst. Bot. Missouri Bot. Gard. 95: 1. Luer, C.A. 2005. Icones Pleurothallidinarum XXVII: Dryadella and Acronia sect. Macrophyllae-Fasciculatae (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 103: 11. Luer, C. A. 2006. Icones Pleurothallidinarum XXVIII. Reconsideration of Masdevallia and the systematics of Specklinia and vegetatively similar genera (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 105: 21. Luer, C. 2007. lcones Pleurothallidinarum XXIX. A third centry of Stelis of Ecuador and Systematics of Apoda-LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.48 LANKESTERIANA

PAGE 56

Prorepentia and Systematics of Miscellaneous small genera, addenda: new genera, species, and combinations (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Gard. 112. Luer, C. 2010. Icones Pleurothallidinarum XXXI: Lepanthes of Bolivia; Systematics of Octomeria Species North and West of Brazil; Addenda and Corrigenda. Monogr. Syst. Bot. Missouri Bot. Gard. Botany 120. Luer, C. 2011. Miscellaneous new species in the Pleurothallidinae (Orchidaceae) excluding species from Brazil. Harvard Papers in Botany 16(2): 311-360. Mytnik-Ejsmont, J. 2011. A monograph of the subtribe Oakeley, H. F. 2008. Lycaste Ida and Anguloa: the essential guide. The author, Beckenham, Kent, UK. Ojeda, I., G. Carnevali Fernndez-Concha & G. A. RomeroGonzlez. 2005. New species and combinations in Heterotaxis Lindley (Orchidaceae: Maxillariinae). Novon 15: 572. Ojeda, I., G. Carnevali Fernndez-Concha & G. A. Romero-Gonzlez. 2009. Nitidobulbon, a new genus of Maxillariinae (Orchidaceae). Novon 19: 96. Ormerod, P. 2002. Taxonomic changes in Goodyerinae (Orchidaceae: Orchidoideae). Lindleyana 17: 189 238. Ormerod, P. 2004. Notulae Goodyerinae. Oasis (Dora Creek) Suppl. 3: 3. Ormerod, P. 2005. Studies of Neotropical Goodyerinae (Orchidaceae). Harvard Pap. Bot. 9: 391. Ormerod, P. 2007. Studies of Neotropical Goodyerinae (Orchidaceae) 2. Harvard Pap. Bot. 11(2): 145. Ormerod, P. 2008. Studies of Neotropical Goodyerinae (Orchidaceae) 3. Harvard Pap. Bot. 13: 55. Ormerod, P. 2009. Studies of Neotropical Goodyerinae (Orchidaceae) 4. Harvard Pap. Bot. 14(2): 111. Pridgeon, A. M. & M. W. Chase. 2001. A phylogenetic Lindleyana 16(4): 235. Pridgeon, A. M. & M. W. Chase. 2002. Nomenclatural notes on Pleurothallidinae (Orchidaceae). Lindleyana 17: 98. Pupulin, F. 2002a. Exploring for orchids. Orchids 71: 1016. Pupulin, F. 2002b. Catlogo revisado y anotado de las Supplements (Part 1)/Additamenta ad orchideologiam mesoamericanam (pars 1). Orchidee (Hamburg) 54: 467. Supplements (Part 2)/Additamenta ad orchideologiam mesoamericanam (pars 2). Orchidee (Hamburg) 54: 561. Pupulin, F. 2003c. Two new species of Stellilabium sect. Taeniorhachis (Orchidaceae: Telipogoninae) from Costa Rica. Harvard Pap. Bot. 8: 29. Pupulin, F. 2004. A note on Prosthechea (Orchidaceae: Laeliinae), with a new species. Selbyana 25(1): 17. Pupulin, F. 2005a. Vanishing beauty: native Costa Rican orchids. Vol. 1. AciantheraKegeliella Editorial Universidad de Costa Rica, San Jos, Costa Rica. 421 pp. Pupulin, F. 2005b. Dichaea viridula (Orchidaceae: Zygopetalinae), a new species in the D. tenuifolia complex. Harvard Pap. Bot. 10(1): 83. Pupulin, F. 2005c. Ciliate dichaeas; Dichaea hystricina and Dichaea ciliolata : two species in one and an interesting variation. Orchids (West Palm Beach) 74: 678. Pupulin, F. 2006. Encyclia ossenbachiana (Orchidaceae: Laeliinae), a new species from Costa Rica. Selbyana 27(1): 4. Pupulin, F. 2007. Contributions toward a reassessment of Costa Rican Zygopetalinae (Orchidaceae). 3. A systematic revision of Dichaea in Costa Rica. Harvard Pap. Bot. 12: 15-153. Pupulin, F. & M. A. Blanco. 2002. A new species of Stellilabium section Taeniorhachis (Orchidaceae) from Costa Rica. Lankesteriana 5: 27. Pupulin, F. & D. Bogarn. 2004. Two new species of Lepanthes (Orchidaceae: Pleurothallidinae) from Costa Rica. Kew Bull. 59: 559. Pupulin, F. & D. Bogarn. 2005a. The genus Brassia in Costa Rica: a survey of four species and a new species. Lindleyana in Orchids 74(3): 202. Pupulin, F. & D. Bogarn. 2005b. The genus Scelochilus Determining the number of species in Central America. Orchids (West Palm Beach) 94(7): 526. Pupulin, F. & D. Bogarn. 2007. A second new species in the genus Restrepiella (Orchidaceae: Pleurothallidinae). Willdenowia 37: 323. Pupulin, F. & D. Bogarn. 2010a. Illustrations and studies in Neotropical Orchidaceae The Lepanthes jimenezii group (Pleurothallidinae). Harvard Pap. Bot. 15(1) 111. Pupulin, F. & D. Bogarn. 2010b. The genus Campylocentrum (Orchidaceae: Angraecinae) in Costa Rica: a review. Harvard Pap. Bot. 15(2): 353. Pupulin, F. & D. Bogarn. 2011a. Two new Lepanthes from Costa Rica. Lindleyana in Orchids (West Palm Beach) 80(3): 169. species of Lepanthes from Costa Rica. Orchidee (Hamburg) 62(6): 469-477.LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica49

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Pupulin, F. & D. Bogarn. 2012. Lepanthes novae Tapantienses. Orchid Digest. 76(1): 20-29. Pupulin, F., D. Bogarn & M. Fernndez. 2010a. A note on Pleurothallis luctuosa (Orchidaceae: Pleurothallidinae), with a new species. Ann. Naturhist. Mus. Wien. B, 112: 239. Pupulin, F., D. Bogarn & M. Fernndez. 2010b. On the identity of Myoxanthus scandens (Orchidaceae: Pleurothallidinae), with a new species. Lankesteriana 9(3): 467. Pupulin, F., D. Bogarn & D. Jimnez. 2009. New species and records in Mesoamerican Lepanthes. Orchid Digest 73: 136. Pupulin, F., D. Bogarn & C. M. Smith. 2010. Two new species of Lepanthes from Costa Rica close to L. schizocardia (Orchidaceae: Pleurothallidinae). Lankesteriana 9(3): 423. Pupulin, F., R. L. Dressler & H. Medina. 2009. A revision (Orchidaceae: Zygopetalinae). Orchid Digest 73: 32 51. Pupulin, F. & A.P. Karremans. 2010a. Crossoglossa sotoana (Orchidaceae: Malaxideae), a new species honoring the Mexican botanist, Miguel ngel Soto Arenas. Lankesteriana 9(3): 443. Pupulin, F. & A.P. Karremans. 2010b. Two new species and a new record of Epidendrum (Orchidaceae: Laeliinae) from Costa Rica. Selbyana 30(2): 195. Pupulin, F., H. Medina & D. Bogarn. 2010. Two Lepanthes (Orchidaceae: Pleurothallidinae) with a reduced corolla. Orchideen J. 17(3): 117. Pupulin, F. & G. Merino. 2008. Two new species of Kefersteinia (Orchidaceae: Zygopetalinae). Willdenowia 38: 1. Pupulin, F. & G. Rojas. 2006. A new species of Sigmatostalix (Orchidaceae: Oncidiinae) from Costa Rica, and its concolorous form. Lindleyana in Orchids 76: 677 683. Pupulin, F., G. Rojas & J. D. Ziga. 2007. Three new species of Acianthera (Orchidaceae: Pleurothallidinae) from Costa Rica. Harvard Pap. Bot. 12(1): 163. Pupulin, F. & J. D. Ziga. 2007. The upside-down Pleurothallis of Mesoamerica, with a new species from Costa Rica. Lindleyana in Orchids 77: 834. Romero-Gonzlez, G. A. & G. Carnevali FernndezConcha. 1999. Notes on the species of Cyrtopodium (Cyrtopodinae, Orchidaceae) from Florida, the Greater Antilles, Mexico, Central America and northern South America. Harvard Pap. Bot. 4: 327. Romero-Gonzlez, G. A. & C. H. Dodson. 2010. A la tercera se gana: the validation of Benzingia (Orchidaceae: Zygopetalinae). Lankesteriana 9: 526. Romowicz, A. & D. L. Szachetko. 2006. Genera et species orchidalium. 12. Oncidieae. Polish Bot. J. 51: 43. Rutkowski, P., J. Mytnik & D. L. Szlachetko. 2004. New taxa and new combinations in Mesoamerican Stenorrhynchidinae (Orchidaceae, Spirantheae). Ann. Bot. Fenn. 41: 479. Salazar, G. A. 1990. Cattleya aurantiaca. Icon. Orchid. 1: plate 7. Snchez, L. & Hgsater, E. 2007. Rchb.f. in Hgsater, E. & L. Snchez Saldaa (eds.) Icon. Orchid. 9: pl. 937. Snchez, L. & E. Hgsater. 2008a. Epidendrum dorotheae P.H.Allen, in Hgsater, E. & L. Snchez Saldaa (eds.) Icon. Orchid. 11: pl. 1119. Snchez, L. & E. Hgsater. 2008b. Epidendrum oerstedii Rchb.f. in Hgsater, E. & L. Snchez Saldaa (eds.) Icon. Orchid. 11: pl. 1151. Snchez, L. & E. Hgsater. 2010. Epidendrum ciliare L. in Hgsater, E. & L. Snchez Saldaa (eds.) Icon. Orchid. 13: pl. 1317. Santiago, E. & E. Hgsater. 2010. Epidendrum cystosum Ames in Hgsater, E. & L. Snchez Saldaa (eds.) Icon. Orchid. 13: pl. 1325. Smith, C. M. & F. Pupulin. In prep. Contributions to a taxonomic revision of Masdevallia (Orchidaceae: Pleurothallidinae) in Costa Rica. A note on Masdevallia attenuata Solano, R., G. A. Salazar & R. Jimnez. 2011. New combinations in Orchidaceae of Mexico. Acta Bot. Mex. 97: 49. Soto Arenas, M. A. & R. L. Dressler. 2010. A revision of the Mexican and Central American species of Vanilla Plum. ex Mill. with a characterization of their ITS region of the nuclear ribosomal DNA. Lankesteriana 9(3): 285. Szlachetko, D. L. 2003. Senghasia, eine neue Gattung der Zygopetaleae. J. Orchideenfr. 10: 332. Szlachetko, D. L. 2004a. Matriaux pour la rvision des Habenariinae (Orchidaceae, Orchidoideae) 4. Richardiana 4: 52. Szlachetko, D. L. 2004b. Matriaux pour la rvision des Habenariinae (Orchidaceae, Orchidoideae) 5. Richardiana 4: 103. Szlachetko, D. L. 2006. Genera et species orchidalium. 11. Oncidieae. Polish Bot. J. 51: 39. Szlachetko, D. L. 2007. Notes sur lalliance Stanhopea (Stanhopeinae, Vandoideae). Richardiana 7: 45. Szlachetko, D. L. & M. Kras. 2004. Notes sur le genre Habenella Richardiana 6: 33. Szlachetko, D. L. & M. Kulak. 2006. Nouvelles combinaisons dans le genre Zosterophyllanthos Szlachetko & Margnska (Orchidaceae, Pleurothallidinae). Richardiana 6: 183. Szlachetko, D. L., M. Kulak & A. Romowicz. 2006. Notes LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.50 LANKESTERIANA

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sur le genre Senghasia Szlachetko (Orchidaceae, Huntleyinae). Richardiana 6: 180. species orchidalium. 3. Polish Bot. J. 46: 113. the genera Malaxis Sol. ex Sw. and Microstylis (Nutt.) Eaton (Orchidaceae, Epidendroideae). Acta Soc. Bot. Poloniae 75: 229. Szlachetko, D. L., J. Mytnik-Ejsmont, M. Grniak & M. Maxillarieae. Polish Bot. J. 51: 57. Szlachetko, D. L., J. Mytnik-Ejsmont & A. Romowicz. 2006. Genera et species orchidalium. 14. Oncidieae. Polish Bot. J. 51: 53. Spiranthinae, Stenorrhynchidinae and Cyclopogoninae. Pp. 130 in: P. Rutkowski, D. L. Szlachetko & M. Grniak (eds.), Phylogeny and taxonomy of the subtribes Spiranthinae, Stenorrhynchidinae and Cyclopogoninae (Spirantheae, Orchidaceae) in Central and South America. Wydawnictwo Uniwersytetu genres dans le complexe Maxillaria (Orchidaceae). Richardiana 7: 26. Williams, N. H., M. W. Chase, T. Fulcher & W. M. Whitten. 2001. Molecular systematics of the Oncidiinae based on evidence from four DNA sequence regions: expanded circumscriptions of Cyrtochilum Erycina, Otoglossum, and Trichocentrum and a new genus (Orchidaceae). Lindleyana 16: 113. Williams, N. H., W. M. Whitten & R. L. Dressler. 2005. Molecular systematics of Telipogon (Orchidaceae: Oncidiinae) and its allies: nuclear and plastid DNA sequence data. Lankesteriana 5: 163. Whitten, W. M., N. H. Williams, R. L. Dressler, G. Gerlach & F. Pupulin. 2005. Generic relationships of Zygopetalinae (Orchidaceae: Cymbideae): combined molecular evidence. Lankesteriana 5(2): 87. Withner, C. L. & P. A. Harding. 2004. The Cattleyas and their relatives: the debatable Epidendrums. Timber Press, Portland, OR/Cambridge, UK.LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. KARREMANS et al. New species and records of Orchidaceae from Costa Rica51

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When Chase and others transferred orchid genera Cochlioda Lindl., Odontoglossum Kunth, Sigmatostalix Rchb.f., and Solenidiopsis Senghas into Oncidium Sw., in Lindleyana (Chase et al. 2008), based on molecular evidence (Williams et al. 2001a, 2001b, Chase et al. 2009), a rather strange situation developed, seen from a taxonomic point of view. Many different looking plants (some mistakenly from the distantly related genus Cyrtochilum Kunth) with very other that it becomes virtually impossible to visually Oncidium, and to separate it from many other genera in the Oncidiinae. In addition, some of the transferred Odontoglossum species, such as O. contaypacchaense D.E.Benn. & Christenson, O. machupicchuense D.E.Benn. & Christenson, O. pseudomelanthes D.E.Benn. & Christenson and O. rubrocallosum D.E.Benn. & Christenson, display all the critical features for belonging in Cyrtochilum (a transfer will be made) and should not have been included in the Oncidium transfer at all. A plausible explanation why this happened anyway is that no DNA data were analyzed prior to the transfer due to a lack of available plant material. It seems they were transferred because they were originally described as Odontoglossum species, like so many other and similar former odontoglossums that now reside in Cyrtochilum (Dalstrm 2001a). In a matter of speaking, this maneuver puts user-friendly and practical Oncidiinae taxonomy out of business. I therefore prefer to treat the visually recognizable species in genera Cochlioda, Odontoglossum and Solenidiopsis as a separate and single genus/clade rather than sinking them into a large waste-basket Oncidium. This is done in accordance with the same DNA data published by Chase and others. The reason behind combining Cochlioda, Odontoglossum and Solenidiopsis is to avoid having to create several new monotypic genera for species such as O. povedanum P.Ortiz (Fig. 1E, 3E), and O. tenuifolium Dalstrm (Fig. 1F, 3F), due to their apparently separate cladistic paths. These latter Andean species display many typical morphological and ecological Odontoglossum features and are therefore easiest treated as such. The oldest name for this large complex LANKESTERIANA 12(1): 53. 2012.NEW COMBINATIONS IN ODONTOGLOSSUM (ORCHIDACEAE: ONCIDIINAE) AND A SOLUTION TO A TAXONOMIC CONUNDRUM STIG DALSTRM 2304 Ringling Boulevard, unit 119, Sarasota FL 34237, U.S.A. Research Associate: Lankester Botanical Garden, University of Costa Rica and Andean Orchids Research Center, University Alfredo Prez Guerrero, Ecuador National Biodiversity Centre, Serbithang, Thimphu, Bhutan stigdalstrom@juno.com ABSTRACT. Oncidium koechliniana demonstrates a unique combination of features Cochlioda and Solenidiopsis to Odontoglossum, which is executed here. Distinguishing features to separate Odontoglossum from Oncidium RESUMEN. Oncidium koechliniana Cochlioda y Solenidiopsis a Odontoglossum, transferencias que se hacen en este artculo. La caractersticas distintiva para separar Odontoglossum de Oncidium travs de ilustraciones. KEY WORDS : DNA, Machu Picchu, Odontoglossum, Oncidium, Oncidiinae

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.54 LANKESTERIANAof species is Odontoglossum (Kunth 1816), which has nomenclatural priority. The general difference between Cochlioda and Odontoglossum is traditionally based on the bright Cochlioda, that suggest bird pollination (although no actual evidence of bird pollination has been documented, or seen by the author), as opposed to sparsely documented bee pollination for Odontoglossum (van der Pijl & Dodson 1966). Another and more distinct morphological difference is the divided stigma of Cochlioda, which consists of a single stigma variably divided into two lobes by the in-bent rostellum. The stigma of all known Odontoglossum species is more or less uniformly rounded. Odontoglossum sanguineum (Rchb.f.) Dalstrm [syn: Oncidium strictum (Cogn.) M.W.Chase & with a rounded stigmatic surface, which supports the nomenclatural transfer executed in this paper. This species has an impressive synonymy list and has jumped back and forth between genera Cochlioda, Mesospinidium Rchb.f. and Symphyglossum Schltr. Odontoglossum based on molecular evidence (Dalstrm 2001, Williams et al. 2001a), before being lumped into Oncidium (Chase & Williams 2008) under the name Oncidium strictum (from the synonym Cochlioda stricta Cogn.), since the name Oncidium sanguineum was already occupied. Solenidiopsis show a similar Cochlioda (as do the distantly related members of genera Oliveriana Rchb.f., and Systeloglossum Schltr.), but lack the bright colors, which may indicate a return to a bee pollination syndrome. Traditionally, the three here accepted species of Solenidiopsis; S. galianoi Dalstrm & Nuez, S. peruviana (Schltr.) D.E.Benn. & Christenson, and S. tigroides (C.Schweinf.) Senghas (Fig. 1H, 3H), all as opposed to Cochlioda and Odontoglossum, which these genera are very similar both vegetatively and in Cochlioda (rosea) was actually described as Odm. roseum by Lindley (1845), and later transferred to Cochlioda by Bentham and Hooker (1881). Similarly, Solenidiopsis tigroides was also originally described as an Odontoglossum by Schweinfurth (1945). In other words, the link to Odontoglossum has always been strong for these plants. Until now, however, it has been possible to separate them as distinct genera based on combinations divided stigma etc. This is no longer possible due to the discovery of from the Machu Picchu sanctuary in Peru; Oncidium koechliniana Collantes & G.Gerlach (2011), (Fig. 1I, 3I). Although described as an Oncidium due to the recent transfers by Chase and others (Gerlach, pers. comm.), this species displays all the features that justify a placement in an expanded Odontoglossum clade. It lowermost, like Cochlioda and Odontoglossum. It has a divided stigma similar to Cochlioda and Solenidiopsis. The coloration (greenish yellow with brown spots) is very much like an Odontoglossum or Solenidiopsis. The column has large spotted Solenidiopsis-like wings, but also a well developed hood, like Cochlioda and species in the Odontoglossum astranthum Linden & Rchb.f. (Fig. 1D, 3D) complex. It has a richly pubescent callus like many species in all three Odontoglossum species, and long and narrow leaves like some Odontoglossum and Solenidiopsis species. Although the exact position in a molecular based unknown, there is little doubt about its nomenclatural status as an (expanded) Odontoglossum. Distinguishing features for Odontoglossum Higher altitude (rarely lower than 1500 -3000 m), strictly Andean (here including Sierra Nevada de Santa Marta) plants with relatively thin roots and glossy, distinctly compressed (ancipitous) pseudobulbs (except for Odontoglossum praestans Rchb.f. & Warsz., which has weakly edged, walnutshaped pseudobulbs), and generally strongly scented Odontoglossum crispum Lindl. and possibly O. nobile Rchb.f.), with an ovary that continues in a more or less straight line into the base, and to a various extension of the variously elongate column. The lip is generally parallel with the column

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DALSTRM Nomenclatural transfers in Odontoglossum55 FIGURE 1. Lateral views of lip-column structures. A Odontoglossum cristatum Lindl. (OIC 7394, SEL). B O. blandum Rchb.f. (Dalstrm 2486, SEL). C O. mirandum Rchb.f. (Dalstrm 987, SEL). D O. astranthum (Dalstrm 925, SEL). E O. povedanum P.Ortiz (Escobar s.n. SEL). F O. tenuifolium Dalstrm 2019, SEL). G O. noezlianum (Dalstrm 2611, SEL). H O. tigroides (Dalstrm 2483, SEL). I O. koechlinianum view (Fernndez 37, USM). All illustrations by the author.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.56 LANKESTERIANA FIGURE 2. Lateral views of lip-column structures. A Oncidium mantense Dodson & R.Estrada (Estrada s.n., SEL). B O. hastilabium (Lindl.) Beer (Dalstrm 939, SEL). C Oncidium cf. citrinum Lindl. (OIC 15210, SEL). D O. andradeanum Dodson & D.E.Benn. (OIC 12087, SEL). E O. toachicum Dodson (OIC 5786). F O. hapalotyle Schltr. (OIC 4059, SEL). G O. bryolophotum Rchb.f. (Alfaro 25500, SEL). H O. acinaceum Lindl. (Harling 11265, SEL). All illustrations by the author.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DALSTRM Nomenclatural transfers in Odontoglossum57near the base and with few exceptions connected with the column either by lateral seams or by a central and longitudinal ridge (suture), and very differently from the main bulk of Oncidium species. The column of Odontoglossum has a pollinarium with a well developed and elongate stipe, placed on a relatively huge ovate viscidium, sometimes as long as the stipe. The placement and shape of the pollinarium generally presents the viscidium from above, and hidden from a frontal view, as opposed to Oncidium species, where the pollinarium generally consists of a much smaller and rounded viscidium positioned so that it is clearly visible from a front view (See Figures 1-3). There are a very few exceptions from this where some Central American Oncidium species have developed a similar morphology (and pollination syndrome?), although in these cases, the distinct geographic distribution should make it easy to distinguish the groups. In addition, very few (if any) of these intermediate Oncidium species display both a hidden and Odontoglossum-shaped ovate viscidium. Concerning the taxonomic status of a small group of species, such as Oncidium chrysomorphum Lindl., O. tipuloides Rchb.f. and O. hapalotyle Schltr. (syn: O. trinasutum Kraenzl.), etc., that seem to wobble somewhere between Odontoglossum and Oncidium depending on which molecular cladogram is studied, it is uncertain at this time how to best handle them. Some species in this group (herbarium specimens) are An attempt to rehydrate these latter specimens for more data is clearly needed in order to give them a stable home in the DNA trees. New combinations and list of transferred species Odontoglossum galianoi (Dalstrm & P. Nuez) Dalstrm, comb. nov. Basionym: Solenidiopsis galianoi Dalstrm & P. Nuez, Selbyana 23: 197. 2002. Oncidium galianoi (Dalstrm & P. Nuez ) M.W.Chase & N.H.Williams, Lindleyana 21(3): 24. 2008. TYPE : Peru. Cuzco: Province of Paucartambo, District of Challabamba, the Biosphere Reserve of Manu, between Pillahuata and Nueva Esperanza, alt 2800-3200 m, 18 Feb. 2001, P. Nuez, W. Galiano, E. Suclli, A. Rodriguez & F. Carazas 28694 (holotype, CUZ). Odontoglossum koechlinianum (Collantes & G.Gerlach) Dalstrm, comb. nov. Basionym: Oncidium koechliniana Collantes & G.Gerlach., OrchideenJournal (2): 79-81. 2011. TYPE : Peru. Cusco: Province of Urubamba, District of Machu Picchu, Quebrada Alccamayo, alt. 2500 m, 11 Dec. 2003, Moiss Quispe & Carmen Soto 148. Flowered in cultivation at Inkaterra, Machu Picchu (holotype, USM). Odontoglossum mixturum (Dalstrm & Snnemark) Dalstrm, comb. nov. Basionym: Cochlioda mixtura Dalstrm & Snnemark, Selbyana 22(2): 135. 2001. Oncidium mixturum (Dalstrm & Snnemark) M.W.Chase & N.H.Williams, Lindleyana 21(3): 25. 2008. TYPE : Bolivia. Chapare, along road between Cochabamba and Villa Tunari, in wet cloudforest at 2000 2100 m, Mar. 7, 1998, S. Dalstrm & J. Snnemark 2342 (holotype, SEL). Odontoglossum noezlianum Mast., Gard. Chron., III, 8: 570. 1890. Basionym: Lindl., Fol. Orch. 4 (Cochlioda): 1. 1853; [non Odontoglossum Lindl. = (Lindl.) Kraenzl., Notizbl. Bot. Gart. BerlinDahlem 7:99. 1917]. (Lindl.) Rchb.f., Gard. Chron. 12: 393. 1872. TYPE : Peru. Amazonas: Chachapoyas, Mathews s.n. (holotype, K-L; isotype, BM, W). Cochlioda noezliana Rolfe, Lindenia 6: 55, pl. 262. 1891. Oncidium noezlianum (Rolfe) M.W.Chase & N.H.Williams, Lindleyana 21(3): 25. 2008. TYPE : Peru. J. Ntzli s.n. (holotype, specimen unknown; illustration in original publication). Cochlioda miniata L.Lind., Lindenia, 12: 71, pl. 562. 1896. Oncidium miniatum (L.Lind.) M.W.Chase & N.H.Williams, Lindleyana 21(3): 25. 2008. TYPE : Peru[?], L. Linden s.n. (holotype

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.58 LANKESTERIANA FIGURE 3. Pollinaria, lateral views. A Odontoglossum cristatum (OIC 7394, SEL). B O. blandum (Dalstrm 2486, SEL). C O. mirandum (Dalstrm 987, SEL). D O. astranthum (Dalstrm 925, SEL). E O. povedanum (Escobar s.n. SEL). F O. tenuifolium (Dalstrm 2019, SEL). G O noezlianum (Dalstrm 2611, SEL). H O. tigroides (Dalstrm 2483, SEL). I O. koechlinianum (Fernndez 37, USM). J O. crispum Lindl. (Dalstrm 936, SEL). K O. epidendroides Kunth (Dalstrm 1496, SEL). L O. juninense Schltr. (Dalstrm 2378, SEL). M Oncidium mantense (Estrada s.n. SEL). N O. hastilabium (Dalstrm 939, SEL). O O. andradeanum (OIC 12087, SEL). P. O. toachicum (OIC 5786, SEL). Q O. hapalotyle (OIC 4059, SEL). R O. bryolophotum (Alfaro 25500, SEL). S O. acinaceum (Harling 11265, SEL). All illustrations by the author. specimen unknown; illustration in original publication). Rolfe, Orchid Rev. 19: 144. 1911. (Rolfe) M.W.Chase & N.H.Williams, Lindleyana 21(3): 24. 2008. TY PE : Peru. H. A. Tracy s.n. (holotype, specimen unknown). Cochlioda beyrodtiana Schltr., Orchis 13: 5. 1919. Oncidium beyrodtioides M.W.Chase & N.H.Williams, Lindleyana 21(3): 22. 2008. TYPE : Peru. O. Beyrodt s.n. (holotype, B, destroyed; photo at F, SEL). Odontoglossum peruvianum (Schltr.) Dalstrm, comb. nov. Basionym: Solenidium peruvianum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 9: 113. 1921, illustration in Mansf., Repert. Spec. Nov. Regni Veg. Beih. 57: t. 129, nr. 507. 1929. Solenidiopsis peruviana (Schltr.) D.E. Benn. & Christenson,

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DALSTRM Nomenclatural transfers in Odontoglossum59Brittonia 46: 44. 1994. Oncidium peruvianoides M.W.Chase & N.H.Williams, Lindleyana 21(3): 26. 2008. TYPE : Peru. Loreto: near Moyobamba, Filomeno s.n. (holotype, B, destroyed; lectotype: Tabula 129, 507). Senghas, Orchidee (Hamburg) 40(6): 205. 1989. Oncidium (Senghas) M.W. Chase & N.H. Williams, Lindleyana 21(3): 24. 2008. TYPE : Peru. Piura: near Huancabamba, ca. 2600 m, B. Wurstle s.n. BGH Nr. 0-18778 (holotype, HEID). Odontoglossum roseum Lindl. in G.Benth., Pl. Hartweg. 151. 1845. Cochlioda rosea (Lindl.) Benth. & Hook.f., J. Linn. Soc. 18: 327. 1881. Mesospinidium roseum (Lindl.) Rchb.f., Gard. Chron. 12: 392. 1872. Oncidium roseoides M.W. Chase and N.H. Williams, Lindleyana 21(3): 26. 2008. TYPE : Ecuador. Loja: Quebradas de Las Juntas, T. Hartweg 57 (holotype, K-L). Odontoglossum tigroides C.Schweinf., Amer. Orchid Soc. Bull. 14: 22, Fig. 167. 1945. Solenidiopsis tigroides (C. Schweinf.) Senghas, Orchidee (Hamburg) 37(6): 274. 1986. Oncidium tigroides (C. Schweinf.) M.W. Chase & N.H. Williams, Lindleyana 21(3): 27. 2008. TYPE : Peru. Yanano, about 1800 m, on mossy tree, Macbride 3840 (holotype, AMES). Solenidiopsis rhombicalla D.E. Benn. & Christenson, Brittonia, 46(1): 44. 1994. Oncidium rhombicallum (D.E. Benn. & Christenson) M.W. Chase & N.H. Williams, Lindleyana 21(3): 26. 2008. TYPE : Peru. Amazonas: Bongara, km 358 along road from Olmos to Jumbilla, 1450 m, July 1965, D.E. Bennett et al. 2066 (holotype, AMES; isotype, AMES). Odontoglossum vulcanicum (Rchb.f.) Dalstrm, comb. nov. Basionym: Mesospinidium vulcanicum Rchb.f., Gard. Chron. 12: 393. 1872. Cochlioda vulcanica (Rchb.f.) Benth. & Hook.f., J. Linn. Soc. 18: 327. 1881. Oncidium vulcanicum (Rchb.f.) M.W.Chase & N.H.Williams, Lindleyana 21(3): 27. 2008. TYPE : Ecuador. Tungurahua: R. Spruce 6243 (holotype, W; isotype, K-L, K). Excluded species (nomen nudum): Cochlioda chasei D.E. Benn. & Christenson, Brittonia 46: 26 (1994). Oncidium chasei (D.E. Benn. & Christenson) M.W. Chase & N.H. Williams, Lindleyana 21(3): 22. 2008. TYPE : Peru. Amazonas: Bngara, reportedly from Rio Nieva above 1700 m, 20 Nov. 1987, M. Arias ex D. Bennett & M. Chase 4080 (holotype, USM in original publication, but no type exists there!). This appears to be a confused concept based on a drawing of Odontoglossum tigroides and a lost Cochlioda specimen. According to Bennett (pers. comm.) no dried specimen was ever made. According to Chase (pers. comm.), no living material was ever introduced to horticulture by him. According to Christenson (pers. comm.), the type specimen was actually deposited at MOL, Lima, but no such dried or alcohol preserved specimen exists there either (Trujillo pers. comm., and pers. obs.). See Dalstrm (2001) for further discussion. ACKNOWLEDGMENTS The author wish to thank Jos Koechlin, owner of the Inkaterra hotels, who generously support orchid research and conservation in the Machu Picchu area. The author also wish to thank Ricardo Fernndez for assisting in making herbarium material available at the orchid herbarium of the University of San Marcos (Lima), and Wesley Higgins for assisting, reviewing and commenting on the manuscript. The authors also wish to thank the Manuel Arias Silva family together with Saul Ruiz for additional logistic support in Peru. LITERATURE CITED Bentham, G. 1881. Notes on Orchideae. In Hooker, J.D., J. Linn. Soc. 18: 327. Chase, M. W., N. H. Williams, K. M. Neubig & W. M. Whitten. 2008. Taxonomic transfers in oncidiinae to accord with Genera orchidacearum, vol. 5. Lindleyana in Orchids (West Palm Beach) 77.(12): 20-31. Chase, M.W. 2009. Subtribe Oncidiinae. In Pridgeon, A.M., M.W. Chase, P.J. Cribb, & F.N. Rasmussen

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60 LANKESTERIANA[eds]. Genera Orchidacearum, Vol. 5. Epidendroideae Oxford University Press. Collantes, B. & G. Gerlach. 2011. ber zwei neue Orchideen aus Machu Picchu (Peru) aus der Subtribus Oncidiinae. OrchideenJournal (2): 79-81. Dalstrm, S. 2001. New species and combinations in the Oncidiinae (Orchidaceae) and a synopsis of the Cochlioda clade (Oncidiinae). Selbyana 22(2): 135145. Kunth, C. S. 1816. In F. W. H. von Humboldt, A. J. A. Bonpland & C. S. Kunth, Nov. Gen. Sp. 1: 350. Lindley, J. 1845. Odontoglossum roseum. In G. Bentham, Plantae Hartwegianae: 151. Societate Linnaeana Londinensi. Schweinfurth, C. 1945. Odontoglossum tigroides. Amer. Orchid Soc. Bull. 14: 22, Fig. 167. Van der Pijl, L. & C. H. Dodson. 1966. Orchid Flowers: Their Pollination and Evolution. University of Miami Press, Coral Gables, Florida. Williams, N.H., M.W. Chase, T. Fulcher & W.M. Whitten. 2001a. Molecular systematics of the Oncidiinae based on evidence from four DNA sequence regions: expanded circumscriptions of Cyrtochilum, Erycina Otoglossum, and Trichocentrum and a new genus (Orchidaceae). Lindleyana 16(2): 113. Williams, N.H., M.W. Chase & W.M. Whitten. 2001b. Phylogenetic positions of Miltoniopsis Caucaea, a new genus, Cyrtochiloides and Oncidium phymatochilum (Orchidaceae: Oncidiinae) based on nuclear and plastid DNA data. Lindleyana 16(2):272. LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.

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Introduccin. La tribu Cranichideae Lindl. representa al grupo de las orquideas terrestres, subtropicales. Algunos de sus representantes tienen races carnosas, hojas reunidas en una roseta basal y subtribus: Achlydosinae, Chloraeinae, Cranichidinae, Galeottiellinae, Goodyerinae, Manniellinae, Prescottiinae, Pterostylidinae and Spiranthinae (Dressler 1993) con aproximadamente 210 especies en 17 gneros (Pridgeon et al. 2003) y una distribucin cosmoplita (Cribb 2003). En Suramrica tienen una distribucin restringida a pases andinos como Ecuador, Per, Colombia y Venezuela, mientras que LANKESTERIANA 12(1): 61. 2012.MORFOANATOMA EN CRANICHIDEAE (ORCHIDACEAE) DE LA ESTACIN LOMA REDONDA DEL P ARQUE NACIONAL SIERRA NEVADA, MRIDA, VENEZUELA BLANCA A. DUGARTE CORREDOR 1,2 & REBECA LUQUE ARIAS 11 Centro Jardn Botnico de Mrida, Facultad de Ciencias. Universidad de los Andes, Mrida, estado Mrida, 5101, Venezuela2 Autor para correspondencia: blancadu@ula.ve RESUMEN Se realiz un estudio morfoanatmico de los rganos vegetativos y del pednculo de tres representantes de la subtribu Cranichidinae s.l. (Orchidaceae): Aa paleacea Myrosmodes paludosa y Pterichis de la estacin Loma Redonda del Telefrico de Mrida, del Parque Nacional Sierra Nevada. Estas la presencia de caracteres adaptativos al pramo. Se recolectaron plantas adultas. Se determinaron caracteres en las paredes radiales, espirantosomas en la corteza y hojas glabras arrosetadas. Se reporta por primera vez corteza radicular; nmero de corridas de clulas de la empalizada y esponjoso; diferentes tipos de estelas en el pednculo. Algunos de estos caracteres se consideran adaptativos al ambiente: races rollizas, corteza compacta; hojas pequeas y brillantes de cutcula gruesa, desarrollo de empalizada; pednculo con epidermis y cutcula gruesa Se considera Myrosmodes paludosa morfolgicos y otra de caracteres anatmicos. ABSTRACT A morpho-anatomic study of vegetative organs and peduncle was conducted in three representatives of the subtribe Cranichidinae s.l. (Orchidaceae): Aa paleacea Myrosmodes paludosa, of the Sierra Nevada National Park in Merida, Venezuela at Loma Redonda Cable car station. These three species were studied with the aim of determining possible diagnostic characters and adaptations to the paramo. Adult exodermis scalariform with thickenings in radial walls, spiranthosomes in the cortex and glabrous clustered determined were: thickness of cortex root; number of runs of the palisade and of the spongy; in the peduncle different types of steles. Some of following characters are considered adaptive to the environment: gross root, compact cortex, small shiny leaves of thick cuticle, development of palisade; peduncle with cuticle and thick epidermis. Myrosmodes paludosa was considered the most xeromorphic species. A key using morphological and anatomical characters is presented. PALABRAS CLAVE / KEY WORDS : Aa paleacea Cranichideae, Myrosmodes paludosa, Orchidaceae, Pterichis Venezuela

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.62 LANKESTERIANAen Centroamrica se han reportado representantes de los gneros Aa (Costa Rica) (Trujillo & Delgado 2011) y Pterichis (Costa Rica y Jamaica) (Dueas & Fernndez-Alonso 2009). Los gneros Aa Rchb.f. y Myrosmodes Rchb.f. estn dentro de la subtribu Prescottiinae y el gnero Pterichis Lindl. dentro de la subtribu Cranichidinae (Dressler 1993). Ambas subtribus comparten ciertas caractersticas que las diferencian de las dems, como entre ellas porque las Prescottiinae tienen velamen tipo Spiranthes mientras que las Cranichidinae tienen velamen tipo Calanthe (Salazar et al. 2009). Figueroa et al. (2008), estudiando engrosamiento de las paredes secundarias de las clulas del velamen, tipo de tilosomas y espacios supraendodermales embargo este grupo es muy diverso y algunos representantes no han sido incluidos en ellos por ecologa tiene un papel fundamental en sus estructuras (Salazar et al. 2009). Estudios realizados por Salazar et al. dos clados altamente apoyados: las Prescottinae de elevaciones altas conformada por los gneros Aa, Altensteinia, Gomphichis, Myrosmodes, Porphyrostachys y Stenoptera y las Prescottinae de elevaciones bajas conformada por los gneros Prescottia y Pseudocranichis (lvarez 2005). Venezuela es uno de los pases ms ricos en especies de orqudeas. El Nuevo Catlogo de la Flora Vascular de Venezuela (2009) reporta 213 gneros y 1506 especies. Alrededor de 283 especies (+1 subespecie) son endmicas. El pas presenta ambientes variados como son tierras ridas, bosques en todos los pisos altitudinales, sabanas, pramos y otros (Lasser 1969). Las caractersticas morfolgicas de las plantas del pramo son por lo general de hbito arrosetado con races gruesas para almacenaje, desarrollan hojas 1974, Dahlgren 1985, Lindorf et al. 1991, Escalona 1999, Rojas 2000). Anatmicamente, las plantas desarrollan una epidermis de paredes engrosadas, cutculas engrosadas y tejidos compactos (Pyykk 1966, Rada et al. 1985, Lindorf et al. 1991, Roth 1990, Luque et al. 1999). El presente estudio pretende evaluar las caractersticas morfoanatmicas de Aa paleacea, Myrosmodes paludosa (subtribu Prescottiinae) y Pterichis de determinar las semejanzas y diferencias entre ellas y la presencia de caracteres que indiquen estrategias adaptativas al medio ambiente pramo. Este trabajo aportar datos importantes que pueden ser tomados en cuenta para el estudio de estos grupos. Materiales y mtodos R E A D E E STUDIO La estacin Loma Redonda del telefrico de Mrida se encuentra en el Parque Nacional Sierra Nevada, al sureste de la ciudad de MridaVenezuela, entre los 8 y 8N y 71 y 71W y a unos 4045 m de elevacin (Petit 1984), con una temperatura media anual de 3,0 C una precipitacin media anual de 1.553 mm y con un rgimen unimodal tropical (Monasterio & Reyes 1980). Pertenece al ncleo central de los Andes venezolanos y en la zona se observan caracteres ambientales estresantes como cambios bruscos de temperatura, humedad, hielo y deshielo, radiaciones altas y bajas, estrs hdrico y fuertes vientos casi constantes durante todo el ao. Se estudiaron dos representantes de la subtribu Prescottiinae: Aa paleacea y Myrosmodes paludosa y un representante de la subtribu Cranichidinae: Perichis (Tabla 1). claves de Garay (1978). Para los estudios anatmicos se utilizaron hojas completamente expandidas, races y una solucin de formaldedo 37%, cido actico glacial y ethanol 70% (FAA) (Johansen 1940). Se realizaron cortes transversales a mano alzada en la regin media de todos los rganos. Para separar la epidermis del Johansen (1940), tanto los cortes como las epidermis fueron coloreados con la doble tincin de azul de astra 0,5% y fuchsina bsica 0,2% segn Luque et al (1996) y fueron montados en glicerina 50% (Johansen 1940). El tamao de las clulas de los diferentes tejidos en cada rgano se obtuvieron midiendo largo por ancho de 10 clulas en tres muestras diferentes y se promediaron los valores obtenidos. Para la densidad estomtica

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae63(DE) se utiliz la formula de Salisbury (1927) segn Wilkinson (1979) y para el ndice estomtico (IE) se utiliz la frmula sugerida por Wilkinson (1979). Los datos utilizados fueron valores promedios obtenidos en tres muestras diferentes. Los cortes anatmicos fueron observados en un microscopio ZEISS AXIOSCOP 20 y se tomaron micrografas con una cmara MC80 adosada al microscopio. Los datos cuantitativos de las caractersticas anatmicas son mostrados en las tablas 2, 3 y 4. Se presenta as mismo una clave de caracteres morfolgicos y otra de caracteres anatmicos que ser til para cuando Resultados MORFOLOGA Para la caracterizacin morfolgica se describieron la raz, hoja y pednculo de cada especie y para el tamao se midieron los rganos de cada indivduo y se promediaron los valores obtenidos. Aa paleacea: hierbas terrestres de 35,0-60,0 cm de alto (Figs. 1A, 1B), crece en suelos pedregosos o en suelos con hojarasca y humus, solitarias o en grupo; races rollizas y suculentas de 4,0-7,5 cm de largo por 0,47 cm de dimetro; hojas envainadoras, dispuestas en rosetas (Fig. 1A), lanceolada hasta elptico-lanceolada, largo y 1,4-1,6 de ancho. Pednculo lateral erecto de unos 18,0-42,0 cm de largo por 0,3 cm de dimetro, revestido por numerosas bracteas (Fig. 1B; br); pequeas glabras no-resupinadas (Fig. 1A, 1B). Spalo dorsal y ptalos libres de la columna. Labio calceolado Myrosmodes paludosa: hierbas terrestres de 2,0-12,0 cm de alto (Fig. 1C-1E), crecen en suelos pedregosos o hmedos, solitarias o en grupo (Fig. 1C, 1D); races rollizas y suculentas de 4,5-8,0 cm de largo por 0,4 cm de dimetro (Fig. 1E); hojas envainadoras, dispuestas en rosetas (Fig. 1C), elptico-oblanceolada hasta cm de largo y 1,4-1,6 de ancho. Pednculo lateral erecto de 2,0-5,0 cm de largo por 0,44 cm de dimetro, revestido por brcteas (Fig. 1E; br), con entrenudos acortados solapndose las brcteas; espiga densa de no-resupinadas (Figs. 1C-1E). Spalo dorsal y ptalos adnados al dorso de la base de la columna. Labio : hierbas terrestres de 20,0-48,0 cm de alto (Fig. 1F, 1G), crecen en suelos con hojarasca y humus, solitarias (Fig. G); races rollizas y suculentas de 3,0-7,5 cm de largo por 0,38 cm de dimetro; hojas pecioladas, dispuestas en rosetas (Fig. G), oblongos de unos 4,0-15,5 cm de largo y 2,5-2,8 cm de ancho. Pednculo terminal erecto de aproximadamente 30,0 cm de largo por 0,32 cm de dimetro glabro en la base y piloso el resto, revestido por bracteas foliosas (Fig. 1F; br); espiga subdensa de 5,0-10,0 cm de largo de (Fig. 1F, 1G). Spalos semejantes libres, el dorsal pndulo y los laterales erectos (Fig. 1F, 1G); ptalos ms angostos que el spalo dorsal. Labio spero y algo adnado a la base de la columna. Se pudo observar en campo que M. paludosa tiene un perodo de crecimiento vegetativo mayor, esto es Taxn Testigo Herbario Zona de colecta Aa paleacea (Kunth) Rchb. f. Estrada et al. 1027a MERC Al oeste de la estacin dentro del bosque Dugarte & Contreras 102 de Polylepis sericeae Suelos con Dugarte & Contreras, 106 abundante humus Dugarte & Contreras 107 Myrosmodes paludosa (Rchb. f.) Garay Estrada et al. 1023 MERC Al oeste de la estacin hacia y por los Dugarte & Contreras 100 alrededores del bosque de Polylepis Dugarte & Contreras 101 sericeae Suelos pedregosos y Dugarte & Contreras 103 encharcados Dugarte & Contreras 104 (Lindl.) Schltr. Len & Gaviria 491 MERC Al oeste de la estacin dentro del bosque Estrada et al. 1027b de Polylepis sericeae Suelos con abundante humus TABLA 1 Lista de las especies estudiadas de Cranichideae s.l.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.64 LANKESTERIANA Especie Epidermis Exodermis Corteza Parenquimtica Endodermis L (m) A (m) L (m) A (m) L (m) A (m) L (m) A (m) Aa paleacea 82,14 89,29 110,71 67,86 111,90 86,90 21,64 34,43 Myrosmodes paludosa 64,29 55,95 72,62 84,52 161,90 129,76 43,28 48,19 52,38 72,62 76,19 60,71 163,09 115,48 39,34 45,25 Especie Epidermis Adaxial Epidermis Abaxial Parenquima Empalizada Parenquima Esponjoso L (m) A (m) L (m) A (m) L (m) A (m) L (m) A (m) Aa paleacea 80,48 80,48 36,19 35,71 86,67 47,62 39,52 34,29 Myrosmodes paludosa 46,67 41,90 45,24 30,95 101,43 57,14 55,71 42,86 40,00 62,38 29,52 34,76 47,62 34,28 37,62 33,33 Especie Epidermis Adaxial Estoma Epidermis Abaxial Estoma L (m) A (m) L (m) A (m) D.E (est/mm2) I.E (%) L (m) A (m) L (m) A (m) D.E (est/mm2I.E (%) Aa paleacea 106,54 118,7 92,06 62,62 66,82 58,88 35,37 15,87 Myrosmodes paludosa 131,17 84,35 58,5 47,96 12,5 10 63,95 42,86 46,6 39,8 84,21 21,62 115,89 82,71 57,94 55,14 2,86 1,75 108,88 57 68,69 52,8 45,16 18,42 TABLA 2. Medidas de las clulas en los diferentes tejidos de la raz. L: largo, A: ancho, m: micras. TABLA est/mm2: estomas por milmetro cuadrado TABLA 3. Medidas de las clulas en los diferentes tejidos de la hoja en corte transversal. L: largo, A: ancho, m: micras.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae65 FIGURA 1. Caractersticas morfolgicas. A, B: Aa paleacea C-E: Myrosmodes paludosa. F, G: (br = bractea). Fotografas por B. Dugarte Corredor (A-E) y J. Estrada (F, G).porque las plantas se pueden conseguir solo en estado A. paleacea y P. no se encuentran en estado ANATOMA Aa paleacea: en corte transversal se evidencia la raz con velamen de una corrida de clulas (Fig. 2A, 2B; Ve) con pelos radicales (Fig. 2A; PR); interna al velamen se observa la exodermis formada por una corrida de clulas (Fig. 2A, 2B; Ex), con las paredes periclinal externa y radial engrosadas, en las paredes radiales se observa un Corteza parenquimtica, compacta de 13 a 14 corrida de clulas (Fig. 2A, 2C, 2D; P) observndose los amiloplastos (Fig. 2C; am) con uno a cuatro granos de almidn (Fig. 2C; al). Cilindro central rodeado por la endodermis (Fig. 2D; En) con bandas de Caspary (Fig.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.66 LANKESTERIANA FIGURA 2. Corte transversal de la raz. A-D: Aa paleacea E-I: Myrosmodes paludosa. J-O: (Ve = velamen; PR = pelo radical; Ex = exodermis; P = parnquima; am = amiloplasto; al = almidn; En = endodermis; bC = 2D; bC) interna a ella el periciclo de una corrida de clulas (Fig. 2D; Pe) y se observa la estela poliarca con 10-16 polos del xilema (Fig. 2D; Xp) alternando con el hoja muestra: hoja bifacial hipostomtica, epidermis paredes periclinales externas papilosas (Fig. 3A, 3B, 4A; Pl), cubierta por una cutcula ms o menos gruesa y estriada (Fig. 3B, 4A; Cut), paredes periclinales internas un poco cncavas. En vista paradermal se

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae67 FIGURA 3. Corte transversal de la hoja. A-D: Aa paleacea E-H: Myrosmodes paludosa. I-L: (Ead en empalizada; Pes = parnquima esponjoso; NM = nervio medio; Ca = cmara aerfera; N = ncleo; VP = vaina observan clulas polidricas de 5 a 7 caras, de paredes ms o menos gruesas y rectas (Fig. 4A). Epidermis de clulas cuadradas (Fig. 3A; Eab). En vista paradermal se observa formada por clulas 4-5-6 caras de paredes ms o menos gruesas, de paredes sinuosas (Fig. 4B); estomas anomocticos rodeados por 3-4 (5) clulas anexas (Fig. 4B; Est). Densidad estomtica de 45,12 est/mm2 diferenciado, compacto; parnquima en empalizada

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.68 LANKESTERIANA(Fig. 3A. 3B; Pem) formado por 1-2 corridas de clulas cortas; parnquima esponjoso de 6 corridas de clulas (Figs. 3A, 3B; Pes), se observan cmaras un ncleo grande con nucleolo (Fig. 3C; N) y estn llenas de cloroplastos (Fig. 3B, 3C). Haces vasculares colaterales cerrados (Fig. 3D) rodeados por una vaina parenquimtica simple (Fig. 3D; VP) con almidn. El pednculo en corte transversal se observa de contorno elptico, con epidermis de una corrida de clulas (Fig. 5B; Ep). Cutcula gruesa (Fig. 5B; Cut). Corteza parenquimtica muy reducida de 1 a 2 corridas de clulas (Fig. 5B; P) seguida por un anillo de 5 a 6 corridas de clulas esclerenquimticas, (Fig. 5B; Es). Internamente se observa la atactoestela con haces vasculares colaterales de mayor tamao (Figs. 5A y B; HV) rodeados de una vaina parenquimtica simple, los menores formados por xilema solamente. Se observa lisis de la zona central (Fig. 5A; *). Myrosmodes paludosa: en corte transversal se evidencia la raz con velamen de una corrida de clulas (Fig. 2E, 2G; Ve) con pelos radicales (Fig. 2F; PR); interna al velamen se observa la exodermis formada por una corrida de clulas (Fig. 2E, 2G; Ex), con las paredes periclinal externa y radial engrosadas, en las paredes radiales se observa un Corteza parenquimtica, compacta de 16 a 18 corridas de clulas (Fig. 2E, 2H; P), observndose los amiloplastos (Fig. 2H; am) con uno a cuatro observa el cilindro central rodeado por la endodermis (Fig. 2I; En) con bandas de Caspary (Fig. 2I; bC) seguida internamente el periciclo con una corrida de clulas (Fig. 2I; Pe) y se observa la estela poliarca con aproximadamente 11 polos xilemticos (Fig. 2I; El corte transversal de hoja muestra: hoja bifacial (Fig. 3E-3G; Ead) formada por clulas de paredes engrosadas ms o menos isodiamtricas, con las paredes periclinales internas un poco cncavas, cubierta por una cutcula lisa y gruesa (Fig. 3E, 3G; Cut). En vista paradermal se observan clulas polidricas de 5 a 6 caras, de paredes muy gruesas y rectas (Fig. 4C). Estomas anomocticos y diacticos rodeados por 2-3-4 (5) clulas anexas (Fig. 4C; Est). Densidad estomtica de 17,65 est/mm2, ndice abaxial, en corte transversal, formada por clulas ms o menos isodiamtricas (Fig. 3E, 3F; Eab) cubierta por una cutcula ms o menos gruesa (Fig. 3F; Cut). En vista paradermal se observan clulas de 4-5-6 caras de paredes gruesas, rectas a ligeramente sinuosas (Fig. 4D); estomas anomocticos rodeados por 3-4-5 clulas anexas (Fig. 4D; Est). Densidad estomtica de 41,18 est/mm2, ndice estomtico de 19,87 %. En vista paradermal en las clulas epidrmicas, se puede observar claramente el ncleo compacto; parnquima empalizada (Fig. 3F, 3G; Pem) formado por 4-5 corridas de clulas cortas; parnquima esponjoso de 5-6-7 (8) corridas de clulas (Fig. 3F, 3G; Pes). Nervio Medio (Fig. 3E-3G; NM) formado por un haz colateral. Haces vasculares colaterales cerrados rodeados por una vaina parenquimtica simple (Fig. 3H; VP) con almidn, se (Fig. 3E; Ca). El pednculo en corte transversal se observa de contorno elptico, con epidermis de una corrida de clulas (Fig. 5C; Ep). Cutcula gruesa (Fig. 5C; Cut). Corteza parenquimtica de 7 a 8 corridas de clulas (Fig. 5C; P) seguida por un anillo de 3 corridas de clulas esclerenquimticas (Fig. 5C; Es). Internamente se observa la estela formando anillos concntricos con haces vasculares colaterales (Fig. 5C; HV), haces del anillo ms externo se observan relacionados con el tejido esclerenquimtico. En la Figura 5D se observa con ms detalle el xilema una vaina parenquimtica simple en cuyas clulas se observan granos de almidn (Fig. 5D; al). Se observa mdula parenquimtica compacta (Fig. 5C; M) y muy notable el ncleo en cada clula (Fig. 5C; N). Pterichis multiflora: en corte transversal se evidencia la raz con velamen de una corrida de clulas (Figs. 2J y L; Ve) con pelos radicales (Fig. 2K; PR); interna al velamen se observa la exodermis formada por una corrida de clulas (Figs. 2J y L; Ex), con las paredes periclinales externa y radiales engrosadas, en las paredes radiales se

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae69 FIGURA 4.Vista paradermal de la epidermis. A,C,F: Epidermis adaxial. B,D,G: Epidermis abaxial. Aa paleacea A,B. Myrosmodes paludosa CyD. EyF. (Cut: cutcula; Est: estoma; P: papila; N: ncleo). A,B: Aa paleacea (A: epidermis adaxial; B: epidermis abaxial). C,D: Myrosmodes paludosa (C: epidermis adaxial; D: epidermis abaxial). E,F: (E: epidermis adaxial; F: epidermis abaxial). (Cut = cutcula; Est = estoma; P = papila; N = ncleo).observa un engrosamiento escalareiforme (Fig. 2L; flecha). Corteza parenquimtica, compacta de 13 a 15 corrida de clulas (Figs. 2J y M; P) observndose los amiloplastos (Fig. 2K y M; am). Es notoria la presencia de hifas (Fig. 2O; Hf). Se puede observar el cilindro central rodeado por la endodermis (Fig. 2N; En) con bandas de Caspary (Fig. 2N; bC), interna a ella el periciclo de una corrida de clulas (Fig. 2N; Pe). Estela con 8 polos xilemticos (Fig. 2N; Xp) alternando con el floema primario (Fig. 2N; Fl). El corte transversal de hoja muestra: hoja bifacial anfiestomtica, epidermis de la superficie adaxial (Fig. 3I-3K; Ead) formada por clulas rectangulares con las paredes periclinales externas papilosas (Fig. 3I 3J; Pl), las internas un poco cncavas, cubierta por una cutcula gruesa y estriada (Fig. 3J; Cut). En vista paradermal se observan clulas de 5 a 6 (7) caras de paredes gruesas, rectas a sinuosas (Fig. 4E) estomas anomocticos rodeados por 4-5 clulas anexas (Fig. 4E; Est). Densidad estomtica de 15,71 est/mm2, ndice estomtico de 6,62 %. Epidermis de la superficie abaxial, en corte transversal, formada por clulas rectangulares (Fig. 3I; Eab). En vista paradermal se observan clulas de 5 a 6 (7) caras de paredes muy gruesas, rectas a sinuosas (Fig. 4F). Estomas anomocticos rodeados por (3) 4-5 clulas anexas (Fig. 4F; Est). Densidad estomtica de 68,65 est/mm2, ndice estomtico de 24,14 %. En vista paradermal en las clulas epidrmicas, se puede observar claramente el ncleo (Fig. 4E, 4F; N). Mesfilo poco diferenciado, compacto; parnquima en empalizada (Fig. 3I-3K; Pem) formada por 2 corridas de clulas cortas; parnquima esponjoso de 7-8 corridas de clulas (Fig. 3I, 3J; Pes). Se puede notar algunas clulas con su ncleo (Fig. 3J; N). Nervio Medio (Fig. 3J, 3L; NM) formado por un haz colateral. Haces vasculares colaterales cerrados (Fig. 3I; HV) rodeados por una vaina parenquimtica simple (Fig. 3L; VP) con almidn. El pednculo en corte transversal se observa de contorno elptico, con epidermis de una corrida de clulas (Fig. 5F; Ep) con tricomas pluricelulares (Fig. 5F, 5G; Tri). Cutcula gruesa (Fig. 5F; Cut). Corteza parenquimtica de 3 a 4 corridas de clulas (Fig. 5F; P) seguida por un anillo de 5 a 6 corridas de clulas esclerenquimticas (Fig. 5F; Es). Internamente se observa la atactoestela con haces vasculares colaterales rodeados de una vaina parenquimtica simple (Fig. 5E; HV). Se observa lisis en la zona central (Fig. 5E; *).

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.70 LANKESTERIANA FIGURA 5. Corte transversal del Pednculo. A,B: Aa paleacea C,D: Myrosmodes paludosa. E,G: (Ep = epidermis; cut = cutcula; P = corteza parenquimtica; Es = clulas esclerenquimticas; HV = haces colaterales; M = mdula; N = Ncleo; al = almidn; = lisis medular; Tri = tricoma). Discusin Las especies estudiadas son plantas terrestres caracterizadas por tener hojas en rosetas basales y races gruesas de almacenaje, caractersticas representativas de la tribu Cranichideae (Dahlgren 1985). Las races rollizas y suculentas, son rganos de reserva apropiados para que estas plantas puedan vivir en las condiciones del pramo y soportar los cambios estacionales. El velamen observado es de tipo spiranthes y coincide con los tipos de velamen propuestos por Porembski y Barthlott (1988) as como tambin una de las caracterstica de los representantes de la tribu Prescottiinae como A palacea y M paludosa, no obstante el velamen que muestra P con lo descrito por Salazar (2009) donde indica que el velamen para los representantes de la tribu Cranichidinae a la cual est incluido este gnero es de tipo Calanthe. La exodermis adems del engrosamiento de las clulas en las paredes periclinal externa, muestra un engrosamiento escalareiforme en las paredes radiales. En un estudio comparativo de las tribus Cranichidae, Diceratosteleae y Tropidieae realizado por Stern et al. (1993b) este tipo de engrosamiento fue encontrado slo en especies de la tribu Cranichideae, estos autores suponen que funcionan como estructuras de rigidez que previenen el colapso de las clulas exodrmicas. Solereder & Meyer (1930) (segn: Stern et al. 1993b) describieron este tipo de engrosamiento, pero no lo ilustraron ni lo comentaron. Aunque no se trata exactamente de deposicin de bandas de Caspary, podramos considerar que este engrosamiento tambin raz tal como lo indican (Perumalla & Peterson 1986, Hose et al. 2001, Garca 2004). Este carcter fue muy

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae71til para Figueroa et al. (2007) porque les permiti delimitar al clado Prescottiinae Spiranthinae Cranichidinae. La corteza compacta evita la acumulacin de agua intercelular por lo que incrementa su resistencia a la congelacin. El nmero de corridas de clulas es menor para A paleacea, segn Fanh (1967), Jenik (1978), Lindorf (1991), un nmero menor de corridas de clulas favorecera el rpido flujo de agua al interior del cilindro central. Estas especies crecen en microambientes similares, la diferencia observada en la corteza puede deberse quizs a que M. paludosa presenta un perodo de crecimiento vegetativo mayor. Las micorrizas en la corteza de P ubicadas en las clulas ms externas de la corteza, coincide con lo observado por Stern et al. (1993b). Algunas orqudeas en condiciones naturales germinan slo despus de establecer simbiosis con el micelio de un hongo (Dalhgreen 1985). Esta interaccin ha sido considerada como un mecanismo de adaptacin que ha permitido a las orqudeas mantenerse en condiciones climticas extremas y tener una distribucin a nivel mundial (Garay 1963). Para el ao 1902 Noel Bernard (segn Arber 1961) describi que la mayora de las semillas de orqudeas no germinan naturalmente en ausencia de micorrizas y para la misma poca Hans Buergeff (Arditti 1992) demostr que la interaccin micorriza-orqudea eran (Rasmussen & Whigham 1998) que algunas especies de hongos, otras (Arditti 1992) tienen una relacin general con muchas especies de hongos. Esta variedad de tipos de interaccin micorrzica en orqudeas podra explicar el hecho de no haberse observado en A paleacea ni en M paludosa o posiblemente deba repetirse la observacin con tcnicas especializadas en micorrizas. Los amiloplastos son llamados espirantosomas por Stern et al. (1993a) y fueron observadas en las subtribu Diceratosteleae, Tropidieae y Cranichideae donde el almidn es sintetizado y reservado en amiloplastos tpicos; adems estos espirantosomas no fueron observados en ninguna Orchidoideae ni en clulas de otras orqudeas que tenan disponibles para el momento de sus estudios. Los espirantosomas son cuerpos esfricos que se encuentran en el citosol, cada uno conteniendo numerosos granos diminutos de almidn que constituiran reserva de carbohidratos. El gran desarrollo observado de las bandas de Caspary parece ser una caractersticas bastante comn en las plantas que crecen en los pramos, observndose no solamente en raz sino tambin en tallo y hoja (Luque 2000). Segn Van Fleet (1961) el desarrollo de las bandas de Caspary est fuertemente luz, temperatura, nutricin, entre otros; por lo tanto se considera que en estas especies las condiciones de cambios de temperatura diaria y exposiciones de desarrollo de grandes bandas de Caspary. La diferencia que se observa entre el nmero de polos xilemticos, siendo mayor en A paleacea, seguida de M paludosa y P hace que sea ms xilema para su rpida conduccin (Luque 2004). En cuanto a la morfologa foliar, M. paludosa presenta hojas ms adosadas al suelo, ms pequeas y brillantes y la lmina ms gruesa que A. paleacea y que presentan hoja ms alargadas y P. es la que presenta la roseta ms laxa con las hojas ms separadas del suelo; M. paludosa crece en lugares un poco ms expuestos y las caractersticas mencionadas le proporcionan rigidez, ayudan a evitar de los vientos excesivos que se da en este tipo de ambiente (Eames & MacDaniels 1947, Fahn 1974, Dahlgren 1985, Lindorf 1991, Escalona 1999, Rojas 2000). Anatmicamente la epidermis de paredes celulares interpretado como un mecanismo para el control de la transpiracin, adems sirve de proteccin contra la luz intensa, evitando que los tejidos internos puedan calentarse excesivamente; tambin ayuda a evitar los daos mecnicos producidos por el viento y previenen el colapso o marchitamiento de los tejidos de la hoja cuando hay prdida de agua en exceso (Pyykk 1966, Lindorf et al. 1991, Roth 1991, Ely 1996, Torres 1998, Escalona 1999, Rojas 2000). Las clulas papilosas de la epidermis de la tambin llamadas clulas en forma de lente, se

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.72 LANKESTERIANAconcidera que su funcin puede ser concentrar la subyacente (Gaulhofer 1908, Wilkinson 1979, Lindorf et al. 1991, Roth 1991). Plantas de las especies mencionadas crecen, en la Estacin Loma Redonda, dentro del bosque de Polylepis, por lo tanto ms protegidas, y las papilas epidrmicas favoreceran la captacin y aprovechamiento de la luz. Myrosmodes paludosa y presentan tiene de ah los valores de densidad estomtica e ndice estomtico tan bajos para esta especie en esta densidad estomtica lo que puede considerarse como una adaptacin a las altas altitudes y al estrs hdrico, la reduccin de la densidad estomtica funcionara como un mecanismo de control transpiratorio (Colmenares et al. 2005). Presentan estomas anomocticos, sin embargo, M. paludosa tambin presenta estomas diacticos, segn Stern et al. (1993 b), en los representantes de la tribu se pueden conseguir diferentes tipos de estomas, inclusive en la misma hoja como es el caso de M. paludosa. Se reporta, en este trabajo, por primera vez pertenecientes a las subtribus Prescottiinae y et al. 1999) y adems evita que se acumule agua intercelular por lo que incrementa su resistencia a la congelacin (Rada et al. 1985). El parnquima en empalizada, a pesar de estar formado por clulas casi isodiamtricas o cuadradas, se observa bien diferenciado del parnquima esponjoso que est formado por clulas redondeadas a ovaladas. Segn Haberlandt (1884), la forma de las clulas del parnquima fotosinttico puede variar desde clulas isodiamtricas, como las de las especies estudiadas, hasta clulas alargadas en sentido paralelo y tambin clulas braciformes y en forma de embudo. Algunos autores (Pyykk 1966, Esau 1976, Roth 1990, Lindorf et al. 1991, Dengler 1994, Ely 1996) consideran que clulas de la empalizada cortas, como las que presentan las especies estudiadas, son un luz, el desarrollo del parnquima en empalizada es en el suelo. Estas observaciones contrastan con lo expresado por Stern et al. (1993b), en su estudio sobre anatoma comparada de los rganos vegetativos de Spiranthoideae, consideran que los representantes de la subtribu Prescottiinae a la cual pertenecen A. paleacea y M. paludosa y representantes de la subtribu Cranichidinae al que pertenece presentan Myrosmodes paludosa presenta la mayor proporcin de parnquima en empalizada, un carcter xeromorfo importante ya que el alto porcentaje de este tejido se fundamenta en el hecho de que el desarrollo es estimulado por la exposicin a luz. Sin embargo, A. paleacea y por crecer en sitios protegidos (como es el caso del bosque de Polylepis sericea) presentan el parnquima en empalizada de una a dos corridas de clulas, el menor nmero de capas de parnquima en empalizada podra evitar la presencia de muchas paredes transversales que Lindorf 1991). Los haces colaterales cerrados son caractersticas propias en monocotiledneas (Arber 1925, Esau 1976, Tomlinson 1969). Vaina vascular parenquimtica de una corrida de clulas con almidn, que segn Alexandrov (1926), Rhoades & Carvalho (1944), es formado por los cloroplastos durante la fotosntesis (Esau 1976). Esta vaina parenquimtica tambin la consideran, en monocotiledneas, una envoltura uniestrata comparable a la endodermis (Lindorf et al. 1991). El pednculo se presenta lateral para A. paleacea y M. paludosa mientras que terminal para P. caractersticas representativas para las subtribus a las cuales pertenecen (Dressler 1993). Anatmicamente, se evidencia la epidermis de una corrida de clulas engrosadas y cutcula engrosada, este carcter favorece la proteccin contra los efectos del viento y daos mecnicos y se consideran que son caractersticas xeromorfas. Stern et al. (1993b) sealan en su estudio que las especies presentan clulas ser debida a que las especies aqu estudiadas crecen en el pramo que es un medio con condiciones ms extremas.

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LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae73 Se observa una corteza formada por un tejido parenquimtico que puede actuar como reservante de agua y solutos y un tejido esclernquimtico que est relacionado con una funcin mecnica y posiblemente como reservorio hdrico, de almidn o aceites y proporciona al tallo resistencia frente a diferentes excesos evitando que sufran dao (Esau 1976, Lindorf et al. 1991, Rojas 2000); observaciones similares fueron realizadas por Stern et al. (1993b) para los representantes de la subtribu Diceratosteleae pero no as para los de Chranichideae, estas diferencias pueden ser debidas tambin al hecho de que entre el gran nmero de especies estudiados por los autores citados no se encontraba ninguno de los pramos. Los haces vasculares colaterales cerrados caractersticos de monocotiledneas estn dispuestos ya sea en una atactostela en A paleacea y P y en 2 anillos concntricos en M paludosa, en todos los casos el anillo ms externo parece estar asociado al anillo esclerenquimtico ms interno de la corteza. Esta observacin coincide con Stern et al., (1993b), posiblemente se trate de una zona generatriz de haces que se observa tambin en otras orqudeas de pramo como Alteinstenia Kunth y Gomphichis bogotensis Renz (observacin personal). Lisis y/o rompimiento mecnico en la mdula parenquimtica de A. paleacea y En muchos tallos la mdula se destruye en forma parcial o total durante el crecimiento, segn Lindorf et al. (1991) esto ocurre debido a la expansin radial y segn Flores (1999) el colapso de debe a la presin que ejercen los nuevos tejidos. El anlisis de las plantas estudiadas indica la interpretarse como adaptativas al pramo. CLAVE MORFOLGICA 1 Hoja peciolada. Roseta basal creciendo separada del suelo. Brcteas foliosas en el pednculo. Flores amarillas con rayas rojas................................................................................................................. 1 Hoja ssil. Roseta basal creciendo adosada al suelo. Brcteas hialinas en el pednculo. Flores blancas.... 2 ....................................................................................................................................................... A. palacea entrenudos cortos............................................ M. paludosa CLAVE ANATMICA clulas. Corteza parenquimtica de la raz de 16-18 corrida de clulas, corteza esclerenquimtica del pednculo de 3 corrida de clulas....................................................................................... M. paludosa de clulas. Corteza parenquimtica de la raz de 13-15 corrida de clulas, corteza esclerenquimtica del pednculo de 5-6 corrida de clulas............................................................................................ 2 2 Hoja hipostomtica. Polos xilemticos de la raz de 10-16, corteza parenquimtica del pednculo de 1-2 corrida de clulas.............................................................................................................. A. paleacea del pednculo de 3-4 corrida de clulas..................................................................................... Claves para la separacin de los representantes de la tribu Cranichideae de la Estacin Loma Redonda del telefrico de Mrida-Venezuela con base a caracteres morfolgicos y anatmicos AG RAD E CI M I E NTOS Este trabajo se realiz en el Laboratorio de Anatoma Vegetal del Centro Jardn Botnico de la Facultad de Ciencias de la Universidad de los Andes. Los autores desean parcial a travs del proyecto N C-119503. A la gerencia del Telefrico de Mrida por permitir el ascenso a la estacin Loma Redonda y al personal por facilitar el traslado. Tambin agradecen al profesor Javier Estrada, y de manera especial a valiosa colaboracin durante su ejecucin.

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LITERATURA CITADA lvarez, A. 2005. Phylogenetics of Prescottiinae Dressler y systematics of Gomphichis lindl. (Orchidaceae). A dissertation submitted to the Graduate Faculty in degree of Doctor of Philosophy, The City University of New York. Arber, A. 1961[1925]. Monocotiledons. A morphological study. Cambridge Univ Press, Cambridge. pp 258. Arditti, J. 1992. Fundamentals of orchid biology. John Wiley & Sons, New York. Carnevali, G., G. Gerlach & G. Romero. 2005. Orchidaceae. Pp. 753-789 in: O. Hocke, P.E. Berry & O. Huber (eds.), Nuevo Catlogo de la Flora Vascular de Venezuela. Fundacin Instituto Botnico de Venezuela Dr. Tobas Lasser. Caracas-Venezuela. of dna data. The Board of Trustees of the Royal Botanic Gardens, Kew 2005. Colmenares-Arteaga, M., F. Rada & R. Luque. 2005. Anatoma foliar de Polylepis sericea Wedd. (Rosaceae) a dos altitudes en los Altos Andes Venezolanos. Plantula 3(3): 141-148. Cribb, P. 2003. Subtribe Cranichidinae, Description and in: A.C. Pridgeon, P.J. Cribb, M.W. Chase & F.N. Rasmussen (eds.), Genera Orchidaceaerum 3. Oxford University Press, Oxford. Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo. 1985. The fami lies of the Monocotyledons. Springer-Verlag, Berlin. development. pp. 100-136 in: M. Iqbal (ed.), Growth patterns in vascular plants. Dioscorides Press. Portland. orchid family. Dioscorides Press, Portland. Dueas, G.H.C. & J.L. Fernndez-Alonso. 2009. Sinopsis de la subfamilia Spiranthoideae (Orchidaceae). Rev. Acad. Colomb. Cienc. 33(127): 157-182. Eames, A.J. & L.H. MacDaniels, 1947. An introduction to plant anatomy. McGraw-Hill Book Company, New York. Ely, F. 1996. Anatoma foliar de Melastomateceae a lo largo de un gradiente altitudinal en el Parque Nacional Sierra Nevada de Mrida. Tesis de Grado de Licenciatura. Facultad de Ciencias, Universidad de los Andes. Mrida-Venezuela. Esau, K. 1976. Plant anatomy. Barcelona-Espaa. Escalona, A. 1999. Anatoma foliar y del xilema de arbustos con aspecto de Gimnosperma en el Parque Nacional Sierra Nevada. Mrida-Venezuela. Tesis de grado de Licenciatura. Facultad de Ciencias, Universidad de los Andes. Mrida-Venezuela. Estelita, M.E.T. & W. Handro. 1983. Leaf ultrastructure in species of Gomphrena and (Amaranthaceae). Can. J. Bot. 62: 812-817. Fahn, A. 1967. Plant anatomy. 4ta ed. (1979). Pergamom Press. Canad. Fahn, A. 1974. Plant anatomy. 2da ed. Ediciones H. Blume. Madrid-Espaa. Figueroa, C., G.A. Salazar, H.A. Zavaleta & E.M. Engleman. 2007. Root character evolution and systematics in Cranichidinae, Prescottiinae and Spiranthinae (Orchidaceae, Cranichideae). Ann. Bot. 1 -12. Flores, E.V. 1999. La planta. Estructura y funcin. Vol. II. Libro Universitario Regional. Costa Rica. Gaulhofer, K. 1908. Die perzeption der Lichtrichtung im Laubbltte mit Hilfe der Randtpfel, Randspalten und der windschiefen Radialwnde. Sber. Akad. Wiss. Wien (Math Nat. K.) 1(117): 153-190. Garay, L.A. 1978. Orchidaceae: Cypripedioideae, Orchidoideae, Neottiodeae. Pp. 3-304 in: Harling, G. & B. Sparre (Eds.). Flora of Ecuador 9. distribution of orchids. Proc. 4th Int Orchid Conference, Singapore: 170-187. Garca, M.C.LL. 2004. Crecimiento, morfoanatoma radical, acumulacin de solutos orgnicos y relaciones inicas en dos genotipos de caa de azcar (Sacharum spp) con tolerancia salina contrastante. Tesis de Doctorado. Facultad de Agronoma, Universidad Central de Venezuela. Haberlandt, G. 1990[1884]. Physiological plant anatomy today & tomorrows. XVII Printers & Ruble, New Delhi. Hose, E., D.T. Clarkson, E. Steudle, L. Shreiber, & W. Hartung, 2001. The exodermis: a variable apoplastic barrier. J. Experim. Bot. 52: 2245-2264. Jenik, J. 1978. Roots and root systems in tropical trees: morphological and ecological aspects. In: Tomlinson, P.B. & M.H. Zimmermann (Eds.), Tropical trees as living systems. Cambridge University Press, Cambridge. Johanssen, D.A. 1940. Botanical microtchnique. Mc Graw Hill Book. New York. Lasser, T. 1969. Flora de Venezuela. Vol. XV. I parte. Instituto Botnico. Direccin de Recursos Naturales Renovables. Ministerio de Agricultura y Cra. Caracas. Lindorf, H., L. Parisca, & P. Rodrguez. 1991. Botnica, Central de Venezuela. Ediciones de la Biblioteca. Caracas-Venezuela. Luque, R. A. 2004. Estructura primaria del sistema radical de Coespeletia Cuatrec. Interciencia 29(1): 13-18. Luque, R.A. 2000. El sistema radical de Coespeletia Cuatrec. del pramo de Piedras Blancas, Mrida. LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012.74 LANKESTERIANA

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Trabajo de Ascenso a la categora de Prof. Asociado. Universidad de los Andes. Luque, R.A., N.L. Menezes, & J. Semir. 1999. Anatoma foliar de Lychnophora Mart. (Vernonieae: Asteraceae). Plantula 2(3): 141-152. Luque R.A., H. Sousa, & J. Kraus. 1996. Mtodos de visando a substituo do azul de astra por azul de alcio 8 GS ou 8GX. Acta Bot. Brasil. (10): 199-212. Monasterio, M. & S. Reyes. 1980. Diversidad ambiental y variaciones en la vegetacin de los pramos de los Andes Venezolanos. Pp. 47-92 in: Monasterio, M. (ed.), Estdios ecolgicos de los pramos andinos. Mrida: Universidad de Los Andes Perumalla, C.J. & C.A. Peterson. 1986. Deposition of Casparian bends and suberin lemellae in the exodermis and endodermis of young corn and onion roots. Canad. J. Bot. 64(7-9):1873-1878. Petit, A.P.M. 1984. Variacin altitudinal de la vegetacin en los Andes Venezolanos. Tesis de grado. Facultad de Ciencias Forestales y Geografa, Universidad de los Andes. Mrida-Venezuela. Porembski S. & W. Barthlott. 1988. Velamen radicum Nordic J. Bot. 8: 117. Pridgeon A.M, P.J. Cribb, M.W. Chase, F.N. Rasmussen, eds. 2003. Genera Orchidacearum. Vol. 3. Orchidoideae. Part 2. Vanilloideae. Oxford University Press, Oxford. Pyykk, M. 1966. The leaf anatomy of East Patagonia xeromorphic plants. Ann. Bot. Fenn. 3: 453-622. Rada. F., G. Goldstein, A. Azcar, & F. Meinzer, 1985. Freezing avoidance in Andean giant rosette plants. Pl. Cell Environ. 8: 501-507. Rasmussen, H.N. & D.F. Whigham, 1998. The underground phase: a special challenge in studies of terrestrial orchid populations. Bot. J. Linn. Soc.126: 4964. Ricardi, M.H. 1988. Familias de monocotiledneas Venezolanas. 1ra Edicin. Consejo de Publicaciones de la Universidad de los Andes. Mrida-Venezuela. Rojas. M. 2002. Anatoma en rganos vegetativos y Orthrosanthus S. y Sisyrinchium L. (Iridaceae) que crecen en el Parque Nacional Sierra Nevada de Mrida. Tesis de Grado de Licenciatura. Facultad de Ciencias, Universidad de los Andes. Mrida-Venezuela. Roth, I. 1990. Leaf structure of a Venezuelan cloud forest in relation to the microclimate. Encyclopedia of Plan Anatomy. Gebrder Bontraeger. Berlin. Stuttgart. Roth, I. 1991. Anatoma de las plantas superiores. Universidad Central de Venezuela. Ediciones de la Biblioteca.Caracas. Saas, J.E. 1958. Botanical Microtechnique, 3ra edicin. Editorial Lona State Collage Press. New York. Salazar, G.A., L.I. Cabrera, S. Madrin & M.W. Chase. 2009. Phylogenetic relationships of Cranichidinae and Prescottiinae (Orchidaceae, Cranichideae) inferred from plastid and nuclear DNA sequences. Ann. Bot.104: 403. Stern, W.L., H.C. Aldrich, L.M. Mcdowell, M.W. Morris & A.M. Pridgeon. 1993a. Amyloplasts from cortical root cells of Spiranthoideae (Orchidaceae). Protoplasma 172: 49-55. Stern, W.L., M.W. Morris, W.S. Judd, A.M. Pridgeon, & R.L. Dressler. 1993b. Comparative vegetative anatomy and systematic of Spiranthoideae (Orchidaceae). Bot. J. Linn. Soc. 113: 161-197. Tomlinson, P. 1969. Anatomy of the Monocotyledons. Vol. III. Pp. 92-126 en: Metcalfe, C. (ed.), CommelinalesZingiberales. Clarendon press. Oxford. Torres, F. 1998. Anatoma foliar en Coespeletia Cuatrec del pramo de Piedras Blancas. Plan estructural bsico y de los Andes. Mrida-Venezuela. Torres, F.R., J.C. Gaviria & J. Peraza. 1996. Anatoma foliar de Oritrophium (H.B.K.) Cuatrec. del pramo de Sierra Nevada, Estacin Loma Redonda. Plantula 1(1): 65-73. Trujillo, D. & A.R. Delgado. 2011. Aa from lomas format ions. A New Orchidaceae record from the desert coast of Peru. Lankesteriana 11(1): 33-38. Van Fleet, D.S. 1961. Histochemistry and function of the endodermis. Bot. Rev. 27(2): 165-219. Wilkinson, H. 1979. The plant surface (mainly leaf): stoma. 2da ed. Vol. I. in: Metcalfe, C. & L. Chalk, (eds.), Anatomy of the dicotyledons. Clarendon Press. Oxford.LANKESTERIANA 12(1), April 2012. Universidad de Costa Rica, 2012. DUGARTE CORREDOR & LUQUE ARIAS Morfoanatoma en Cranichideae75