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Universidad De Costa Rica Sistema De Estudios De UNIVERSIDAD DE COSTA RICA SISTEMA DE ESTUDIOS DE POSGRADO REVISIÓN SISTEMÁTICA DEL GÉNERO MYOXANTHUS (ORCHIDACEAE: PLEUROTHALLIDINAE) EN COSTA RICA Tesis sometida a la consideración de la Comisión del Programa de Estudios de Posgrado en Biología para optar al grado y título de Maestría Académica en Biología GUSTAVO ROJAS ALVARADO Ciudad Universitaria Rodrigo Facio, Costa Rica 2020 DEDICATORIA Esta obra está dedica a mis padres Luz y Fernando, quienes siempre me han brindado su apoyo incondicional y a quienes debo mi especial interés y aprecio por la naturaleza, y sobre todo por las orquídeas. También a la familia Orozco Murillo y abuela Celi, a quienes debo su invaluable y especial apoyo durante mis estudios universitarios. ii AGRADECIMIENTOS Agradezco especialmente a mi tutor Adam Karremans, por su gran apoyo y enseñanzas desde el inicio de esta investigación. A Federico Albertazzi su apoyo inicial como tutor temporal, y por su especial ayuda en el trabajo de laboratorio y durante todo el proceso de Maestría. A Mario Blanco por su gran conocimiento transmitido y también por esas buenas retroalimentaciones sobre morfología vegetal. A Franco Pupulin y Diego Bogarín por sus valiosas enseñanzas en todo lo relacionado al mundo de las orquídeas; al último también por sus consejos en la elaboración de ilustraciones botánicas digitales y su ayuda brindada en los análisis filogenéticos. A todo el personal docente de la Escuela de Biología, de los cuales aprendí enormemente en sus clases. También agradezco a los profesores Gustavo Vargas, Luis Jorge Poveda y Rafael Murillo de la Universidad Nacional quienes me transmitieron gran parte de su conocimiento y apoyo durante el proceso de ingreso a la Maestría. Agradezco a Barbara Gravendeel, Richa Kusumawati, Dewi Pramanik y Bertie Joan van Hueven de la Universidad de Leiden, Países Bajos, por permitir el uso, guiar y asistir en la obtención de las imágenes en el microscopio electrónico de barrido. A Hannia Ramírez del Sistema de Estudios de Posgrado en Biología; en el Jardín Botánico Lankester (JBL) a Karen Barquero, Lizbeth Oses y Maricruz Bonilla del Departamento Administrativo; a Jorge Brenes, Giovanni Meza y Gerson Villalobos del Departamento de Horticultura; a Melissa Díaz y Grettel Salguero del Departamento de Investigación; y a mis compañeros José Esteban Jiménez, Marco Cedeño e Isler Chinchilla por su apoyo durante el proceso de la Maestría, actividades de campo, y aporte de material fotográfico de múltiples herbarios de diferentes países de América. A Jaime Aguilar† del JBL por sus inducciones en la documentación de plantas. Agradezco a los directores y personal de los herbarios USJ, CR, AMES, SEL, AMO, MO, UVAL, KEW, y W por brindar acceso a sus colecciones, aportar y permitir la reproducción de imágenes de especímenes utilizados en esta investigación. Agradezco también a todos mis amigos y demás personas que de una u otra forma me brindaron su apoyo constante durante esta etapa universitaria. Agradezco especialmente a Vero, una de las personas más importantes para mí y que siempre estuvo en los buenos y malos momentos y nunca dejó darme ánimo para perseguir cada una de mis metas. iii ÍNDICE GENERAL DEDICATORIA .............................................................................................................................. ii AGRADECIMIENTOS ................................................................................................................. iii HOJA DE APROBACIÓN ............................................................................................................. iv RESUMEN ................................................................................................................................... viii ÍNDICE DE FIGURAS ................................................................................................................... ix ÍNDICE DE TABLAS .................................................................................................................. xvi PREFACIO ...................................................................................................................................... 1 LITERATURA CITADA ................................................................................................................. 5 ARTÍCULO 1: Revision of the Costa Rican species of Myoxanthus (Pleurothallidinae: Orchidaceae) ................................................................................................................................... 10 Abstract:...................................................................................................................................... 11 Resumen: .................................................................................................................................... 11 Keywords: ................................................................................................................................... 11 Introduction .................................................................................................................................... 11 The genus Myoxanthus in Costa Rica: ....................................................................................... 13 Distribution: ................................................................................................................................ 14 Materials and Methods ................................................................................................................... 15 Taxonomy ...................................................................................................................................... 16 Key to the Costa Rican species of Myoxanthus .......................................................................... 17 1. Myoxanthus affinis .................................................................................................................. 18 2. Myoxanthus cereus ................................................................................................................. 22 3. Myoxanthus colothrix ............................................................................................................. 24 4. Myoxanthus congestus ............................................................................................................ 27 5. Myoxanthus exasperatus......................................................................................................... 31 6. Myoxanthus hirsuticaulis ........................................................................................................ 35 7. Myoxanthus parahybunensis .................................................................................................. 38 8. Myoxanthus scandens ............................................................................................................. 42 v 9. Myoxanthus sotoanus ............................................................................................................. 45 10. Myoxanthus trachychlamys .................................................................................................. 48 Literature cited ............................................................................................................................... 52 ARTÍCULO 2: Infrageneric evaluation of Myoxanthus (Pleurothallidinae: Orchidaceae) ......... 111 Abstract:.................................................................................................................................... 112 Resumen: .................................................................................................................................. 112 Keywords: ................................................................................................................................. 112 Introduction .................................................................................................................................. 112 Materials and methods ................................................................................................................. 114 DNA extraction: ....................................................................................................................... 114 ADN amplification and sequencing: ........................................................................................ 114 Editing of sequences and analysis: ........................................................................................... 114 Scanning electron microscopy (SEM): ..................................................................................... 115 Results .......................................................................................................................................... 115 Discussion .................................................................................................................................... 117 Characterization of discrete groups .......................................................................................... 118 Clade A ................................................................................................................................. 118 Myoxanthus lonchophyllus group: ................................................................................... 118 Clade B ................................................................................................................................. 119 Myoxanthus congestus group: .........................................................................................
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