Centro De Investigación Científica De Yucatán, A.C. Posgrado En Ciencias Biológicas

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Centro De Investigación Científica De Yucatán, A.C. Posgrado En Ciencias Biológicas ~ .:S POSGRADO EN ~ ( CIENCIAS CICY ( ) BIOLÓGICAS Centro de Investigación Científica de Yucatán, A.C. Posgrado en Ciencias Biológicas EVOLUCIÓN DE CARACTERES DE HISTORIA DE VIDA Y TASAS DE DIVERSIFICACIÓN DE LA FAMILIA BROMELIACEAE Tesis que presenta GUADALUPE JEANETT ESCOBEDO SARTI En opción al título de DOCTOR EN CIENCIAS (Ciencias Biológicas: Opción Recursos Naturales) Mérida, Yucatán, México Septiembre 2013 CENTRO DE INVESTIGACIÓN CIENTÍFICA DE YUCATÁN, A.C. POSGRADO EN CIENCIAS BIOLÓGICAS :-...:: :S (_ .>O')Gf<A )0 ~ ~ ) CIENCIAS CICY RECONOCIMIENTO ( B )LOC1I A Por medio de la presente, hago constar que el trabajo de tesis titulado "Evolución de ca­ racteres de historia de vida y tasas de diversificación de la familia Bromeliaceae" fue realizado en los laboratorios de la Unidad de Recursos Naturales del Centro de Inves­ tigación Científica de Yucatán , A.C. bajo la dirección de la Dra. lvón Mercedes Ramírez Morillo, dentro de la opción Doctorado después de Maestría, perteneciente al Programa de Posgrado en Ciencias Biológicas de este Centro. Atentamente, ........_..""""' Vázquez Flota Coordinador de Docencia Centro de Investigación Científica de Yucatán, A.C. Mérida, Yucatán , México; a 8 de septiembre de 2013 DECLARACIÓN DE PROPIEDAD Declaro que la información contenida en la sección de Materiales y Métodos Experimentales, los Resultados y Discusión de este documento proviene de las actividades de experimen­ tación realizadas durante el período que se me asignó para desarrollar mi trabajo de tesis, en las Unidades y Laboratorios del Centro de Investigación Científica de Yucatán, A.C., y que a razón de lo anterior y en contraprestación de los servicios educativos o de apoyo que me fueron brindados, dicha información, en términos de la Ley Federal del Derecho de Autor y la Ley de la Propiedad Industrial, le pertenece patrimonialmente a dicho Centro de Investigación. Por otra parte, en virtud de lo ya manifestado, reconozco que de igual manera los productos intelectuales o desarrollos tecnológicos que deriven o pudieran derivar de lo correspondiente a dicha información, le pertenecen patrimonialmente al Centro de Investi­ gación Científica de Yucatán , A.C. , y en el mismo tenor, reconozco que si derivaren de este trabajo productos intelectuales o desarrollos tecnológicos, en lo especial, estos se regirán en todo caso por lo dispuesto por la Ley Federal del Derecho de Autor y la Ley de la Propiedad Industrial , en el tenor de lo expuesto en la presente Declaración. Firma: ~seo Yar=/:1 Nombre: Guadalupe Jeanett Escobedo Sarti AGRADECIMIENTOS Gracias a Dios por la vida, por la oportunidad que me dio de seguir estudiando. Gracias a CONACyT por otorgarme la beca no. 200523, para realizar mis estudios de Doc­ torado y gracias al Posgrado en Ciencias Biológicas, opción Recursos Naturales del CICY. Gracias a mi familia, por todo su amor, su apoyo, muchas gracias por financiar parte de mi estancia en Mérida, gracias por financiar los boletos de avión para realizar mi trabajo de campo en Oaxaca durante el primer año del posgrado. Gracias a Barbara Bodenhorn y a Laura Barraza, por invitarme a participar en los inter­ cambios culturales y científicos del 2006-201 O, y por financiar en el 2008, mi estancia y transporte a Barrow, Alaska a través de la National Science Foundation de los Estados Uni­ dos, por medio del proyecto Award 081635 (Aiaska/Mexico cultural-scientific interchange) 2008-2012. Gracias al CICY, por la beca de movilidad académica CICY 2011-11 , que me otorgó para realizar una estancia en el Instituto de Biología de la Universidad Nacional Autónoma de México, gracias a la Dra. Susana Magallón por abrirme las puertas de su laboratorio. Gracias a CONACyT por la beca mixta y al CICY por la beca de movilidad académica CICY 2012-11 que me otorgaron, este financiamiento me permitió real izar mi estancia doctoral en Harvard University Herbaria y en el Jardín Botánico de Missouri. Gracias Dr. Gustavo Romero por fungir como asesor externo durante la estancia, por las facilidades que me brindó para realizar parte de mi investigación en HUH. Gracias a las investigadoras que conformaron el primer comité tutorial: Oras. lvón Ramírez Morillo, Demetria Mondragón Chaparro, Luz María Calvo-lrabién y Azucena Canto Aguilar. En particular, gracias Dra. Demetria Mondragón Chaparro por la asesoría que me brindó para desarrollar el primer proyecto de tesis doctoral : "Dinámica poblacional de Tillandsia carlos-hankii, y su biología reproductiva", así como por el tiempo que dedicó a discutir ar­ tículos referentes al tema (vía Skype); muchas gracias por financiar las sal idas a campo en Oaxaca, así como por su apoyo técnico y logístico. Gracias a las Oras. Luz Maria Calvo­ lrabién y Azucena Canto, por las recomendaciones que hicieron durante los primeros dos exámenes tutoriales. Gracias a los investigadores que integraron el segundo com ité tutorial: Dra. lvón Ramírez Morillo, Dra. Demetria Mondragón Chaparro, Dr. Rodrigo Duno y Dr. Germán Carnevali. En particular, al Dr. Germán Carnevali le agradezco que me haya dicho que existía el método de superárbol, y que era una alternativa para iniciar un nuevo proyecto de tesis doctoral. Al Dr. Rodrigo Duno gracias por las ideas que aportó durante cada examen tutorial, por su entusiasmo y todas las sugerencias. Muchas gracias Dra. Susana Magallón Puebla, por la asesoría teórica y metodológica que me brindó, por toda la literatura que me facilitó, por platicarme de las tasas de diversificación y ayudarme a hacer el análisis. Le agradezco mucho las "pistas" que me dio para mejorar este proyecto de tesis, en verdad gracias por todo el apoyo, el tiempo, y por su paciencia. Muchas gracias Carlos Luis Leopardi Verde por enseñarme la teoría referente a los modelos de evolución de caracteres que utilicé en el capítulo IV de la tesis. Te agradezco mucho el tiempo que dedicaste a discutir diferentes artículos (de métodos filogenéticos, métodos de súper-árbol, modelos de evolución, tasas de diversificación, entre otros); muchas gracias por enseñarme a usar los diferentes programas que utilice para el desarrollo de este estudio, gracias por prestarme tu laptop cuando mi máquina no era suficiente para hacer algunos análisis o cuando el programa no funcionaba en mi laptop. Gracias por enseñarme a trabajar con ejemplares de herbario, gracias por el material fotográfico, muchísimas gracias por todo. A Eric Gouda (Utrecht University Botanic Gardens) por el material fotográfico de Bromelia­ ceae que me facilitó. A la Dra. Demetria Mondragón Chaparro, Dra. lvón Ramírez Morillo, Dra. Miriam Ferrer Ortega, Dra. Susana Magallón Puebla y al Dr. Rodrigo Duno, les agradezco el tiempo que dedicaron a revisar el documento final de la tesis, gracias por las sugerencias que hicieron para mejorar el escrito. Gracias a los técnicos de la línea de Sistemática y Florística, Lilia Can ltzá, Ma. Dolores Chan Puc, Amilcar Castillo, Silvia Hernández Aguilar y José Luis Muñoz Tapia, por su ayuda para conseguir algunas publicaciones. Gracias Dr. Juan Manuel Dupuy, por su apoyo como coordinador del posgrado en ciencias biológicas, opción recursos naturales del CICY. Gracias al personal de la biblioteca, a Narce Gamboa, Miriam Juanqui, Sergio Pérez por su am istad y su apoyo con la literatura. Gracias a las familias: Gamboa Angulo, Valencia Rito, Vargas Alonso y Soloveichik Puente por su am istad , hospitalidad y apoyo. Gracias a los compañeros y amigos del posgrado, por su amistad y hospitalidad. DEDICATORIA "Bendice alma mía al Señor, y no olvides ninguno de sus beneficios, Él es quien te corona de favores y misericordias" Salmos 103:1 Dedico esta tesis a los motores de mi vida: Al Dios Todopoderoso del que habla la Biblia (ese es el Dios en quien yo creo). Gracias porque fuiste mi refugio, mi consuelo, mi amigo, mi maestro, quien abrió mi entendimiento cuando lo único que leía en los artículos y libros eran palabras que no entendía. Para mí, esta tesis es un milagro, pues después de un año de estar en el posgrado, me di cuenta que debía empezar de cero, y que no tenía ni la más remota idea para desarrollar una nueva tesis doctoral, que no había financiamiento, que no tenía el conocimiento, que aun el lenguaje académico en esta área me era ajeno ... Pero Dios, tu estabas ahí, cuando te pedí con todo mi corazón y fe tu ayuda, tu respuesta llegó, casi casi como un plan de auxilio en caso de desastre (similar al DN-111 E de la Secretaria de la Defensa Nacional); fuiste acomodando las piezas del rompecabezas, libros, artículos, amigos, asesoría, etc. Gracias por este milagro, gracias por la oportunidad me diste de seguir estudiando. A mi familia, por su amor, apoyo, consejos, comprensión, ayuda en los momentos difíciles y por proveer los recursos necesarios para estudiar. Gracias por impulsarme y gracias por no dejarme tirar la toalla, gracias por la confianza que tuvieron en mí, sin su apoyo hubiera sido imposible. Los amo. Índice ÍNDICE GENERAL PÁGINA LISTA DE FIGURAS . iii LISTA DE CUADROS . V LISTA DE ABREVIATURAS ................................. vii RESUMEN . 1 ABSTRACT . 3 CAPÍTULO J. INTRODUCCIÓN GENERAL . 5 Objetivos . 6 Objetivo general . 6 Objetivos específicos . 7 CAPÍTULO JI. INTEGRATING PHYLOGENETIC INFORMATION FOR THE FAMILY BROMELIACEAE (POALES, MONOCOTYLEDONAE) . 11 Abstract . 11 lntroduction . 11 Materials and Methods . 14 Results .... .... ... ...... .. ... .. ... .. .. 15 Discussion . 15 Conclusions . 31 Acknowledgements . 32 CAPÍTULO 111. A PHYLOGENY OF BROMELIACEAE (POALES, MONOCOTYLEDO­ NAE) DERIVED FROM AN EVALUATION OF NINE SUPERTREE MET- HODS ..................................... 41 Abstract . 41 lntroduction . 41 Materials and Methods . 44 Results .. ..... .. ........ ... ..... .. .. ...... .. 50 Discussion . 55 Conclusions . 61 Acknowledgements . 62 Índice CAPÍTULO IV. EVOLUCIÓN DE CARACTERES DE HISTORIA DE VIDA Y TASAS DE DIVERSIFICACIÓN DE LA FAMILIA BROMELIACEAE . 71 Resumen . 71 Introducción . 71 Materiales y métodos 76 Resultados . 79 Discusión . 86 Conclusiones . 99 CAPÍTULO V. DISCUSIÓN Y CONCLUSIÓN GENERAL ................. 107 ANEXO 1. Matriz de estados de carácter utilizada para la reconstrucción de la evolución de los caracteres presentados en el capítulo IV ....
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