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Estudio Filogenético Del Grupo De" Rhizocarpon Geographicum

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Estudio Filogenético Del Grupo De UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE FARMACIA DEPARTAMENTO DE BIOLOGÍA VEGETAL II ESTUDIO FILOGENÉTICO DEL GRUPO DE "RHIZOCARPON GEOGRAPHICUM" (LÍQUENES, RHIZOCARPACEAE, ASCOMYCOTA) ANÁLISIS CONTRASTADO DE LOS CARACTERES MORFOLÓGICOS Y LOS PATRONES BIOGEOGRÁFICOS PHYLOGENETIC STUDY IN THE RHIZOCARPON GEOGRAPHICUM GROUP (LICHENS, RHIZOCARPACEAE, ASCOMYCOTA) CONSTRACTING ANALYSIS OF MORPHOLOGICAL CHARACTERS AND BIOGEOGRAPHIC PATTERNS TESIS DOCTORAL DE: BEATRIZ ROCA VALIENTE DIRIGIDA POR: ANA M. CRESPO DE LAS CASAS SERGIO PÉREZ ORTEGA LEOPOLDO GARCÍA SANCHO Madrid, 2013 ©Justo Aquilino Corral Aragón, 2013 Universidad Complutense de Madrid Facultad de Farmacia Departamento de Biología Vegetal II Estudio filogenético del grupo de Rhizocarpon geographicum (Líquenes, Rhizocarpaceae, Ascomycota). Análisis contrastado de los caracteres morfológicos y los patrones biogeográficos Memoria de Tesis Doctoral Beatriz Roca Valiente Tesis dirigida por: Dra. Ana M. Crespo Dr. Sergio Pérez Dr. Leopoldo García de las Casas Ortega Sancho F. de Farmacia (UCM) MNCN (CSIC) F. de Farmacia (UCM) En, Madrid, a 22 de Febrero de 2013 Dña. Ana M. Crespo de las Casas, Doctora en Ciencias Biológicas y Catedrática de Botánica de la Facultad de Farmacia de la Universidad Complutense de Madrid, Don Sergio Pérez Ortega, Doctor en Ciencias Biológicas e Investigador del Departamento de Biología Ambiental del Museo Nacional de Ciencias Naturales (CSIC) y Don Leopoldo García Sancho, Doctor en Ciencias Biológicas y Catedrático de Botánica de la Facultad de Farmacia de la Universidad Complutense de Madrid. Informan: La presente Tesis Doctoral titulada “Estudio filogenético del grupo de Rhizocarpon geographicum (Líquenes, Rhizocarpaceae, Ascomycota). Análisis contrastado de los caracteres morfológicos y los patrones biogeográficos”, ha sido realizada bajo su dirección por la Licenciada en Ciencias Biológicas Beatriz Roca Valiente, para optar al Grado de Doctor con mención de Doctorado Europeo, y al estar concluida, autorizan su presentación para que sea juzgada por el tribunal correspondiente. Y para que así conste firman el presente informe en Madrid 22 de Febrero de 2011. Vº Bº Director de Tesis Vº Bº Director de Tesis Dra. Ana M. Crespo de las Casas Dr. Sergio Pérez Ortega Vº Bº Director de Tesis Dr. Leopoldo García Sancho RECONOCIMIENTOS Este trabajo se ha realizado en el Departamento de Biología Vegetal de la Facultad de Farmacia de la Universidad Complutense de Madrid, gracias a la financiación proporcionada por una ayuda para la Formación de Personal Investigador BES-2007-16763 asociada al proyecto POL 2006-08405, concedido ambos por el desaparecido Ministerio de Ciencia e Innovación del estado español. El mismo programa FPI financió dos estancias breves en el extranjero: -National Museum of Nature and Science, ( 国 立 科 学 博 物 館, TNS), en Tsukuba, Japón. -Naturmuseum Senckenberg, en Frankfurt am Main, Alemania. A mi padre, Ángel Roca. Cuando te contestes con ser simplemente tú mismo, y no te compares ni compitas, todos te respetarán. Lao Tse. Tao Te Ching (ca. s. V a.C) AGRADECIMIENTOS No puedo dar por concluida la tesis sin agradecer a tantas personas que han estado a mi lado durante todos estos años y que me han ayudado de una u otra manera. Gracias: A mis directores de tesis, Ana, Leo y Sergio, primero por darme la oportunidad de iniciarme en la investigación científica, por la confianza que depositaron en mí, por su paciencia en mis buenos y malos momentos, por su disposición a enseñar y aprender, por sus consejos y por invertir mucho de su tiempo y esfuerzo en la supervisión y revisión de este trabajo. De mis dos estancias en el extranjero: al Dr. Yoshihito Ohmura por su colaboración y la acogida en Japón. Al Dr. Christian Printzen por sus comentarios e ideas sobre mi trabajo y estar siempre disponible para la discusión. Debo agradecer en ambos casos el apoyo de “las” técnicos de laboratorio, sin él no podría haber aprovechado tanto mis estancias, no nombro aquí a nadie por miedo a olvidarme de alguien. Especialmente debo recordar a Fernando Fernández-Mendoza porque hizo que mi estancia en Frankfurt fuera más enriquecedora de lo que hubiera esperado, profesional y personalmente. Muchas Gracias. 感 謝. Danksagung. A la Unidad de Genómica (Parque Científico de Madrid, UCM), especialmente a Rosana y a Tomas, y a Ricardo García Mata del Servicio Informático de Apoyo a la Docencia y a la Investigación (UCM). Al personal del Departamento de Biología Vegetal II, especialmente a Charo, Pilar y Manolo por estar siempre dispuestos a ayudarme, (aunque fuera en la hora de la fruta, tengo un ojo…). Al Dr. Pepe Pizarro, conservador del herbario MAF, por su disposición y apoyo indispensables para la tesis. A la Dra. Ana Pintado porque fue la primera que confió en mí, por su sinceridad y su animó para que solicitará la beca, muchas gracias. A la Prof. Paloma Cubas por compartir tantos conocimientos y discutir mis resultados. Al Prof. Víctor Rico por sus recomendaciones, su disposición y su ayuda. Al Dr. Divakar y a la Dra. Del Prado por enseñarme en el laboratorio y prestarme ayuda siempre que la pedí. Al Dr. Ruibal por todo que es mucho. Al Prof. José Antonio Molina por querer trabajar conmigo, por darme la oportunidad de participar en sus proyectos y ser un poquito más multidisciplinar. A mis compañeros becarios, “pre y post”, de la sala blanca, del herbario, del laboratorio de fisiología arriba y abajo, (estamos un poco desperdigados¡¡). Muchas gracias por las ideas, las colaboraciones, las comidas, los cafés, las cañas, los cotilleos, las salidas de campo, los viajes dentro y fuera (también gracias por los monetes, los elefantes, y los cerdos despistados en las dehesas)… vamos por estar ahí dispuestos a compartir. Probablemente sois los que mejor habéis comprendido todo “esto”. Con especial cariño a Celia (sin ti los “tomaca” ya no son lo mismo), a Ana Lumbr. (mi sitio siempre será tu sitio) y a Mer. Si algo me llevó son unos cuantos buenos amigos, ellos ya lo saben (^_^). A mi familia, numerosa, pero en especial a mi madre, Carmen, y a mis hermanas porque han sido, son y serán un apoyo fundamental. A mis sobrinxs, que sólo con pensar en ellos me sale una sonrisa y se me olvidan los males. A mi padre que se “fue” cuando todo esto no hacía nada más que comenzar, ojala estuvieras aquí… A él (mi padre) y a mi hermana Susana, que me enseñaron a luchar, a decir lo que pienso, a demostrar lo que valgo y a no agachar nunca la cabeza. A mis amigxs por escucharme y comprenderme en todos los buenos y los malos momentos de los últimos 7 años, que no han sido pocos. A las “biolocas” especialmente por los últimos años duros que hemos pasado. A David, mi compañero, mi amigo, mi sol… por quererme y cuidarme y dármelo todo sin pedirme nada. Porque la adversidad nos une más y cada día lo demostramos. A todos los que se acordaron de mí en sus salidas de campo y me trajeron material. A los que se han ofrecido a leerla, a los que no sabíais o no comprendíais lo que hacía, para qué servía, ni qué es un liquen, a los que me preguntaban: ¿para qué sirve doctorarse?, ¿y cuándo acabas?, ¿ya tienes fecha?, ¿otra vez te vas?, Espero que vuestras dudas se resuelvan, Porque ya está!!! A todxs Gracias!!! ÍNDICE / INDEX ABREVIATURAS ……………………………………………………………………………..… i ABREVIATURAS PARA LAS LOCALIDADES ESTUDIADAS …………………………………..... ii VOCABULARIO UTILIZADO EN LATÍN Y EN INGLES ………………………………………..… iii ABSTRACT (Resumen amplio en Ingles) …………………………………………………… 1 INTRODUCTION ………………………………………………….…………………………… 1 AIMS AND STRUCTURE ……………………………………….……………………………… 2 METHODOLOGY AND RESULTS ……………………………………………………………… 3 CONCLUSIONS ………………………………….…………………….……………………… 4 INTRODUCCIÓN GENERAL / GENERAL INTRODUCTION ……………………..………. 5 A.- El género Rhizocarpon Ramond ex DC ..………………………………..………. 7 B.- Las especies amarillas del género Rhizocarpon ………….……...................... 12 C.- La delimitación de especie; su interpretación en hongos liquenizados y dentro del grupo de Rhizocarpon geographicum …….………….......................................... 18 D.- Caracteres utilizados en la delimitación específica dentro del grupo de Rhizocarpon geographicum ………………………………………………………... 21 E.- Ecología y distribución ….……………..…..................................................... 31 F.- Biogeografía ….……………......................................................................... 32 G.- Análisis de los caracteres moleculares del micobionte ….……………............. 34 H.- Análisis de los caracteres moleculares del fotobionte ….………….................. 36 J.- Relevancia del estudio del grupo de Rhizocarpon geographicum ……………... 37 OBJETIVOS / AIMS ……………..……………………………………………….……...…. 41 METODOLOGÍA GENERAL / GENERAL METHODS ……….……………………………. 47 A.- Material biológico ……….…...……………………..………………..…………. 49 B.- Método de extracción de ADN ……….…...………………………....……...…. 50 C.- Loci utilizados, amplificación de las secuencias de ADN y condiciones de PCR …………………………………………..………………….. 51 D.- Purificación y secuenciación de los productos de PCR ………........................ 54 E.- Alineamiento de secuencias, modelos evolutivos de sustitución de nucleótidos y análisis filogenéticos ……………………………………………......................... 56 CAPÍTULO 1 / CHAPTER 1 ……………………………………..………...................... 59 Descripción detallada de las especies y revisión de las especies tipo del grupo de Rhizocarpon geographicum / Species description and type species revision in the Rhizocarpon geographicum group CAPÍTULO 2 / CHAPTER 2 ……………………………………………...…………....... 91 Variabilidad Genética
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