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Isabel Sanmartín Bastida, María Luisa Alarcón

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Isabel Sanmartín Bastida, María Luisa Alarcón ISABEL SANMARTÍN BASTIDA, MARÍA LUISA ALARCÓN CAVERO Y JUAN JOSÉ ALDASORO CERTIFICAN Que el trabajo titulado “Macroevolución y microevolución en sistemas insulares: el patrón Rand Flora en el género Canarina”, que presenta Mario Mairal Pisa, ha sido realizado bajo su dirección y autorizan su presentación al tribunal calificador para optar al grado de Doctor en el Programa de Doctorado de Conservación de Recursos Naturales por la Universidad Rey Juan Carlos. Y para que así conste, firman la presente en Madrid, a ___ de ______ de 2015. Directora de tesis Isabel Sanmartín Bastida Codirectora Codirector María Luisa Alarcón Cavero Juan José Aldasoro Martín Macroevolution and microevolution in island systems: the Rand Flora pattern in the genus Canarina Macroevolución y microevolución en sistemas insulares: el patrón Rand Flora en el género Canarina Tesis Doctoral Mario Mairal Pisa Julio 2015 Directora Isabel Sanmartín Bastida Codirectores María Luisa Alarcón Cavero Juan José Aldasoro Martín Departamento de Biología y Geología, Física y Química Inorgánica y Analítica Este trabajo ha sido realizado en el Real Jardín Botánico de Madrid, CSIC, bajo la dirección de la doctora Isabel Sanmartín Bastida, científica titular del Departamento de Conservación y Biodiversidad, y la codirección de los doctores María Luisa Alarcón Cavero y Juan José Aldasoro Martín, ambos del Departamento de Biodiversidad del Instituto Botánico de Barcelona, CSIC-ICUB. Además, parte de los resultados fueron obtenidos en las estancias realizadas en el Jardín Botánico ‘Viera y Clavijo’- Unidad Asociada CSIC (Cabildo de Gran Canaria; Gran Canaria, España) bajo la dirección del doctor Juli Caujapé Castells, en la Université de Lausanne (Lausanne, Suiza) bajo la dirección del doctor Nadir Álvarez y en la Université de Fribourg bajo la dirección del doctor Loïc Pellissier y la doctora Myriam Heuertz. La realización de la tesis ha sido posible gracias a una ayuda predoctoral de Formación del Personal Investigador (FPI, BES-2010-037261) concedida a Mario Mairal Pisa por el Ministerio de Ciencia e Innovación (ahora Ministerio de Economía y Competitividad) y a los proyectos i) Reconstrucción del Origen de la Rand Flora Afro-Mediterránea con datación filogenética e inferencia biogeográfica (proyecto CGL2009-13322-C03-01) y ii) The assembly of African floras from lineages to populations: effects of historical climate changes and species ecology (AFFLORA) (proyecto CGL2012-40129-C02-01) concedidos a Isabel Sanmartín por la misma entidad. Esta tesis doctoral opta a la mención de Doctorado Internacional. Para poder optar a esta mención, el documento ha sido evaluado por los doctores Thomas Couvreur del Institut de Recherche pour le Developpement y Luis Valente de la Universität Potsdam. ÍNDICE / INDEX AGRADECIMIENTOS / ACKNOWLEDGEMENTS..................................................................... 1 2 RESUMEN .......................................................................................................................... 1 7 I. Summary (in English) ......................................................................................................... 17 II. Síntesis (en español) ......................................................................................................... 23 III. Antecedentes ..................................................................................................................... 29 1. Biogeografía: dispersión vs. vicarianza ........................................................................................................... 29 1.1. Patrones de diversidad: las disyunciones biogeográficas ............................................................................................ 31 1.2 Origen evolutivo de una enigmática distribución geográfica: La “Rand Flora” Africana ............................................ 35 2. La filogeografía para detectar la diversidad que “no se ve” ........................................................................... 48 2.1. Buscando refugios de biodiversidad ........................................................................................................................... 50 3. Biología de islas .............................................................................................................................................. 53 3.1. Islas oceánicas e islas dentro de un continente: las Islas Canarias y las sky islands Afromontanas............................. 59 3.2. El archipiélago de islas Afromontanas ........................................................................................................................ 63 3.3. Las Islas Canarias ....................................................................................................................................................... 76 4. El género Canarina ......................................................................................................................................... 90 IV. Objetivos e hipótesis ..................................................................................................... 96 V. Metodología ................................................................................................................. 101 1. Las reconstrucciones filogenéticas ............................................................................................................... 101 1.1 Métodos de construcción de árboles. ..........................................................................................................................102 2. El uso de marcadores moleculares .............................................................................................................. 105 2.1 Secuencias de ADN ..................................................................................................................................................106 2.2. Polimorfismos de ADN..............................................................................................................................................107 3. El reloj molecular y el registro fósil .............................................................................................................. 110 4. En busca de la inferencia filogeográfica… ..................................................................................................... 114 4.2 Estimadores de la diversidad genética.........................................................................................................................116 4.3 Aproximaciones de vanguardia en filogeografía .........................................................................................................116 5. Los modelos de distribución de especies ..................................................................................................... 122 VI. Conclusions (in English) ........................................................................................... 124 VII. Conclusiones (en español) ...................................................................................... 128 VIII. Bibliography ............................................................................................................... 133 IX. List of papers included in this phD………………………………………………………………………....166 CAPÍTULOS / CHAPTERS ................................................................................................... 171 CHAPTER 1 Ancient vicariance and climate-driven extinction explain continental-wide disjunctions in Africa: the case of the Rand Flora genus Canarina (Campanulaceae). ................................. 173 CHAPTER 2 Geographic barriers and biota evolve together: continental rifting and climate change may explain patterns of genetic variation in the Afromontane epiphyte Canarina eminii. ......... 197 CHAPTER 3 Paleo-islands as refugia and sources of genetic diversity within volcanic archipelagos: The case of the widespread endemic Canarina canariensis (Campanulaceae) ........................... 235 CHAPTER 4 A tale of two forests: human-induced population decline threatens levels of genetic biodiversity in Afro-Macaronesian evergreen forest archipelago endemics (Canarina, Campanulaceae)……………………………………………………………………………………………………………….259 CHAPTER 5 Living on the edge: timing of Rand Flora disjunctions congruent with ongoing aridification in Africa ..................................................................................................................................... 297 CHAPTER 6 Lineage-specific climatic niche drove the temporality of vicariance. ................................... 315 ANEXOS / ANNEXES ......................................................................................................... 3 41 SUPPLEMENTARY DATA CHAPTER 1...................................................................................... 343 SUPPLEMENTARY DATA CHAPTER 2 ..................................................................................... 365 SUPPLEMENTARY DATA CHAPTER 3 ..................................................................................... 373 SUPPLEMENTARY DATA CHAPTER 5 ..................................................................................... 391 AGRADECIMIENTOS / ACKNOWLEDGEMENTS La participación de tantas personas en esta tesis tiene un valor especial, muchísima gente me ha acompañado y ayudado en su elaboración y con vosotros quiero compartir mi ilusión y alegría en estas páginas. ¡Aunque durante el camino ha habido obstáculos, gracias a vosotros no ha habido límites!
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