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Determinants of Phenotypic Variation

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General section DETERMINANTS OF PHENOTYPIC VARIATION Determinants of phenotypic variation in the Iberian wall lizard species IN THE IBERIAN WALL LIZARD SPECIES complex (Podarcishispanicus)….................................................................... 15 COMPLEX (PODARCIS HISPANICUS) Chapter I: Determinantes de la variación fenotípica en el complejo de especies de Altitudinally divergent adult phenotypes in Iberian wall lizards are not la lagartija ibérica (Podarcis hispanicus) driven by egg differences nor hatchling growth rates..................................... 89 Chapter II: Environmental–determined growth may explain altitudinal body size differences in Iberian wall lizards: a reciprocal transplant experiment......... 121 Chapter III: 2015 TESIS DOCTORAL Conspicuous blue tails, dorsal morphs and escape behaviour in hatchling Iberian wall lizards (Podarcis hispanicus)..................................................... 143 Chapter IV: Dorsal pattern polymorphism in female Iberian wall lizards: differences in morphology, dorsal coloration, immune response and reproductive investment .................................................................................................... 181 Chapter V: Giménez Altitudinal variation in morphology and coloration in the Brown lizard (Podarcis liolepis) ......................................................................................... 217 Chapter VI: s Ortega ú Interpopulational variations in sexual chemical signals of Iberian wall lizards may allow maximizing signal efficiency under different climatic Jes conditions....................................................................................................... 245 Jesús Ortega Giménez Tesis doctoral 2015 el complejo de estFACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Zoología y Antropología Física TESIS DOCTORAL Determinants of phenotypic variation in the Iberian wall lizard species complex (Podarcis hispanicus) Determinantes de la variación fenotípica en el complejo de especies de la lagartija ibérica (Podarcis hispanicus) Memoria presentada por el licenciado Jesús Ortega Giménez para optar al grado de Doctor en Ciencias Biológicas, dirigida por el Doctor José Martín Rueda y la Doctora Pilar López Martínez del Departamento de Ecología Evolutiva del Museo Nacional de Ciencias Naturales- Consejo Superior de Investigaciones Científicas. Madrid, 2015 Jesús Ortega Giménez © Jesús Ortega Giménez, 2015 La foto que abre la sección general fue realizada por Jesús Ortega Giménez. Queda prohibida la reproducción o publicación total o parcial, así como la producción de obras derivadas sin la autorización expresa de los autores. La presente Tesis Doctoral ha sido financiada por una beca de la Junta para la Ampliación de Estudios e Investigaciones Científicas (JAE), concedida por el Consejo Superior de Investigaciones Científicas (CSIC). Asimismo, los estudios realizados han sido financiados por el Ministerio de Ciencia e Innovación a través de los proyectos MICIIN-CGL2011- 24150/BOS. La Comunidad de Madrid ha otorgado los permisos necesarios para la captura de las lagartijas. La biología es la ciencia. La evolución es el concepto que hace a la biología singular. Jared Diamond FACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Zoología y Antropología Física TESIS DOCTORAL Determinants of phenotypic variation in the Iberian wall lizard species complex (Podarcis hispanicus) Determinantes de la variación fenotípica en el complejo de especies de la lagartija ibérica (Podarcis hispanicus) Memoria presentada por el licenciado Jesús Ortega Giménez para optar al grado de Doctor en Ciencias Biológicas, dirigida por el Doctor José Martín Rueda y la Doctora Pilar López Martínez del Departamento de Ecología Evolutiva del Museo Nacional de Ciencias Naturales- Consejo Superior de Investigaciones Científicas. Madrid, 2015 El doctorando Jesús Ortega Giménez VºBº del Director VºBº del Director VºBº del Tutor INDEX José Martín Rueda Pilar López Martínez José A. Díaz INDEX Agradecimientos/Acknowledgements................................................................... 9 General section Introduction ............................................................................................................. 17 Objectives ................................................................................................................ 35 Material and methods .............................................................................................. 37 Results and discussion ............................................................................................. 49 Conclusions .............................................................................................................. 61 References ................................................................................................................ 64 Chapter I Altitudinally divergent adult phenotypes in Iberian wall lizards are not driven by egg differences nor hatchling growth rates ......................................... 89 Chapter II Environmental–determined growth may explain altitudinal body size differences in Iberian wall lizards: a reciprocal transplant experiment…..…. 121 Chapter III Conspicuous blue tails, dorsal morphs and escape behaviour in hatchling Iberian wall lizards (Podarcis hispanicus)............................................................ 143 Chapter IV Dorsal pattern polymorphism in female Iberian wall lizards: differences in morphology, dorsal coloration, immune response and reproductive investment ............................................................................................................. 181 Chapter V Altitudinal variation in morphology and coloration in the Brown lizard (Podarcis liolepis) ................................................................................................... 217 Chapter VI Interpopulational variations in sexual chemical signals of iberian wall lizards may allow maximizing signal efficiency under different climatic conditions................................................................................................................ 245 Resumen extenso en español................................................................................. 281 Abstract………………………………………………………………………...... 315 Resumen………………………………………………………………………..... 319 Agradecimientos/Acknowledgements Ha sido largo camino y es momento de pararse por un momento a reflexionar. Recuerdo dos sabios consejos que me dieron en mi infancia: para saber algo tienes que leer, luego leer y luego leer más; si quieres ser bueno en algo hay que practicar, practicar y practicar. No puedo dejar de pensar que la constancia y tener claro un objetivo son condiciones indispensables para prosperar, no solo en una carrera científica, sino en cualquier otra disciplina o ámbito de la vida. No obstante, el mérito no es solo mío pues no habría podido llegar hasta aquí sin la ayuda y apoyo de unas cuantas personas. Mención especial se merece mi familia, mis padres Jesús y Maribel y mi hermana Alba. Muchas gracias por vuestro apoyo y por vuestro aguante ante situaciones cuanto menos pintorescas. Recuerdo como el congelador poco a poco paso a convertirse en una especie de morgue en la que tenían cabida animales de muy diversa índole, reptiles en su mayoría, que pasaban por allí a la espera de ser diseccionados. Muchas gracias por permitirme tener y criar en casa infinidad de bichos. Fuisteis especialmente pacientes con mi afición por la cría de gecos, lo que no fue precisamente un capricho pasajero. Durante ese tiempo aprendí muchísimas cosas, algunas de las cuales me han permitido llevar a cabo esta tesis de una manera más fácil. Sin embargo, hay dos lecciones con las que me quedo: la importancia de pensar a largo plazo y la utilidad de ser capaz de trabajar de una manera independiente. No me puedo olvidar de Elena, mi mujer. Gracias por comprender que el camino que he elegido, a pesar de ser muy gratificante, está lleno de dificultades y hay veces que es necesario decidir con la cabeza y no con el corazón. Tu apoyo y cariño han sido decisivos no ya solo para acabar la tesis sano y salvo, sino para tirar para adelante y, en definitiva, ser feliz. La verdad que quedaría muy bien agradeciendo su buen hacer a todos los profesores con los que me he topado, tanto en el colegio como en la facultad, pero lo mío no es ser 9 politicamente correcto. En el colegio conocí a grandes profesores como Donato, José Luis, Enrique e Hilario, se que no era difícil destacar pero os mereceis mi más sincero respeto y agradecimiento, profesionales como vosotros no abundan. En otra categoría se encuentran personajes como Manolo, profesor de biología, quien fue capaz de hacer odiar una asignatura tan apasionante como la biología a toda una promoción, salvo al que escribe estas líneas. Es curioso que todas y cada una de las mentes reprimidas (léase monjas) que me han dado clase hayan dejado mucho que desear, ¿casualidad? lo dudo mucho. No es normal que la misma persona imparta religión, geografía e historia, lengua castellana y francés. Demasiados dogmas y muy poca formación. Vergüenza ajena, a todas luces. No obstante, uno no puede negar el gran valor didáctico que tienen los malos ejemplos, de todo se aprende. Fue muy triste comprobar cómo este mismo patrón se repite en la facultad, abundan los mediocres. Por suerte conocí a Juan Mateo Pérez Zaballos, quien impartía Modelos Adaptativos en Zoología, asignatura extinta en el plan actual, una pena. Me ofreció la oportunidad
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