Amphibia: Anura, Caudata) a Través De Métodos Filogeográficos

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Amphibia: Anura, Caudata) a Través De Métodos Filogeográficos UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Zoología y Antropología Física TESIS DOCTORAL Estudio de patrones de diversidad intraespecífica en anfibios (Amphibia: anura, caudata) a través de métodos filogeográficos MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR Ernesto Recuero Gil DIRECTORES: Gabriela, dir Parra Olea Mario, dir García París Madrid, 2015 © Ernesto Recuero Gil, 2011 Universidad Complutense de Madrid - Facultad de Ciencias Biologicas Estudio de patrones de dîversidad intraespecifica en anfibios (Amphibia: Anura, Caudata) a través de métodos fîlogeogrâfîcos. Ernesto Recuero Gil 2010/2011 UNIVERSIDAD COMPLUTENSE 532998816X CSIC . o T C mnc® Dpto. Zoologi'a y Antropologi'a Fi'sica Dpto. Biodiversidad y Biologia Evolutiva Facultad de Ciencias Biolôgicas Museo Nacional de Ciencias Naturales Universidad Complutense de Madrid Consejo Superior de Investigaciones Cientificas Estudio de patrones de dîversidad intraespecifica en anfîbios (Amphibia: Annra, Candata) a través de métodos filogeogràfîcos. Memoria presentada por ERNESTO RECUERO GIL para optar al grado de Doctor en Biologia directores de tesis Dra. Gabriela Parra Olea Dr. Mario Garcia Paris CO/Wp Madrid, 2010/2011 A mi familia. Por to do. Indice Agradecimientos ...........................................................................................7 I. Introduccion general ..................................................................11 IL Capitules ♦♦♦ Capitule 1 Phylogeography of Pseudacris regilla (Anura: Hylidae) in western North America, with a proposai for a new taxonomie rearrangement Molecular Phylogenetics and Evolution, 39: 293-304 .............. 47 Capitule 2. Mitochondrial differentiation and biogeography of Hyla meridionalis (Anura: Hylidae): an unusual phylogeographical pattern Journal of Bio geography, 34: 1207-1219 ................................. 65 <♦ Capitule 3. Evolutionary history of Lissotriton helveticus: multilocus assessment of ancestral vs. recent colonization of the Iberian Peninsula En preparacion ..............................................................................83 ❖ Capitule 4 Urban aquatic habitats and conservation of highly endangered species: the case of Ambystoma mexicanum (Caudata, Ambystomatidae) Annales Zoologici Eennici, 47: 223-238 ................................ 159 < Capitule 5. Polymorphic microsatellite markers for Mexican salamanders of the genus Amby stoma Molecular Ecology N otes,! \ 818-820 ................................... 179 <♦ Capitule 6 Insight on the conservation genetics of threatened populations of Mexican Ambystoma (Caudata: Ambystomatidae) En preparacion ........................................................................ 187 III. Discusion general ................................................................. 255 IV. Cenclusienes.......................................................................... 285 Agradecimientos Una de las conclusiones de este trabajo bien puede ser que realizar una tesis doctoral es un excelente medio para conocer gente. Efectivamente a lo largo de estos anos son muchas las personas que han intervenido de forma directa o indirecta en todo el proceso, y otras tantas que, podriamos decir, se han visto salpicadas por el mismo. Aprovecho este rinconcito para expresar mi mas profundo agradecimiento a todas ellas, sea cual sea su papel en los avatares de esta tesis. Un recuerdo muy especial para Marina, a la que tanto echamos de menos y cuya ausencia se hace notar cada dia. Muchas gracias por todo Marina, ojalâ hubieras estado con nosotros para disfrutar de estos momentos Un millôn de gracias para mis directores, Gabriela y Mario, sin los cuales nada de esto h abri a sido posible. Muchas gracias por vuestro apoyo, confianza y por supuesto por todos los buenos ratos que hemos compartido. Muchas gracias también a todos los que han contribuido a la realizaciôn de cada uno de los trabajos incluidos aqui, pero muy especialmente a mis coautores: Ihigo, Gabriela, Mario, Ainhoa, Xabi, Annie, Juan, Kelly. Espero que seguirâ habiendo ocasiones para trabajar juntos. A mi querido grupo herpetoentomolôgico, por el que tantos van y vienen, convirtiendo esta experiencia en algo aun mas interesante y divertido, gracias de corazôn. A José y Helena, que, me acabo de enterar hace un momento, estais en camino de ampliar la familia ;Felicidades pareja!. A Eva, espero que todo te siga yendo bien. Te lo mereces. A Ainhoa, creo que tu seras la siguiente, ^no?. Mucho ânimo y a por ello. A Guillermo, a puntito de regresar del periplo americano. Nos veremos pronto companero. A Judit, siempre bienvenida, pronto nos darâs clases de gramâtica espanola. Kôszônôm. A Jesus, me imagine que tan liado como siempre, espero que también sigas feliz y con la energia de te caracteriza. Te sigo debiendo una visita. A Jorge, parece que fue ayer cuando empezaste a venir por el Museo. Veras, en menos de lo que te piensas estaras tu escribiendo los agradecimientos de tu tesis. A David, sinceramente, es una alegrfa tenerte de vuelta por Madrid. Se te ha echado de menos tronco. A Inigo, igualmente, un placer volver a tenerte por aquf cerca. Espero que no tardemos mucho en poder felicitarte por haber conseguido una plaza. A Chiara, gracias a tu ânimo y apoyo por fin he terminado con la tesis. Un bacio bella. Agradecido quedo por supuesto a todos los amigos y companeros que he encontrado en el Museo: Cristina, Elena, Fernando, Patri, Raquel, Silvia, Lourdes, Felipe, Annie, Pepe, Anabel, Carolina, Jesus, Iker, David, Regina, Diego, Lukas, Fede, Teresa, Carlos, Ivan, Angela, Anabela, Jose, Sebastian, Fafa,... A Paty y Carlos, companeros de fatigas en el principio del fin, habéis conseguido que haya sido mas facil y menos estresante. Muchas gracias. Un montôn de gracias a la gente que me ha hecho sentir como en casa en mis estancias en México: Ruth, Paty, Edna, Noemi, Lisbeth, Juan Carlos, Gerardo,... Ha sido un placer haberles conocido. A mis amigos del alma, que estoicamente me han soportado estos anos y siempre han estado ahi cuando se les necesita: Raul, Pedro, Inigo, Santi, David, Cebollo, Noemi, Diego, Carlos, Rafa, Javi, Miguel Àngel, Nacho, Juan, Itziar, Luis, Alex. Chavales, esto habrâ que celebrarlo. Finalmente, a mi familia, a quien dedico esta tesis, con todo mi carino. Muchfsimas gracias por todo, hacéis que el Mundo sea mucho mejor. I. Introduccion General 11- -12- Intfodiiccinii La biologia evolutiva aborda el estudio de la evoluciôn desde dos perspectivas. Por un lado trata de reconstruir e interpretar la historia evolutiva de los organismos. Por otro busca elucidar los procesos y mecanismos generadores de dicha historia (Futuyma, 1986; O’hara, 1988). A menudo ambas Imeas de estudio se mantienen separadas por las dificultades a la hora de integrar diferentes ramas de investigacion, debido principalmente a importantes diferencias en las metodologias y objetivos propios (Schlichting & Pigliucci, 1998). En algunos casos, sin embargo, el estudio de la historia evolutiva y sus procesos causales van unidos de la mano. Asi, por ejemplo, el estudio de la diversidad intraespecifica suele implicar una parte descriptiva en la que se establecen las caracteristicas de los diferentes linajes asi como las relaciones entre ellos y otra parte que trata de explicar los procesos que han dado origen a dicha diversidad (Carstens et a i, 2009) . Consecuencia directa de la evoluciôn es la diversidad que caracteriza a los seres vivos. Sin embargo, esta diversidad no siempre résulta évidente. Suele ser asi el caso de la existente entre poblaciones de una misma especie o incluso de organismos estrechamente relacionados. El desarrollo de técnicas - 13- Introduccion moleculares, especialmente desde el ultimo cuarto del siglo XX, ha permitido explorar un vasto campo de diversidad, a menudo criptica, que en muchos casos resultaba inconcebible no hace muchos anos (Beheregaray & Caccone, 2007; Trontelj & Fiser, 2009). Se trata de una revolucion que ha afectado a todos los niveles y muy especialmente en aspectos bâsicos como por ejemplo en el eterno debate sobre el concepto de especie (Wiens, 2004a; Vences & Wake, 2007). De este modo el estudio de los patrones de variacion genética intraespecifica se ha convertido en un elemento clave que esta permitiendo un progreso considerable en nuestros conocimientos sobre los factores que influyen en las estructuras poblacionales, en la definiciôn de linajes y en los procesos de especiaciôn. En las ultimas décadas este tipo de estudios se han visto favorecidos notablemente no solo por el continue desarrollo de las técnicas moleculares que permiten cada vez mas la generaciôn de gran cantidad de datos acordes a los objetivos planteados en cada proyecto, sino también gracias a la apariciôn de nuevos marcos conceptuales en los que analizar e interpretar dichos resultados (Avise, 2000; Wiens, 2004a; Buckley, 2009; Hickerson et a l, 2010). - 14- Introduction Filogeograjïa y diversificaciôn. La filogeografia es una disciplina que estudia los patrones de distribuciôn geogrâfica de los linajes genéticos, normalmente a nivel intraespecifico o entre especies proximamente emparentadas, asi como los procesos que generan dichos patrones (Avise, 2009; Nielsen & Beaumont, 2009). Al estudiar la diversidad intraespecifica median te una aproximaciôn filogeogrâfica, la distribuciôn espacial de los linajes encontrados debe analizarse en un trasfondo causal que permita conjeturar que procesos histôricos y demogrâficos se encuentran detrâs (Buckley, 2009).
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