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Crustacea: Decapoda: Galatheidae) from the Solomon and Fiji Islands (South-West Pacific)

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Crustacea: Decapoda: Galatheidae) from the Solomon and Fiji Islands (South-West Pacific) DEPARTAMENTO DE BIOLOGÍA DEPARTAMENTO DE BIODIVERSIDAD Y BIOLOGÍA EVOLUTIVA FACULTAD DE CIENCIAS MUSEO NACIONAL DE CIENCIAS NATURALES (CSIC) Sistemática e historia evolutiva de la familia Galatheidae (Crustacea: Anomura) en el Pacífico Sur Memoria presentada por PATRICIA CABEZAS PADILLA para optar al Grado de Doctora en Ciencias Biológicas Vº Bº Directora de Tesis Vº Bº Director de Tesis Dra. Annie Machordom Barbé Dr. Enrique Macpherson Mayol Vº Bº Tutora de Tesis Prof. Ángela Delgado Buscalioni Madrid, 2010 Para Fer, magnífico compañero de trabajo y mejor amigo Agradecimientos La presente Tesis Doctoral ha requerido un gran esfuerzo y dedicación y su finalización no hubiera sido posible sin la cooperación y el apoyo de las personas que a continuación citaré, muchas de las cuales han sido fundamentales para afrontar los momentos más difíciles. En primer lugar debo de dar las gracias a mis directores de Tesis, Annie Machordom y Enrique Macpherson por la confianza depositada en mí para comenzar este proyecto. Muchas gracias, Annie, por haber sido algo más que una directora a lo largo de estos años, por tu disponibilidad absoluta siempre que lo he necesitado y por haberme guiado hasta el final de este trabajo. A ti, Enrique, gracias por haberme introducido en el mundo de la taxonomía y por tu apoyo constante en la distancia. A todos aquellas personas que hicieron posible disponer del material necesario para llevar a cabo esta investigación: A. Crosnier, B. Richer de Forges, T. Yam Chan, K. Baba, y principalmente al Museo de Historia Natural de París (MNHN) por el préstamo de la mayor parte de los ejemplares aquí estudiados. Fue en el último año de carrera cuando aterricé en el Museo Nacional de Ciencias Naturales y desde entonces han pasado seis años, durante los cuales he tenido la oportunidad de crecer a nivel laboral y personal. El tiempo transcurrido entre los muros de esta institución ha dado para cientos de historias divertidas, las cuales he tenido el privilegio de compartir con los que hoy considero mis mejores amigos. En primer lugar, tengo que dar las gracias a Lourdes Alcaraz por ser mi mentora y por ayudar a estas manitas mías a desenvolverse con soltura y rapidez en el laboratorio. Gracias por tantas y tantas conversaciones sobre la vida, ya sabes que sigo pensando que eres una fuente inagotable de sabiduría. A Fernando Alda, Raquel Alvárez, Carlos Toledo, Chiara Settanni y María Calero tengo tantas cosas que agradecerles que no sé ni por dónde empezar. Sin duda alguna habéis sido el mejor apoyo durante este último año, gracias por levantarme cuando me caí y gracias por vuestra incondicional amistad. Os deseo lo mejor. Amparillo, pero qué buena gente eres, gracias por las risas, por las lágrimas y por todos los momentos que hemos compartido juntas. Guacimara, son muchas las historias que he vivido contigo; sin embargo, sabes que ninguna podrá igualar a aquella memorable noche de vajillas, andamios y cervezas… esa queda para el recuerdo. Meri Joe, un placer compartir contigo hogar y laboratorio, gracias por acogerme en el 8ºD. A mi George, por quedarte solo ante el peligro y aguantarme durante el proceso de escritura. Marta Calvo, sólo puedo decir que eres un “crack” y que ha sido todo un honor ser tu compañera de buceo, de trabajo y de confidencias. María del Mar Cañado, siempre al teléfono cuando se te necesita, aun en la distancia se os siente cerca a ti y a tu isla. Al resto de gentecilla que habitaron o habitan el Museo: Iván Acevedo, Anna Addamo, Pedro Abellán, Regina Cunha, Ainhoa Iraola, Iker Irisarri, Ernesto Recuero, Carlos Pedraza, Silvia Perea, Patricia Ornelas, Felipe Morcillo, Francisco Brusa, José Fernández, Ana Izquierdo, Lucía Arregui, Raquel González, Rodolfo Pérez, Adriana Bermúdez, Pilar Pavón, David Osca, Federico Abascal, Diana Delicado, Pilar Casado, Diego Llusía, Joaquín Soler, Isabel Polo, Sebastián Barrionuevo, Laura Gómez, Angela Abad… por echar un cable cuando ha hecho falta y por las fiestas y risas. Esto ha sido mucho más llevadero en vuestra compañía. A los que se fueron y volvieron: Noemí y Cris, gracias por vuestros consejos durante estos últimos meses y a Iñigo y David por la asistencia con algún que otro programa de ordenador creado para atormentar becarios, y como no por los torreznos y las cañas. Dolores Bragado, Pepe Templado, Rafael Araujo, Pepe Fernández, Anabel Perdices y Mario García-París, todos ellos grandes científicos y grandes personas, un placer haber trabajado con vosotros. Por último, y para terminar con el bloque museístico, me gustaría hacer una mención especial a Marina Alcobendas. Gracias por las sonrisas que nos brindaste durante años y por todos los consejos que otros estudiantes y yo pudimos recibir de ti. Te echamos de menos. Como parte de este período de formación, he tenido la oportunidad de trabajar en distintos países, lo cual me ha enriquecido enormemente a nivel laboral y me ha brindado la oportunidad de hacer grandes amigos: Machel, Jimmy, Rob, Chelsea, Mike, Sun, François, Jenna, Davin, Mariko, Rocío, Kareen, Ana, Irene y Antonio. Un verdadero placer compartir con vosotros horas de trabajo, conversaciones y cervezas desde Oriente a Occidente. A mis amigos de la Facultad Javi, Juan, Virgi, Mentxu, Pableras y las Elenas, puedo prometer y prometo que tras la defensa de esta Tesis no faltaré a nuestra cita semanal en La Bodega. A mi gente zoológica, por su apoyo durante todo este proceso, gracias por estar ahí. Ismael, Vir, Efrén, Charlie y Beti, gracias por vuestra amistad, los viajes, las películas y tantas y tantas horas de charleta. Ézaro, no podía terminar estos agradecimientos sin mencionarte, puesto que el último empujón de este trabajo te lo debo a ti. No te equivocaste cuando auguraste que esto era el comienzo de una gran amistad, y espérate, porque todavía nos queda lo mejor. Como no podía ser de otra manera, el mayor de los agradecimientos es para mi familia, especialmente para mis padres, María y Nicolás, y para mi hermano Roberto. Sois mi mejor apoyo y os quiero, gracias por vuestra infinita paciencia y por haberme apoyado en todas y cada una de las decisiones que he tomado a lo largo de estos años. El aguilucho ha remontado el vuelo. ÍNDICE Resumen .............................................................................................................................. 1 Introducción general ........................................................................................................ 5 Taxonomía .................................................................................................................. 5 Filogenia ..................................................................................................................... 10 Máxima verosimilitud Inferencia Bayesiana Máxima parsimonia Métodos de distancia Medida de soporte de árboles Marcadores moleculares ........................................................................................... 20 Marcadores mitocondriales Marcadores nucleares Biogeografía ............................................................................................................... 23 Familia Galatheidae (Crustacea: Anomura) .......................................................... 26 Caracteres morfológicos Hábitat Relaciones filogenéticas Distribución y origen del grupo Área de estudio .......................................................................................................... 37 Origen del océano Pacífico Origen de las islas Origen de la diversidad del Pacífico Objetivos .............................................................................................................................. 43 Capítulo I ............................................................................................................................. 45 Capítulo II ........................................................................................................................... 57 Capítulo III ......................................................................................................................... 89 Capítulo IV ....................................................................................................................... 149 Capítulo V ......................................................................................................................... 231 Discusión general ........................................................................................................... 281 Evaluación de la variabilidad morfológica .......................................................... 281 Nivel intergenérico Nivel interespecífico Evaluación de la variabilidad molecular .............................................................. 285 Detección de divergencias genéticas Análisis de las relaciones filogenéticas Historia evolutiva de la familia Galatheidae ....................................................... 294 Estasis morfológica Origen y diversificación del grupo Reconstrucción de la historia biogeográfica Consideraciones futuras ......................................................................................... 300 Conclusiones .................................................................................................................... 301 Referencias ....................................................................................................................... 305 Anexo I ............................................................................................................................... 321 Anexo II
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