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Mitogenómica Y Filogenia De Linajes De Gasterópodos Altamente Diversificados (Vetigastropoda, Neritimorpha Y Conoidea)

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Mitogenómica Y Filogenia De Linajes De Gasterópodos Altamente Diversificados (Vetigastropoda, Neritimorpha Y Conoidea) Mitogenómica y filogenia de linajes de gasterópodos altamente diversificados (Vetigastropoda, Neritimorpha y Conoidea) Juan Esteban Uribe Arboleda 2016 MITOGENÓMICA Y FILOGENIA DE LINAJES DE GASTERÓPODOS ALTAMENTE DIVERSIFICADOS (VETIGASTROPODA, NERITIMORPHA Y CONOIDEA) Juan Esteban Uribe Arboleda Tesis doctoral 2016 Mitogenómica y filogenia de linajes de gasterópodos altamente diversificados (Vetigastropoda, Neritimorpha y Conoidea) PROGRAMA DE DOCTORADO EN BIOLOGÍA Y CIENCIAS DE LA ALIMENTACIÓN MEMORIA PRESENTADA POR JUAN ESTEBAN URIBE ARBOLEDA PARA OPTAR AL GRADO DE DOCTOR POR LA UNIVERSIDAD AUTÓNOMA DE MADRID DIRECTORES: RAFAEL ZARDOYA SAN SEBASTIÁN JOSÉ TEMPLADO GONZÁLEZ TUTOR: ÁNGEL ANTONIO LUQUE DEL VILLAR ! ! ! ! ! ! 2! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! ! Para mi madre y mis abuelos ! 3! ! 4! AGRADECIMIENTOS Muchas personas e instituciones han contribuido a esta tesis doctoral, en un sentido u otro, y quiero en estas líneas agradecerles: Inicialmente agradezco a mis directores de tesis, que además de grandes personas son excelentes profesionales con un gran don para enseñar. Gracias de verdad, esto ha sido un experiencia enriquecedora, que seguro, recordaré toda mi vida. A mi tutor de la Universidad Autónoma de Madrid Ángel Luque, que siempre estuvo dispuesto a ayudarme y a corregirme. Al Museo Nacional de Ciencias Naturales de Madrid, por facilitar todo para que nosotros los becarios hagamos lo que hacemos. Incluyendo a todo el personal humano: investigadores, directivos, técnicos, servicios generales, a todos, gracias. Al Ministerio de España por la financiación de esta tesis doctoral a partir de los proyectos (CGL2010-18216;! CGL2013- 45211-C2-2-P) y la beca FPI (BES-2011-051469). Como también, al Muséum National d'Histoire Naturelle de París, institución que me ha recibido en tres ocasiones para hacer mis estancias de investigación que tanto han enriquecido mi formación. Estancias que fueron patrocinadas por becas del Ministerio de España (estancias cortas) y beca de la comunidad europea (Synthesys). Agradezco a los colaboradores expertos, que con sus sabios comentarios han mejorado cada uno de los escritos. Como también a los investigadores y amigos que tantos consejos me han dado: ! 5! Mario García-París, Diego San Mauro, Federico Abascal, Marta Barluenga, Borja Mila, Mónica Suarez, Ignacio Doadrio, Manuel J. Tenorio, Annie Machordom, Elena G. González, Cristina Grande, David Buckley, Iñigo Martínez, entre otros. A toda mis amigos y compañeros del museo, que de una u otra forma han hecho parte de este proceso, y que sin ellos hubiera sido imposible terminar esta tesis doctoral: Jorge Gutiérrez, Silvia Perea, Iván Acevedo, Miriam Casal, Mario García, Samuel Abalde, David Osca, Iker Irisarri, Paula Rodríguez, Lourdes Alcaraz, Michel Domínguez, Guillermo Friis, Manuel Sánchez, Diushi Keri, Ernesto Recuero, Etienne Kornobis, Paloma Mas, Pilar Ochoa, Adelmo Martins (seguro he olvidado a alguien!). Y a todos con los que he compartido, chicos, gracias. También, a la persona que me dio la oportunidad de iniciarme en el camino de la investigación, que tanto disfruto, gracias profe Lyda Castro. A la familia Uribe, que a pesar de la distancia han estado presentes dando palabras de ánimo que me han fortalecido: mi abuela Josefina que es un pilar de vida; mis tíos (Doralba, Ildegar, Cesar, Chela, Carlos), que hicieron de mi infancia el mejor recuerdo; mis primas (Sandrita, Dianita, y Naty), que quiero mucho; y en especial a mi tío Libardo Uribe, que no volveré a ver, pero que me dejó la imagen de su sonrisa y positiva visión de la vida. A la familia Arboleda: mis abuelos, mis alcahuetas incondicionales y comprensivos; las tías (Cristina, Mary Isabel, ! 6! Alejandra, Amanda), cada una con su punto especial; mi tío Humberto, que más que nadie dio ejemplo académico y su peculiar visión de la vida, que tanto me gusta; mis primos con los que crecí (Santy, Valen, Alejo, Susa, July, Laila, Mariana, Naty). La familia Toro, que son parte fundamental del proceso: Alba, Mauricio, Jorge Moreno, Maricel. Y mis primos (Julián, Andrés, Caro, Caliche, Juan Camilo). En especial a mi hermano Jorge Esteban. También, a muchas personas que han hecho parte de mi vida desde que llegué a España: Ferchito-Pilar, la familia de la Iglesia, Pablito, Isidro, a la gente de la “parcela”, María José y Albert y Leire. A todas esas personas que desde Colombia (o cualquier parte) han creído en este proceso y que incondicionalmente me han apoyado: Alexandra Granados, Jan Pájaro, Alberto Yepes, Lisset Gelves, Hadder Jiménez, Merlin, Ivania entre otros. Y como último y más importante: a mi madre Luz Stella (luz de estrella) que siempre ha sido lo que me gustaría ser. A mi abuelo Fabio que es promotor de mi inquietud por los estudios y de terquedad; A Carlos Alberto que ha hecho de padre; y a mis dos hermanos Rosana y David que siempre están en mi mente. También a mi Chabelita, que con su sonrisa me alegra la vida. Gracias, TOTALES. ! 7! ! 8! ÍNDICE DE CONTENIDOS ABREVIATURAS*.................................................................................*11! RESUMEN*.............................................................................................*13! SUMMARY*...........................................................................................*19! 1! INTRODUCCIÓN*..........................................................................*25! 1.1.! Clase*Gastropoda*.......................................................................*25! 1.1.1.! Generalidades!..................................................................................!25! 1.1.2.! Clasificación!......................................................................................!29! 1.1.3.! Filogenias!moleculares!.................................................................!35! 1.2.! Mitogenomas*de*Gastropoda*.................................................*38! 1.2.1.! Generalidades!..................................................................................!38! 1.2.2.! Filogenias!...........................................................................................!44! 1.2.3.! Reordenamientos!...........................................................................!46! 1.3.! Subclase*Vetigastropoda*SalviniKPlawen,*1980*..............*48! 1.3.1.! Superfamilia!Trochoidea!s.!l.!Rafinesque,!1815.!...............!50! 1.3.1.1.! Familia!Trochidae!Rafinesque,!1815.!.................................!52! 1.4.! Subclase*Neritimorpha*Golikov*&*Starobogatov,*1975.*54! 1.5.! Subclase*Caenogastropoda*Cox,*1960*.................................*58! 1.5.1.! Superfamilia!Conoidea!Fleming,!1822.!.................................!60! 1.5.1.1.! Familia!Conidae!Fleming,!1822.!...........................................!62! 2! OBJETIVOS*...................................................................................*65! 2.1.! Subclase*Vetigastropoda*.........................................................*66! 2.2.! Subclase*Neritimorpha*............................................................*68! 2.3.! Subclase*Caenogastropoda*.....................................................*69! 3! RESULTADOS*...............................................................................*71! 3.1.! PUBLICACION*1:*..........................................................................*71! 3.2.! PUBLICACION*2:*........................................................................*127! 3.3.! PUBLICACION*3:*........................................................................*157! 3.4.! PUBLICACION*4:*........................................................................*213! 3.5.! PUBLICACION*5:*........................................................................*265! 4! DISCUSION*................................................................................*315! 5! CONCLUSIONES*........................................................................*331! 6! BIBLIOGRAFÍA*.........................................................................*339! 7! APÉNDICES*................................................................................*353! ! 9! ! ! 10! ABREVIATURAS ADN Ácido desoxirribonucleico ADNmt Ácido desoxirribonucleico mitocondrial ADNr Ácido desoxirribonucleico ribosómico AIC Criterio de información de Akaike ARN Ácido ribonucleico ARNr Ácido ribonucleico ribosómico ARNt Ácido ribonucleico de transferencia atp6-8 Genes mitocondriales que codifican para las subunidades 6 y 8 de la ATPasa mitocondrial BI Inferencia bayesiana CAT modelo mixto Bayesiano con diferentes categorias cob Gen mitocondrial que codifica para el citocromo b cox1-3 Genes mitocondriales que codifican para las subunidades 1, 2 y 3 de la citocromo oxidasa dNTP Desoxirribonucleótidos Trifosfato H3 Gen nuclear que codifica para la histona H3 Kb Kilobases LBA Atracción de ramas largas mya Millones de años atrás MgCl2 Cloruro magnésico MgSO4 Sulfato magnésico ML Máxima verosimilitud mM Milimolar mt Mitocondrial nad1-6-4L Genes mitocondriales que codifican para las subunidades 1-6 y 4L de la NADH deshidrogenasa PCR Reacción en cadena de la polimerasa pb Pares de bases rrnL Gen mitocondrial que codifica para la subunidad mayor del ácido ribonucleico ribosómico rrnS Gen mitocondrial que codifica para la subunidad menor del ácido ribonucleico ribosómico SNG Secuenciación de nueva generación trnA Gen mitocondrial codificante del ácido ribonucleico de transferencia de Alanina trnC Gen mitocondrial codificante del ácido ribonucleico de transferencia de Cisteína ! 11! trnD Gen
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