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Sistemática Molecular Del Género Fissurella En El Pacífico Sudoriental

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Sistemática Molecular Del Género Fissurella En El Pacífico Sudoriental UNIVERSIDAD DE SANTIAGO DE COMPOSTELA FACULTAD DE BIOLOGÍA Departamento de Bioquímica y Biología Molecular SISTEMÁTICA MOLECULAR DEL GÉNERO FISSURELLA EN EL PACÍFICO SUDORIENTAL Memoria que para optar al Grado de Doctor en Bioquímica y Biología Molecular presenta ALBERTO NICOLÁS OLIVARES PAZ Santiago de Compostela – España, 2007 MANUEL REY MÉNDEZ, Profesor Titular del Departamento de Bioquímica y Biología Molecular de la Universidad de Santiago de Compostela, Informa: Que la memoria adjunta, titulada: Sistemática Molecular del Género Fissurella en el Pacífico Sudoriental, que para optar al grado de Doctor en Bioquímica y Biología Molecular, presenta Don Alberto Nicolás Olivares Paz, ha sido realizada bajo mi dirección en los laboratorios del Departamento de Bioquímica y Biología Molecular de la Facultad de Biología. Considerando que constituye trabajo de Tesis, autorizo su presentación en Santiago de Compostela a cinco de enero de dos mil siete. El Doctorando, VºBº del Director Fdo.: Alberto N. Olivares Paz Fdo.: Dr. Manuel Rey Méndez Alberto N. Olivares Paz, 2007 AGRADECIMIENTOS Desde estas páginas deseo expresar mis más sinceros agradecimientos a todas las personas e instituciones que de forma directa o indirecta han colaborado en la realización de este programa de doctorado. Al doctor Manuel Rey Méndez, director de esta tesis por su permanente apoyo y dedicación. Por la oportunidad de haber ingresado a su unidad, por las facilidades otorgadas para la realización de las diferentes actividades llevadas a cabo durante mi estadía en ella, y por su sincera y franca amistad. A Javier Quinteiro, por entregarme horas de su trabajo en la orientación, planificación y diseño de las diferentes actividades experimentales. Por su permanente apoyo, enseñanza y consejos brindados durante la estancia en el laboratorio. Su experiencia, conocimientos y dominio del área temática, fueron el pilar fundamental para concluir esta memoria. A Jorge Rodríguez, por su esmerada ayuda y colaboración brindada en la realización de actividades del laboratorio, algunas de las cuales al principio se me hacían casi imposibles. A los colegas del departamento de Acuicultura de la Universidad de Antofagasta, por la generosidad para permitirme realizar estos estudios de doctorado y a aquellos que además suplieron mi actividad docente durante los años que permanecí alejado de la institución. A Oscar Zúñiga y Gladys Ramos, por su paciencia para ayudarme en la obtención y envío desde Chile de numerosos artículos. A todos los que colaboraron en la ardua labor de búsqueda y obtención de los ejemplares de las diferentes especies de lapas desde distintos puntos de Chile y de otros países, entre ellos y esperando no olvidar a nadie a Jorge Oliva, Eduardo Díaz, Andrés Olguín, Braulio Montanares, Alberto Paredes, 5 Filogenia del género Fissurella Francisco Rivera, José Sánchez, Rodrigo Burgos, Cesar Lodeiros, Julio Pérez y Nieves González. También, para quienes ayudaron en el procesamiento de los animales, disección y almacenamiento de las numerosas muestras: Paula Medina, Katherine Dawson, María Maldonado, Patricia Olivares y Dania Olivares. A mis hijas Dania y Patricia y a mi esposa María Angélica, por el constante estímulo para que nunca claudicara en esta nueva etapa de mi formación académica. Durante el período de realización de esta memoria, se ha recibido financiación para desarrollar diferentes proyectos que, en algunos casos, permitieron o facilitaron el desarrollo de este trabajo. Por ello, quiero agradecer a las diferentes instituciones su ayuda: A la Unión Europea por financiar el proyecto: QLK1-CT-2000-30476 (años 2001 a 2003). A los Ministerios de Ciencia y Tecnología (MCYT) y al de Educación y Ciencia (MEC), por la financiación de los proyectos: AGL2000-3115-CE y AGL2004-06411-CO2-02/ALI (años 2001 a 2007). A la Xunta de Galicia por la financiación de los proyectos: PGIDT01PXI20005PM y PGIDIT05PXIC20002PN (años 2001 a 2008). A las Instituciones que posibilitaron el financiamiento de mi estancia. Mis agradecimientos a la Universidad de Antofagasta, por las facilidades administrativas otorgadas. A MECESUP UANTOF 2004, por la beca otorgada durante los años 2004-2005. A la Universidad de Santiago de Compostela por la beca concedida en los años 2006 – 2007. Al Departamento de Bioquímica y Biología Molecular de la Universidad de Santiago de Compostela por las facilidades que me ha brindado para realizar este Programa de Doctorado. 6 Alberto N. Olivares Paz, 2007 ÍNDICE AGRADECIMIENTOS ................................................................................... 5 RESUMEN...................................................................................................... 13 SUMMARY..................................................................................................... 15 Listado de Figuras....................................................................................................17 Listado de Tablas .....................................................................................................21 INTRODUCCIÓN....................................................................................... 23 I. CONDICIONES DEL MEDIO Y REGISTROS PALEONTOLÓGICOS ....................................................................... 25 I.1. UBICACIÓN GEOGRÁFICA DEL ESTUDIO ........................................................26 I.2. ZONAS BIOCLIMÁTICAS ................................................................................27 I.3. CORRIENTES MARINAS .................................................................................30 I.4. PROVINCIAS BIOGEOGRÁFICAS.....................................................................33 I.5. REGISTROS PALEONTOLÓGICOS DE MOLUSCOS.............................................35 II. ANTECEDENTES GENERALES DE LOS MOLUSCOS............... 38 II.1. FILO MOLLUSCA ..........................................................................................38 II.1.A. Morfología y Sistemática.........................................................................38 II.2. CLASE GASTROPODA ...................................................................................45 II.2.A. Morfología ...............................................................................................45 II.2.B. Sistemática y filogenia de gasterópodos ..................................................47 II.3. SISTEMÁTICA Y FILOGENIA DE VETIGASTROPODA........................................51 II.4. SUPERFAMILIA FISSURELLOIDEA..................................................................54 II.4.A. Morfología ...............................................................................................54 II.4.B. Sistemática...............................................................................................55 II.4.C. Filogenia ..................................................................................................58 III. EL GÉNERO FISSURELLA EN EL PACÍFICO SUDORIENTAL................................................................................... 58 III.1. SISTEMÁTICA Y CLASIFICACIÓN ...................................................................58 III.2. GRUPOS DE ESPECIES....................................................................................63 III.3. ANTECEDENTES BIOLÓGICOS DE FISSURELLA ...............................................65 III.3.A. Taxonomía ...............................................................................................65 III.3.B. Distribución .............................................................................................68 III.3.C. Hábitat......................................................................................................70 III.3.D. Alimentación............................................................................................73 7 Filogenia del género Fissurella III.3.E. Epibiontes ................................................................................................75 III.3.F. Sexo .........................................................................................................76 III.3.G. Parasitismo gonadal .................................................................................77 III.3.H. Reproducción...........................................................................................78 III.3.I. Morfometría.............................................................................................79 IV. FISSURELLA COMO RECURSO ..................................................... 81 IV.1. DURANTE EL HOLOCENO ..............................................................................81 IV.2. EN EL PERÍODO ACTUAL ...............................................................................85 V. LOS CONCEPTOS DE ESPECIE Y EL PROCESO DE ESPECIACIÓN .................................................................................... 90 V.1. CONCEPTO DE ESPECIE .................................................................................90 V.2. MECANISMOS DE ESPECIACIÓN ....................................................................93 V.3. INFERENCIA FILOGENÉTICA..........................................................................96 VI. FILOGENIA MOLECULAR.............................................................. 98 VI.1. ATRIBUTOS DEL ADN..................................................................................98 VI.2. ADN MITOCONDRIAL...................................................................................99 VI.3. MUTACIÓN.................................................................................................103
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