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Patrones Genómicos De Divergencia En El Complejo Anthothoe Del

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Patrones Genómicos De Divergencia En El Complejo Anthothoe Del Universidad de Concepción Dirección de Postgrado Facultad de Ciencias Naturales y Oceanográficas Programa de Doctorado en Sistemática y Biodiversidad Patrones genómicos de divergencia en el complejo Anthothoe del Hemisferio Sur: Implicancias para la delimitación de especies en las anémonas de mar (Orden Actiniaria) Tesis para optar al grado de Doctor en Sistemática y Biodiversidad CARLOS ALBERTO SPANO PÉREZ CONCEPCION - CHILE 2018 Profesor Guía: Dr. Daniel Gómez Uchida Dpto. de Zoología, Facultad de Ciencias Naturales y Oceanográficas Universidad de Concepción Profesor Co-Guía: Dra. Vreni Häussermann Escuela de Ciencias del Mar, Facultad de Recursos Naturales Pontificia Universidad Católica de Valparaíso Para mi esposa Gianinna Clementi. Todo el tiempo que destine a esta tesis fue tiempo que permanecí alejado de ella. Que este trabajo sea siempre un recordatorio de nuestra lucha por estar juntos ii AGRADECIMIENTOS Mi primer y más sentido agradecimiento es a Dios. Todo lo que soy y he podido construir hasta ahora ha sido solo posible gracias al apoyo de mi familia. Ellos siempre han sido mi mayor bendición. Agradezco profundamente al programa de Doctorado en Sistemática y Biodiversidad, a todos mis compañeros y profesores, y en especial a mi tutor Daniel Gómez Uchida. Su acogida, respaldo y amistad han sido fundamentales en el arduo camino al grado Doctoral. Nunca olvidare su ejemplo como profesor y, sobretodo, como persona. Debo incontables gracias a mi extraordinaria comisión de tesis: Vreni Häussermann, Lisa Seeb, Angie Díaz, Pedro Victoriano y Cristián Hernández. Ellos no solo me corrigieron cuando hizo falta sino que también me ayudaron con ánimo y paciencia cada vez que lo necesite. Esta tesis también es fruto de su trabajo académico. Gracias de corazón a cada amistad que forme durante estos últimos cuatro años en Concepción. Mención especial a todo el equipo del GEEC lab: Selim, Pablo, Mauricio, Diego, Mónica, Camila, Celia y Andrea; ellos dieron la cuota de alegría diaria que se hace necesaria para surcar un postgrado. Por la valiosa relación que entablamos, no puedo dejar de nombrar a Cristian Canales, Fernanda Oyarzún y Alejandra Panzera. Curioso es que en tan poco tiempo sienta tanto el aprecio de una amistad. Les agradezco profundamente sea así. Muchas gracias al Departamento de Zoología de la Universidad de Concepción así como a la Dirección de Postgrado por todo el apoyo brindado durante el transcurso del Doctorado. De la misma manera, agradezco al Seeb Lab de la Universidad de Washington, y en especial a Jim Seeb, Garrett McKinney y Carita Pascal. La inolvidable experiencia de mi Pasantía Doctoral fue lo que fue debido a ellos. Finalmente, gracias a CONICYT por haberme adjudicado la beca que hizo todo esto posible. iii Universidad de Concepción RESUMEN Patrones genómicos de divergencia en el complejo Anthothoe del Hemisferio Sur: Implicancias para la delimitación de especies en las anémonas de mar (Orden Actiniaria) Carlos Alberto Spano Pérez Las anémonas de mar (Orden Actiniaria) constituyen un grupo relativamente pequeño de invertebrados marinos caracterizados habitualmente como pólipos solitarios de cuerpo blando y llamativos colores. Si bien son miembros conspicuos del intermareal rocoso, aún es muy poco lo que se sabe respecto a los procesos biológicos que subyacen a su extraordinaria diversidad. En la primera mitad de la presente Tesis Doctoral se repasan las principales causas históricas y contemporáneas de esta situación, así como potenciales alternativas para abordar la delimitación de especies en Actiniaria. Mientras que las convergencias morfológicas y la falta de conocimiento sobre la estructura y evolución de los caracteres diagnósticos conforman algunos de los problemas centrales en este contexto, la distribución geográfica ha ido surgiendo recientemente como una poderosa variable explicativa ante los resultados genéticos. Esta inferencia se evaluó explícitamente en la segunda mitad de la presente tesis, poniendo a prueba la hipótesis de especie para los representantes del género Anthothoe del Hemisferio Sur. Información proveniente de todo el genoma demostró una alta diferenciación entre poblaciones, la cual estuvo mucho más asociada a la localidad de su procedencia que a variedades morfológicas intraespecificas. Procesos vicariantes históricos mediados por eventos dispersivos de larga distancia serían los responsables más probables de la divergencia de los linajes de Anthothoe. Nuestros resultados indican consistentemente que el aislamiento geográfico, incluso a pequeña escala, estarían determinando la formación de especies en anémonas de mar, aún cuando esta diferenciación no sea detectable a través de su apariencia externa. iv TABLA DE CONTENIDOS INDICE DE TABLAS ........................................................................................................ vii INDICE DE FIGURAS ..................................................................................................... viii INTRODUCCION GENERAL ............................................................................................ 1 CAPÍTULO 1...................................................................................................................... 16 Abstract ........................................................................................................................... 17 Introduction .................................................................................................................... 18 The polymorphic clade Cnidaria ................................................................................... 19 The species problem in sea anemones ............................................................................ 21 Taxonomy through time ................................................................................................. 23 Early classifications ...................................................................................................... 24 Descriptions, re-descriptions and multidisciplinarity ..................................................... 26 Species concepts and delimitation .................................................................................. 28 The Actiniaria conundrum ............................................................................................. 29 What is a species in sea anemones? ............................................................................... 33 Integrated approaches and future directions................................................................. 37 The multispecies coalescent ........................................................................................... 38 Testing biogeographic hypotheses for species delimitation ............................................ 40 Conclusions and future perspectives .............................................................................. 44 Acknowledgements ......................................................................................................... 45 References ....................................................................................................................... 46 CAPÍTULO 2...................................................................................................................... 64 Abstract ........................................................................................................................... 65 Introduction .................................................................................................................... 66 v Material and Methods .................................................................................................... 70 Specimen collection and DNA sequencing ..................................................................... 70 Determination of the divergence parameters ................................................................. 70 Lineage tree inference ................................................................................................... 71 Species delimitation and species tree reconstruction ..................................................... 72 Evaluation of biogeographic hypotheses ........................................................................ 73 Population structure and detection of candidate loci for local adaptation ..................... 74 Results ............................................................................................................................. 76 Divergence patterns and lineages tree inference ............................................................ 76 Species delimitation and species tree reconstruction ..................................................... 77 Evaluation of biogeographic hypotheses ........................................................................ 78 Population structure and detection of candidate loci for local adaptation ..................... 78 Discussion ........................................................................................................................ 80 Genomic divergence in the southern Anthothoe complex ............................................... 80 The biogeography of speciation ..................................................................................... 82 Signals of selection ........................................................................................................ 84 Taxonomic considerations and future direction ............................................................. 85 Acknowledgements
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