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Verónica Fernanda Aros Navarro Valdivia – Chile 2016

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Verónica Fernanda Aros Navarro Valdivia – Chile 2016 FACULTAD DE CIENCIAS ESCUELA DE BIOLOGÍA MARINA PROFESORA PATROCINANTE Dra. Leyla Cárdenas T. Instituto de Ciencias Ambientales y Evolutivas PROFESOR CO-PATROCINANTE Isabel Valdivia R. Instituto de Ciencias Ambientales y Evolutivas PROFESOR INFORMANTE Luis Vargas Chacoff Instituto de Ciencias Marinas y Limnológicas TAXONOMÍA MOLECULAR DE LOS PARÁSITOS Clestobothrium crassiceps Y Anonchocephalus chilensis (CESTODA) EN SUS HOSPEDADORES DE LOS GÉNEROS Merluccius Y Genypterus EN CHILE Memoria de grado presentada como parte de los requisitos para optar al grado de Licenciado en Biología Marina y Título Profesional de Biólogo Marino. VERÓNICA FERNANDA AROS NAVARRO VALDIVIA – CHILE 2016 Dedicado a Mis padres Verónica y Christian & Hermanos Felipe y Melissa AGRADECIMIENTOS En primer lugar, agradecer al Proyecto Fondecyt 1140173 “Host parasite phylogeny, phylogeography and parasite fitness: Understanding the evolution pattern in marine parasites”, por el financiamiento de esta tesis. Agradecer a mi profesora guía Dra. Leyla Cárdenas, por su confianza y creer en mis capacidades, por su disposición siempre a enseñar y ayudar, por promover el trabajo en equipo e individual bajo un clima de respeto y amistad. Agradecer a la Dra. Isabel Valdivia, por toda la ayuda entregada en esta tesis, por su apoyo y disposición a enseñar y transmitir sus conocimientos. Darle las gracias a mi familia, a mi madre Verónica, por estar siempre a mi lado en todo este proceso académico, por tener paciencia y creer que yo podía. A mi padre, Christian, que siempre ha confiado en mí, ayudándome y estando a mi lado. A mis hermanos, Felipe y Melissa, que desde pequeños hemos permanecido juntos y nuestras carreras profesionales no fueron la excepción. Y a mi familia en general, mi abuelo, mi abuela, mis tías y tíos, primos, jamás me he sentido sola, porque he tenido una familia que me respalda. Mis amigas, Kamilla y Roxana. Comenzamos este camino juntas, y así vamos a terminarlo, sin ustedes muchas cosas no hubiesen sido posible, cuando una flaquea las otras la levantan. Siempre. A mis compañeros de laboratorio: Paulina, Mabel, Ramón, Patricia, Romina, Yethro, Tarin, Horacio; quienes me enseñaron y ayudaron cuando lo necesitaba, y a todos los que de alguna forma participaron e influyeron en mis años de estudio, profesores, compañeros de carrera y de facultad. A todos ustedes, GRACIAS ÍNDICE Pág. RESUMEN………………………………………………………………………………....1 ABSTRACT………………………………………………………………………………..2 1. INTRODUCCIÓN………………………………………………………………....3 1.1 Aspectos generales……………………………………………………………..3 1.2 Taxonomía de organismos parásitos………………………………………....4 1.3 Cestodos………………………………………………………………………..5 1.4 Clestobothrium crassiceps………………………………………………….….7 1.5 Anonchocephalus chilensis…………………………………………………....9 1.6 Hospedadores 1.6.1 Género Merluccius……………………………………………………10 1.6.2 Género Genypterus…………………………………………………....12 2. HIPÓTESIS……………………………………………………………………….14 3. OBJETIVO GENERAL………………………………………………………….14 4. OBJETIVOS ESPECÍFICOS……………………………………………………14 5. MATERIAL Y MÉTODOS……………………………………………………...15 5.1 Áreas de estudio y obtención de las muestras………………………………15 5.2 Extracción de ADN…………………………………………………………...16 5.3 Amplificación de ADN por PCR…………………………………………….18 5.4 Edición y ensamblaje………………………………………………………....21 5.5 Análisis filogenéticos………………………………………………………….21 6. RESULTADOS………………………………………………………….………...22 7. DISCUSIÓN……………………………………………………………….………31 8. CONCLUSIONES……………………………………………………….……..…35 9. BIBLIOGRAFÍA……………………………………………………………….....36 !" " RESUMEN Tradicionalmente el estudio de la parasitofauna, ha estado basada en el reconocimiento de especies a través de caracteres morfológicos y morfométricos. Sin embargo, este enfoque ha presentado una limitación debido a la presencia de especies crípticas, es decir, aquellas especies que poseen una morfología altamente similar pero que difieren genéticamente, lo que resulta en una errónea clasificación taxonómica. En la actualidad, con la implementación y desarrollo de técnicas moleculares, el desarrollo de la taxonomía asistida por marcadores genético moleculares ha permitido desarrollar la taxonomía de forma más objetiva y ha revelado una biodiversidad mayor a la supuesta. Clestobothrium crassiceps y Anonchocephalus chilensis son endoparásitos cestodos de la familia Bothriocephalidae. Ambas especies, han sido descritas morfológicamente, estableciéndose la presencia de C. crassiceps en peces del género Merluccius (M. gayi y M. australis) y de A. chilensis en peces del género Genypterus (G. australis, G. blacodes y G. maculatus). El objetivo de esta tesis es evaluar su taxonomía molecular, a través del uso de genes nucleares (18S rRNA y 28S rRNA), y responder la interrogante: ¿Es concordante la taxonomía tradicional con la taxonomía molecular en nuestros modelos de estudio? Los resultados para Clestobothrium crassiceps muestran un patrón de similitud genética de parásitos provenientes de un mismo hospedador, pero revelan dos probables grupos taxonómicos. Se sugiere que en peces del género Merluccius existirían al menos dos especies del género Clestobothrium. Un grupo correspondería a C. crassiceps encontrado en el hospedador Merluccius gayi. El segundo grupo, incorpora las muestras de parásitos colectados desde M. australis (C. splendidum y C. cristinae), sugiriendo que existiría solo un grupo taxonómico que necesita ser renombrado. Por otro lado, para Anonchocephalus chilensis, se observa un solo grupo taxonómico independiente del hospedador o zona geográfica, es decir, parásitos desde G. maculatus, G. chilensis y G. australis, serían un solo taxón. Sin embargo, la única muestra reportada para esta especie desde la base de datos GenBank reveló diferenciación genética entre esta y las muestras de este estudio, dificultando la caracterización taxonómica de este grupo. #" " ABSTRACT " Traditionally the study of parasitic fauna, has been based on the recognition of species through morphological and morphometric characters. However, this approach has presented a limitation because of the presence of cryptic species, that is, species that have a highly similar morphology but differ genetically, resulting in an erroneous taxonomic classification. Today, with the implementation and development of molecular techniques, development of assisted molecular genetic markers taxonomy has allowed to develop the taxonomy more objectively and revealed the alleged greater biodiversity. Clestobothrium crassiceps and Anonchocephalus chilensis are tapeworms of the family that parasitize Bothriocephalidae teleost fish. Both species have been described morphologically, establishing the presence of C. crassiceps in fishes of the genus Merluccius (M. gayi and M. australis) and A. chilensis in fishes of the genus Genypterus (G. australis, G. maculatus and G. blacodes). The objective of this thesis is to evaluate its molecular taxonomy, through the use of nuclear genes (18S rRNA and 28S rRNA), and answer the question: Is the traditional taxonomy consistent with the molecular taxonomy in our study models? The results for Clestobothrium crassiceps show a pattern of genetic similarity of parasites from one host but two likely reveal taxonomic groups. It is suggested that fish of the genus Merluccius exist at least two species of Clestobothrium. A group correspond to C. crassiceps found in the host Merluccius gayi. The second group includes samples of parasites collected from M. australis (C. splendidum and C. cristinae), suggesting that there would be only a taxonomic group that needs to be renamed. Furthermore, for Anonchocephalus chilensis, one host independent taxonomic group or geographical area, that is, parasites from G. maculatus, G. chilensis, G. australis, would be one taxon. However, the only sample reported for this species from the GenBank database revealed genetic differentiation between this and the samples of this study, making it difficult taxonomic characterization of this group. " $" " 1. INTRODUCCIÓN 1.1 Aspectos generales El parasitismo se puede definir como una estrecha relación ecológica, donde existe una dependencia fisiológica obligada desde el parásito hacia su hospedador (de Vienne et al., 2013). Se puede considerar que más del 50% de las especies son parasitas (Price, 1980; Windsor, 1998), estos son cada vez más reconocidos como componentes importantes de la ecología de sus hospedadores, además de su supervivencia, distribución poblacional, así como también, dentro del funcionamiento y estructura de un ecosistema (Marcogliese, 2004; Luque, 2008). El estudio de la parasitofauna a lo largo del tiempo se ha centrado especialmente en el reconocimiento de especies y su clasificación, basado principalmente en taxonomía tradicional, es decir, en caracteres morfológicos y morfométricos (Valdivia et al., 2010). La principal deficiencia dentro de este tipo de enfoque, es la dificultad de diferenciar individuos que presentan morfologías altamente similares (especies crípticas), lo que conlleva a un eventual error en la identificación, nominándolas como una misma especie (Mayr, 1996; Bickford et al., 2006; Pfeninger & Schwenk, 2007). Este problema ha sido evidenciado en el último tiempo, gracias a la implementación de nuevas tecnologías, por ejemplo, la evolución de la microscopía, así como el desarrollo de la sistemática molecular (Palumbi, 1996). Estas herramientas han permitido una reevaluación del estatus taxonómico dentro de muchos taxones, como por ejemplo, insectos, plantas, mamíferos, aves, anfibios, reptiles, así como también parásitos, entre otros (Baker
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