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Tesis Doctoral TESIS DOCTORAL Análisis poblacional morfométrico y genético, áreas de solapamiento y distribución de los pejerreyes de la Provincia Parano-Platense (Atheriniformes, Atherinopsidae) Lic. Mariela V. Cuello Facultad de Ciencias Naturales y Museo Universidad Nacional de La Plata Directora: Dra. Mirta L. Garcia Co Director: Dr. Guillermo Ortí 2020 A Mateo A mi Abuela Tota 2 Agradecimientos En primer lugar quiero agradecer a la Facultad de Ciencias Naturales y Museo y a la Universidad Nacional de La Plata por todos estos años de formación profesional, de grado y postgrado, en forma pública y gratuita. A Mateo, a mi abuela Tota, a mis padres, mis hermanas y sobrinos, por haber sido y ser el sostén de mi vida. Sin ellos, no hubiese podido continuar. Gracias infinitas. A Gustavo, por compartir estos últimos años y hacerme más feliz aún. A mi querida Directora, mi amiga, mi compañera de laboratorio, de charlas, de campañas, de congresos, de compras... Mirta, un agradecimiento muy especial y con muchísimo cariño, por haber creído siempre que esta tesis iba a llegar a su fin, por su apoyo incondicional y por sus enseñanzas, no sólo de Ictiología sino también de vida. A Lau y Belu por contar pejerreyes durante las campañas. A Luis por llevarnos, esperarnos y traernos a todos lados. Y a toda la familia Genchi García, por hacerme sentir parte de ella. A mi codirector Guillermo Ortí, por la posibilidad de realizar una pasantía en su laboratorio de la Universidad de Nebraska, en Lincoln y por las correcciones y aportes al capítulo de análisis molecular; a Julie Sommer por la ayuda con mi entrenamiento en técnicas moleculares. A Agustín Solari, Andrés Jaureguizar, Andrés Milessi, Federico Cortéz y a todos los chicos que participaron de las campañas, por haberse mojado, pasar frío y mil problemas pero siempre volver con material. A Gustavo Somoza y su equipo de laboratorio, por la buena predisposición para recibirme, por su ayuda y la donación de material del Intech. A José y Andrés Neuville, por guardarme muestras cada vez que salían de pesca por las lagunas pampeanas. A la familia Skare por estar siempre dispuesta a transportar frascos por toda la costa argentina. A Lucila Protogino, Darío Colautti, Adriana Almirón, Andrés Milessi, Juan Iwaszkiw, Francisco Firpo, Andrea Tombari y Alonso Medina por el aporte de material. A Lucila Protogino, Amalia Miquelarena, Gustavo Chiaramonte, Marcelo Loureiro y Marlise de Azevedo Bemvenuti por los préstamos de material de las distintas colecciones. A Roberto Menni por sus interesantes charlas en el laboratorio de Ictiología del Museo de La Plata. A Hugo López por darme lugar en la División Zoología Vertebrados durante mi beca doctoral. 3 A Mónica Rodriguez, por “acompañarme y acompañarla”; a Justina, Julia y Daniela por los buenos momentos. A Diego Nadalin por su buena predisposición con el material de colección. A Ceci Morgan, por pasarme los programas. A Vanina Villanova por las secuencias de citocromo b. A Mer Cuello por salvarme con las fotos y la traducción!! A Graciela Bailliet por apoyarme continuamente! A Claudio Bravi, Rita Santos, Laura Jurado, Marisol Schwab, Julieta Beltramo y todo el laboratorio de GMP, por recibirme y hacerme sentir que pertenezco al grupo desde siempre. A Juli Sánchez por su gran ayuda con los programas de análisis molecular. A Amanda y Silvia, por escucharme, aguantarme y motivarme para finalizar esta tesis. Gracias chicas! A los jurados Alejandra Volpedo, Darío Colautti y Javier García de Souza por sus valiosos aportes a este trabajo. Y a todas aquellas personas que desde mi ingreso a la facultad y hasta el día de hoy, han contribuido en mi formación profesional y personal. 4 Resumen Para la Argentina, han sido citadas cinco especies continentales del género Odontesthes: O. bonariensis, O. hatcheri, O. humensis, O. perugiae y O. retropinnis. Históricamente, las entidades presentes en la Provincia Parano-Platense, han presentado serios problemas sistemáticos y nomenclaturales, particularmente en la identificación, ya que las descripciones son poco precisas y además, se ha hallado hibridización entre algunas de las especies. Por otra parte, existen importantes problemas faunísticos, como por ejemplo los debidos a las conexiones de las poblaciones del Río de la Plata y los ambientes costeros restringidos como la Bahía Samborombón y la albúfera de Mar Chiquita. El objetivo general de este trabajo es realizar una revisión taxonómica de las entidades incluidas en el género Odontesthes presentes en la Provincia Parano-Platense, establecer la distribución de cada una de ellas y analizar los ejemplares cuyos caracteres no coiciden con el de las especies descriptas y que en general fueron colectados en ambientes con solapamiento de especies, mediante un análisis morfométrico y estadístico, que se complementa con un análisis molecular. Las muestras provinieron de dos fuentes, por una parte de campañas realizadas en diversas localidades de la Argentina; paralelamente, se revisaron las colecciones ictiológicas de varias Instituciones nacionales e internacionales cubriendo todo el rango de distribución conocido de las especies. Para el análisis morfométrico se aplicaron distintos tratamientos estadísticos; entre las técnicas de análisis multivariado, se emplearon el Análisis de Componentes Principales (ACP) y el análisis Cluster con el propósito de testear las variaciones inter e intraespecíficas y para evaluar si existieron diferencias entre los grupos se aplicó un análisis de similitud (ANOSIM). Los datos morfométricos y merísticos fueron tratados por separado en los análisis. El análisis molecular fue llevado a cabo con el estudio del gen mitocondrial citocromo b, a partir de muestras colectadas en la Bahía Samborombón, definida en este tesis como el mayor área de solapamiento de las especies tratadas. Se han analizado 4.087 ejemplares del género Odontesthes de 92 localidades de Argentina, Brasil y Uruguay y además, 40 ejemplares tipo y paratipos. En este trabajo se provee por primera vez una clave identificatoria con caracteres morfológicos y merísticos que incluye las cinco especies del género Odontesthes presentes en la región de estudio, registrando el mayor rango de tallas analizado. Con respecto al análisis de cada especie se determinó que Odontesthes bonariensis se identifica por la siguiente combinación de caracteres: inserción de la Dorsal 1 a nivel de la vertical proyectada desde el ano, distancia predorsal 1 entre el 50,5 y el 62,6 %LS, de 11 a 13 hileras de escamas en la línea transversal, de 11 a 14 hileras de escamas 5 entre la Dorsal 2 y la Anal, de 20 a 22 escamas alrededor del pedúnculo caudal y de 30 a 49 branquiespinas totales en el primer arco branquial. Odontesthes humensis se caracteriza por tener la inserción de la Dorsal 1 por delante de la vertical que se proyecta desde el ano, la distancia predorsal 1 entre el 53,3 y 60,1 %LS, el ojo grande (18,3 y 35,3 %LC), de 11 a 13 hileras de escamas en la línea transversal, de 5 a 8 escamas entre las aletas dorsales, de 11 a 13 hileras de escamas entre la Dorsal 2 y la Anal y entre 15 y 23 branquiespinas totales en el primer arco. En O. perugiae la aleta Dorsal 1 se ubica por detrás de la vertical que se proyecta desde el ano, hocico largo (22,7-37,6 %LC) mayor a la longitud del ojo, la distancia predorsal 1 es de 55,1 a 62,6 %LS, el hocico largo (22,7-37,6 %LC), de 8 a 9 hileras de escamas en la línea transversal, de 4 a 8 escamas entre las aletas dorsales, de 9 a 10 hileras de escamas entre la Dorsal 2 y la Anal, y entre 19 y 34 branquiespinas totales en el primer arco. Odontesthes retropinnis es una especie poco conocida para nuestro país y se caracteriza por tener la inserción de la aleta Dorsal 1 por detrás de la vertical que se proyecta desde el ano, la distancia predorsal 1 de 55,4 a 73,0 %LS, de 8 a 9 hileras de escamas en la línea transversal, de 3 a 6 escamas entre las aletas dorsales, de 8 a 9 hileras de escamas entre la Dorsal 2 y la Anal y entre 38 y 65 branquiespinas totales en el primer arco. Históricamente, entre los caracteres más importantes en la determinación de estas especies se destaca el número de branquiespinas del primer arco. A partir del análisis de una gran variedad de tallas, se determinó que este carácter por sí sólo no puede utilizarse en la identificación, ya que varía con la ontogenia durante la cual gran parte de las especies superponen el rango de variación del número de branquiespinas. Por otra parte, el análisis de alometrías durante el desarrollo evidenció que esta fuente de caracteres es también útil en la identificación de las especies del género Odontesthes. Los análisis multivariados evidenciaron la presencia de morfotipos intraespecíficos en O. retropinnis e interespecíficos entre O. bonariensis y O. argentinensis, que solapan su distribución en ambientes mixohalinos, como los es la Bahía Samborombón y se determinó que el único carácter que permite la identificación de las especies cuando se hallan en simpatría es el número de escamas alrededor del pedúnculo caudal (14 a 17 en O. argentinensis vs. 20 a 22 en O. bonariensis) En cuanto al análisis molecular de poblaciones que solapan su distribución en el Río de la Plata, se estudiaron ejemplares de O. bonariensis, O. argentinensis y morfotipos intermedios entre estas especies. Se determinó una alta riqueza haplotípica a partir del análisis del gen citocromo b. Por primera vez se determinan tres haplotipos compartidos entre O. bonariensis y O. argentinensis. Este trabajo de tesis representa el primer antecedente con material que incluye gran parte de la variabilidad de formas entre las dos especies, que se da en un ambiente mixohalino.
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