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Filogenia Y Evolución De La Subtribu Melinidinae (Paniceae: Panicoideae: Poaceae)

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Filogenia Y Evolución De La Subtribu Melinidinae (Paniceae: Panicoideae: Poaceae) Tesis Doctoral Filogenia y evolución de la subtribu Melinidinae (Paniceae: panicoideae: poaceae) Salariato, Diego Leonel 2010 Este documento forma parte de la colección de tesis doctorales y de maestría de la Biblioteca Central Dr. Luis Federico Leloir, disponible en digital.bl.fcen.uba.ar. Su utilización debe ser acompañada por la cita bibliográfica con reconocimiento de la fuente. This document is part of the doctoral theses collection of the Central Library Dr. Luis Federico Leloir, available in digital.bl.fcen.uba.ar. It should be used accompanied by the corresponding citation acknowledging the source. Cita tipo APA: Salariato, Diego Leonel. (2010). Filogenia y evolución de la subtribu Melinidinae (Paniceae: panicoideae: poaceae). Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Cita tipo Chicago: Salariato, Diego Leonel. "Filogenia y evolución de la subtribu Melinidinae (Paniceae: panicoideae: poaceae)". Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. 2010. Dirección: Biblioteca Central Dr. Luis F. Leloir, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. Contacto: [email protected] Intendente Güiraldes 2160 - C1428EGA - Tel. (++54 +11) 4789-9293 UNIVERSIDAD DE BUENOS AIRES Facultad de Ciencias Exactas y Naturales FILOGENIA Y EVOLUCIÓN DE LA SUBTRIBU MELINIDINAE (PANICEAE: PANICOIDEAE: POACEAE) Tesis presentada para optar al título de Doctor de la Universidad de Buenos Aires en el área Ciencias Biológicas Diego Leonel Salariato Director de tesis: Osvaldo Morrone Director Asistente: Fernando O. Zuloaga Consejero de Estudios: Patricia S. Hoc Lugar de trabajo: Instituto de Botanica Darwinion – CONICET (IBODA) Buenos Aires, 2010 FILOGENIA Y EVOLUCIÓN DE LA SUBTRIBU MELINIDINAE (POACEAE) RESUMEN FILOGENIA Y EVOLUCIÓN DE LA SUBTRIBU MELINIDINAE (PANICEAE: PANICOIDEAE: POACEAE) La subtribu Melinidinae (Poaceae: Panicoideae: Paniceae) incluye 13 géneros y aproximadamente 190 especies, las cuales se hallan caracterizadas por presentar la vía fotosintética C4 PCK y una anatomía foliar distintiva. El objetivo de este trabajo es estudiar las relaciones filogenéticas y la historia evolutiva de este grupo. Para esto se propone analizar a la subtribu desde tres diferentes niveles taxonómicos. En una primera parte se estudia la relación de Melinidinae con los demás integrantes de la tribu Paniceae, poniendo a prueba su monofilia, resolviendo la relación entre la subtribu y su grupo hermano y reconstruyendo la vía fotosintética ancestral. En una segunda parte se estudian las relaciones filogenéticas entre los distintos géneros incluidos en Melinidinae y se reconstruye la tendencia evolutiva que presenta la subtribu en cuanto al grado de homogenización de sus inflorescencias. Finalmente, se analiza en la tercera parte del trabajo las relaciones existentes entre las especies de Urochloa, género con mayor número de especies dentro de la subtribu, con el objetivo de definir grupos infragenéricos naturales y caracteres morfológicos de valor taxonómico. En cada sección se utilizan métodos de análisis y caracteres adecuados para el nivel taxonómico que se estudia. Los resultados obtenidos muestran que la subtribu Melinidinae resulta monofilética cuando Urochloa venosa es excluida de ésta, mientras que la asignación del grupo y del tipo fotosintético ancestral son ambiguas. Los géneros Urochloa y Eriochloa resultan parafileticos y/o polifiléticos, mientras que géneros como Megathyrsus, Melinis y Moorochloa son monofiléticos. La homogenización de las inflorescencias en la subtribu parece ser la tendencia evolutiva favorecida por sobre el proceso inverso. Finalmente, el género Urochloa es, en su actual circunscripción, un grupo artificial carente de sinapomorfías morfológicas. No obstante, los análisis recuperan dentro de Urochloa varios grupos infragénericos ii FILOGENIA Y EVOLUCIÓN DE LA SUBTRIBU MELINIDINAE (POACEAE) RESUMEN definidos por diferentes caracteres morfológicos. Sobre la base de estos resultados se establece para las especies Urochloa acuminata, U. decidua y U. megastachya el nuevo género Rupichloa. Palabras clave: Brachiaria, Chaetium, Eriochloa, ETS, evolución reticulada, filogenia, fotosíntesis C4 PCK, Leucophrys, Megathyrsus, Melinidinae, Melinis, Moorochloa, Morfología, ndhF, Paniceae, Panicum, Poaceae, radiación rápida, rpl16, Rupichloa, Scutachne, Thuarea, Tricholaena, trnL-F, Urochloa, Yvesia. iii PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (POACEAE) ABSTRACT PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (PANICEAE: PANICOIDEAE: POACEAE) The subtribe Melinidinae (Poaceae: Panicoideae: Paniceae) includes 13 genera and approximately 190 species, which are characterized by having the C4 PCK photosynthetic pathway and a distinctive foliar anatomy. The aim of this work is to study the phylogenetic relationships and the evolutionary history of this group. For this purpose, the subtribe is analyzed through three different taxonomic levels. First, the relations of Melinidinae with other taxa included in the tribe Paniceae are studied, in order to test its monophyly, resolving the relation between the subtribe with its sister group and reconstructing the ancestral photosynthetic pathway. Secondly, the phylogenetic relationships between the genera included in Melinidinae and the evolutionary trend in the homogenization of the inflorescences are analyzed. Finally, the relations among the species of Urochloa, the largest genus in the subtribe, are analyzed in order to define natural infrageneric groups and morphological characters with taxonomic value. In each section, different analyses and characters are used, depending of the studied taxonomic level. The results obtained showed the monophyly of the subtribe Melinidinae when Urochloa venosa is excluded, while the assignation of the sister group and the ancestral photosynthetic pathway are ambiguous. Urochloa and Eriochloa are paraphyletic and/or polyphyletic genera, whereas Megathyrsus, Melinis and Moorochloa are monophyletic. The homogenization of the inflorescences in the subtribe seems to be the favored evolutionary trend over the inverse process. Finally, the genus Urochloa appears as an artificial group without morphological synapomorphies. Nevertheless, the analyses recovered several infrageneric groups defined by different morphological characters. Based in these results, Urochloa iv PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (POACEAE) ABSTRACT acuminata, U. decidua and U. megastachya, are included within the new genera Rupichloa. Key words: Brachiaria, C4 PCK photosyntetic pathway, Chaetium, Eriochloa, ETS, Leucophrys, Megathyrsus, Melinidinae, Melinis, Moorochloa, morphology, ndhF, Paniceae, Panicum, phylogeny, Poaceae, rapid radiation, reticulated evolution, rpl16, Rupichloa, Scutachne, Thuarea, Tricholaena, trnL-F, Urochloa, Yvesia v PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (POACEAE) AGRADECIMIENTOS Quiero expresar mi profundo agradecimiento a mis directores de tesis, los Drs. Osvaldo Morrone y Fernando O. Zuloaga, por darme la oportunidad de iniciarme en la investigación científica, por enseñarme todo lo referido a la sistemática y taxonomía de las gramíneas, por sus consejos, paciencia y por todo el apoyo brindado durante estos años. A la Dra. Liliana Giussani, por enseñarme todo lo referido a las técnicas de sistemática molecular, por su constante apoyo y su guía en lo que refiere a esta área. A la Dra. Lone Aagesen, por haber compartido conmigo sus conocimientos en filogenia, por su apoyo y por todo el tiempo compartido durante estos años. A los Drs. Ana Cialdella, Silvia Denham, Rosa Guaglianone, Raúl Pozner y Renata Reinheimer, por la innumerables enseñanzas y el valioso tiempo cedido para compartir conmigo sus conocimientos. Al Lic. Diego Degennaro, compañero y amigo del cual aprendí tanto. A todo el personal de Instituto de Botánica Darwinion, que me brindo su ayuda incondicional durante todo este periodo y por los buenos momentos vividos. A Fabian Tricarico y Ángel Fusaro, operadores del Microscopio de Barrido del Museo de Ciencias Naturales Bernardino Rivadavia, los cuales siempre me brindaron su gran ayuda y vasta experiencia. Este trabajo no hubiera sido posible sin el apoyo y la financiación del CONICET y la ANPCyT. Los viajes de campaña fueron también financiados por la fundación Myndel Pedersen y la National Geographic Society. Finalmente quiero agradecer a mi familia, quienes me han brindado su apoyo incondicional a lo largo de toda mi vida. A mi compañera Liliana, por su amor, apoyo, paciencia, comprensión, y por compartir esta maravillosa experiencia desde el principio hasta el final. vi PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (POACEAE) DEDICATORIA A mi viejo, Jorge. vii PHYLOGENY AND EVOLUTION OF THE SUBTRIBE MELINIDINAE (POACEAE) INDICE Abreviaturas utilizadas ix Introducción general 1 Capitulo 1: Filogenia y Evolución de Melinidinae dentro de la Tribu Paniceae 7 Introducción 8 Materiales y Métodos 11 Resultados 24 Discusión 34 Apéndice 1 47 Apéndice 2 52 Apéndice 3 60 Capitulo 2: Filogenia y evolución de los géneros incluidos en Melinidinae 61 Introducción 62 Materiales y Métodos 65 Resultados 80 Discusión 99 Apéndice 1 117 Apéndice 2 118 Apéndice 3 119 Apéndice 4 121 Capitulo 3: Filogenia de Urochloa usando métodos de evidencia total 127 127 Introducción 128 Materiales y Métodos 136 Resultados 144 Discusión 175 Apéndice A 188 Apéndice B 232 Apéndice C 239 Apéndice D 248 Apéndice E 250 Apéndice F 283 Conclusiones generales 296 Bibliografía
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