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Tesis Amarilla, Leonardo David.Pdf (5.496Mb) Universidad Nacional de Córdoba Facultad de Ciencias Exactas, Físicas y Naturales Estudio Poblacional y Filogenético en Munroa (Poaceae, Chloridoideae) Lic. Leonardo David Amarilla Tesis para optar al grado de Doctor en Ciencias Biológicas Directora: Dra. Ana M. Anton Co-Director: Dr. Jorge O. Chiapella Asesora de Tesis: Dra. Victoria Sosa Instituto Multidisciplinario de Biología Vegetal CONICET-UNC Córdoba, Argentina 2014 Comisión Asesora de Tesis Dra. Ana M. Anton, IMBIV, Córdoba. Dra. Noemí Gardenal, IDEA, Córdoba. Dra. Liliana Giussani, IBODA, Buenos Aires. Defensa Oral y Pública Lugar y Fecha: Calificación: Tribunal evaluador de Tesis Firma………………………………… Aclaración…………………………………... Firma………………………………… Aclaración…………………………………... Firma………………………………… Aclaración…………………………………... “Tengamos ideales elevados y pensemos en alcanzar grandes cosas, porque como la vida rebaja siempre y no se logra sino una parte de lo que se ansía, soñando muy alto alcanzaremos mucho más” Bernardo Alberto Houssay A mis padres y hermanas Quiero expresar mi más profundo agradecimiento a mis directores de tesis, la Dra. Ana M. Anton y el Dr. Jorge O. Chiapella, por todo lo que me enseñaron en cuanto a sistemática y taxonomía de gramíneas, por sus consejos, acompañamiento y dedicación. De la misma manera, quiero agradecer a la Dra. Victoria Sosa (INECOL A.C., Veracruz, Xalapa, México) por su acompañamiento y por todo lo que me enseñó en cuando a filogeografía y genética de poblaciones. Además quiero agradecer… A mis compañeros de trabajo: Nicolás Nagahama, Raquel Scrivanti, Federico Robbiati, Lucia Castello, Jimena Nores, Marcelo Gritti. A los curadores y equipo técnico del Museo Botánico de Córdoba. A la Dra. Reneé Fortunato. A la Dra. Marcela M. Manifesto. A la Dra. Andrea Puebla y equipo de trabajo. A todas las personas que contribuyeron directa o indirectamente a esta tesis: Alejandra Romanutti, Diego Angulo, Leonardo Galetto, Martina Vaz Torres, Luciana Caeiro. A directivos y personal administrativo y técnico del IMBIV. A las entidades financieras: CONICET, UNC, MINCyT, Red Latinoamericana de Botánica-The Andrew W. Mellon Foundation, CONACyT (México) y Myndel Botanica Foundation A mis eternos profesores: la Dra. Lourdes Hirt y el Dr. Aníbal Amat. A mis hermanos por elección: Gisela Castro, Santiago Benzzo, Cristian de Candia, Leandro Gutiérrez y Sergio Villordo. A los amigos: María Kolman, Juan Urdampilleta, Gabriel Grilli, Ezequiel Saraceno, Laura Stiefkens, Silvana Monteverde, Lucia Acuña, Lorena Poelstra. Finalmente, quiero agradecer a mi familia… Natalia, compañera de vida e incondicional sostén. Mis padres, por la vida y la libertad. Alicia, por la entrega y por ser mi modelo de fortaleza. Évelyn, por las alegrías y por ser el motor de mis fuerzas. Índice Índice………………………………………………………………………………………..... i Resumen...………………………………………………………………………….……...... ii Abstract..…………………………………………………………………………………….. v Introducción………………………………………………………………………………… 1 Capítulo I: Estudios filogenéticos, cromosómicos y morfológicos en Munroa. 13 Antecedentes………………………………………………………………………………. 14 Materiales y Métodos……………………………………………………………………… 20 Resultados………………………………………………………………………………….. 31 Discusión……………………………………………………………………………………. 39 Conclusiones……………………………………………………………………………….. 44 Capítulo II: Estudios evolutivos y de biogeografía histórica en Munroa………. 51 Antecedentes………………………………………………………………………………. 52 Materiales y Métodos……………………………………………………………………... 56 Resultados…………………………………………………………………………………. 60 Discusión…………………………………………………………………………………… 64 Conclusiones…………………………………………………………………………….… 72 Capítulo III: Estudios filogeográficos y de modelado de nicho ecológico en Munroa argentina………………………………………………………………………… 74 Antecedentes………………………………………………………………………………. 75 Materiales y Métodos……………………………………………………………………... 79 Resultados…………………………………………………………………………………. 88 Discusión………………………………………………………………………………….. 101 Conclusiones……………………………………………………………………………… 108 Conclusiones generales……………………………………………………………….. 110 Anexos……………………………………………………………………………………. 114 Bibliografía……………………………………………………………………………….. 133 i Resumen Resumen Munroa comprende cinco especies de distribución disyunta en las regiones áridas y semiáridas de las Américas. Dasyochloa es un género monotípico común en el oeste de Norteamérica y afín a Munroa. La evaluación de los límites genéricos entre Munroa y Dasyochloa mediante el análisis conjunto de evidencia molecular, citológica y morfológica indicó la pertenencia de D. pulchella en Munroa y la transferencia incluyó la nueva combinación Munroa pulchella (Kunth) Amarilla. Munroa. Este género con dos taxones son norteamericanos (M. pulchella, M. squarrosa) y cuatro son sudamericanos (M. andina, M. argentina, M. decumbens, M. mendocina) forma un clado con dos sinapomorfías morfológicas y todas sus especies son diploides (x = 8). M. pulchella es el taxón hermano del resto de las especies y ni los taxones norteamericanos ni los sudamericanos resultaron en grupos monofiléticos. Munroa se habría comenzado a diversificar hace aprox. 7 millones de años (Ma) en América del Norte. Los estudios de evolución morfológica sugieren que los cambios en atributos tales como el hábito, la sexualidad, la desarticulación de la raquilla y la forma y textura de las glumas, han sido importantes para que estos taxones se dispersaran y establecieran en nuevas áreas. Para explicar su distribución disyunta proponemos dos eventos de dispersión a larga distancia; uno de América del Norte hacia América del Sur, seguido por un segundo en dirección opuesta. El establecimiento y diversificación de Munroa en Sudamérica posiblemente fue favorecida por los eventos orogénicos del Mioceno y del Plioceno, como así también por las oscilaciones climáticas del Pleistoceno. En Sudamérica, las especies de Munroa son anuales y ii Resumen crecen en la Zona de Transición Sudamericana (South American Transition Zone, SATZ) en tres provincias biogeográficas: la Puna, la Prepuna y el Monte. Se seleccionó a M. argentina para explorar acerca de las fuerzas evolutivas que modelan la distribución de sus poblaciones en esta zona. Los estudios filogeográficos recuperan tres haplogrupos (PAB, PrA, MA) que se distribuyen principalmente en cada una de las provincias biogeográficas. Los haplogrupos habitarían la SATZ desde hace aprox. 4 Ma y los estudios de paleomodelado mostraron que, en la actualidad, la distribución “potencial” de M. argentina es mayor a la distribución “real”; así mismo estimaron condiciones climáticas favorables en estas áreas durante el Periodo Interglaciar y el Último Máximo Glacial. Las poblaciones de M. argentina habrían experimentado, tempranamente un fuerte aislamiento genético debido a barreras intrínsecas como el tiempo de floración, asociadas al gradiente altitudinal y latitudinal en el que habitan. Este aislamiento genético se refleja en una profunda estructura filogeográfica cuyas poblaciones, sin embargo, son morfológica y cromosómicamente indistinguibles, lo que nos lleva a pensar que M. argentina ofrece otro ejemplo de especiación críptica. iii Abstract Abstract Munroa comprises five species of disjunct distribution in arid and semi-arid regions of the Americas and Dasyochloa, is a monotypic genera common in western North America. The evaluation of generic boundaries between Dasyochloa and Munroa through phylogenetic analyzes, and morphological and cytological studies showed unequivocally that D. pulchella belongs in Munroa, and the transference included the new combination Munroa pulchella (Kunth) Amarilla. Munroa is a monophyletic group and all species are diploid (x = 8) Munroa comprises two North American taxa (M. pulchella, M. squarrosa) and four South American taxa (M. andina, M. argentina, M. decumbens, M. mendocina). M. pulchella is the sister taxon of the rest species and no taxa occurring in North or South America resulted in monophyletic groups. The crown group of Munroa diversified about 7 Ma ago in North America and studies on the evolution morphological traits suggested that hanges in attributes such as habit, the the sexuality, the disarticulation of the rachilla, and the shape and texture of the glumes, have been important for these taxa to disperse and colonize new areas. Two events of long-distance dispersal are needed to explain the current distribution, one of North America to South America, followed by a second in the opposite direction. The arrival and establishment of Munroa in South America was possibly favored by the orogenetic events of Miocene and Pliocene, such as the climatic oscillations of the Pleistocene. In South America, the species of Munroa are annuals and grow in the South American Transition Zone (SATZ) in three biogeographic provinces: the Puna, the Prepuna and Monte. To explore about the v Abstract evolutionary forces affecting gene flow and the distribution of their populations in this zone, phylogeographic, population and ecological niche modeling studies were carried out in a species carefully selected of Munroa: M. argentina. Phylogeographic studies recovered three haplogroups (populations) whose distribution corresponds to the previously identified biogeographic provinces and indicated that these populations inhabit the SATZ about 4 Ma ago. Ecological niche modeling showed favorable climatic conditions in these areas during the Interglacial Period and the Last Maximum Glacial, and currently showing that the "potential" distribution of M. argentina is greater that the "real" distribution. M. argentina have undergone genetic isolation due to intrinsic
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