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Taxonomía Y Filogenia De Trichocereus (Berg.) Riccob

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Taxonomía Y Filogenia De Trichocereus (Berg.) Riccob UNIVERSIDAD NACIONAL DE LA PLATA FACULTAD DE CIENCIAS NATURALES Y MUSEO ―Taxonomía y filogenia de Trichocereus (Berg.) Riccob. (Trichocereeae- Cactaceae)‖ Tesis presentada para optar al grado de Doctor en Ciencias Naturales de la Universidad Nacional de La Plata Bióloga M. Sc. Adriana Sofía Albesiano Hoyos Director: Dr. Roberto Kiesling Co-Directora: Dra. Susana Freire La Plata, Argentina Agosto 3 de 2015 COMISIÓN ASESORA Dr. Roberto Kiesling, Instituto Argentino de Investigaciones de las Zonas Áridas-IADIZA, CONICET. Dra. Susana Freire, Instituto de Botánica Darwinion-San Isidro. Prof. Titular, Cátedra de Botánica Sistemática II, Facultad Ciencias Naturales y Museo, Universidad Nacional de La Plata. DEFENSA ORAL Y PÚBLICA Lugar y Fecha: Calificación: TRIBUNAL Firma:………………………………………………………...Aclaración………… Firma:………………………………………………………...Aclaración………… Firma:………………………………………………………...Aclaración………… TABLA DE CONTENIDO AGRADECIMIENTOS .............................................................................................................................. iii RESUMEN............................................................................................................................................... iv ABSTRACT ............................................................................................................................................ viii I. INTRODUCCIÓN ................................................................................................................................... 1 II. MATERIALES Y MÉTODOS ................................................................................................................. 2 III. HISTORIA DEL GÉNERO / ANTECEDENTES HISTÓRICOS .............................................................. 2 IV. RELACIONES GENÉRICAS ................................................................................................................ 7 V. MORFOLOGÍA DEL GÉNERO Y GRUPOS RELACIONADOS.............................................................. 8 5.1. Caracteres vegetativos ..................................................................................................... 8 Habito ................................................................................................................................. 8 Tallo ................................................................................................................................... 8 Ramas ................................................................................................................................ 8 Costillas .............................................................................................................................. 8 Areolas ............................................................................................................................... 8 Espinas .............................................................................................................................. 8 Madera ............................................................................................................................... 9 5.2. Caracteres reproductivos ................................................................................................. 9 Flores y polinización ........................................................................................................... 9 Apertura floral ..................................................................................................................... 9 Longitud de las flores ........................................................................................................ 11 Frutos ............................................................................................................................... 12 Semillas ............................................................................................................................ 12 Polen ................................................................................................................................ 12 5.3. Números cromosómicos ................................................................................................. 15 VI. HIBRIDACIÓN ................................................................................................................................... 15 VII. DISTRIBUCIÓN GEOGRÁFICA Y ECOLOGÍA ................................................................................. 16 VIII. ETNOBOTÁNICA ............................................................................................................................ 16 IX. TRATAMIENTO TAXONÓMICO ........................................................................................................ 19 9.1. Observaciones de la especie tipo ................................................................................... 20 9.2. Clave para la identificación de las especies .................................................................... 25 9.3. Clave para la identificación de las especies de Argentina: .............................................. 29 9.4. Clave para la identificación de las especies de Bolivia, Ecuador y Perú .......................... 30 9.5. Clave para la identificación de las especies de Chile ...................................................... 31 9.6. Descripción de las especies y taxones infraespecíficos .................................................. 32 9.7. Especies excluidas ....................................................................................................... 168 i X. ANÁLISIS CLADÍSTICO ................................................................................................................... 177 10.1. Introducción ............................................................................................................... 177 10.2. Materiales y Métodos ................................................................................................. 178 10.3. Resultados ................................................................................................................. 182 10.4. Discusión ................................................................................................................... 186 10.5. Evolución de caracteres ............................................................................................. 195 XI. CONCLUSIONES GENERALES ...................................................................................................... 202 XII LITERATURA CITADA .................................................................................................................... 204 XIII ÍNDICE DE NOMBRES CIENTÍFICOS ............................................................................................ 214 XIV. APÉNDICES ................................................................................................................................. 226 14.1. Formulario para la toma de datos morfológicos en campo .......................................... 226 14.2. Especies estudiadas .................................................................................................. 226 14.3. Lista de caracteres morfológicos y sus estados .......................................................... 228 14.4. Índice de figuras ......................................................................................................... 233 14.5. Índice de tablas .......................................................................................................... 238 ii AGRADECIMIENTOS Esta tesis surgió de una serie de ideas compartidas con el Dr. Roberto Kiesling, que conformaron el proyecto ―Taxonomía y Filogenia del género Trichocereus‖. El apoyo constante y decidido de Roberto, como consejero y profesor a lo largo de la investigación fueron decisivos para la realización de este trabajo. A la Dra. Susana Freire por dirigir este proyecto con tanto interés, entusiasmo, y por contribuir a mi crecimiento profesional. Agradezco a Salvador Arias-M., Leopoldo Arrieta, Ayra Faúndez, Luis Faúndez, Omar Ferrari, Rodrigo Flores, Iris Peralta, Patricio Saldivia, Teresa Terrazas, Matin Terry y Monserrat Vázquez, por la ayuda que me brindaron tanto en las labores de campo y en el laboratorio. A los directores y curadores de los herbarios consultados (Stephan Beck, Manuel Belgrano, Norma Deginani, Laura Iharlegui, Eduardo Méndez, Inés Meza, Mélica Muñoz, Nora Muruaga, James Solomon y Luc Willemse), por la gentileza en la resolución de dudas, envió de imágenes y préstamo del material solicitado. A Detlev Metzing por el envío de algunas publicaciones históricas, a Daniel Schweich por el préstamo de dos de sus fotos de campo. A Cecilia Scoones, por la elaboración de las ilustraciones. A la Sociedad Alemana de Cactus y a la Sociedad Inglesa de Cactus, por los subsidios que posibilitaron el desarrollo de este trabajo. A la Organización de Estados Americanos-OEA por el otorgamiento de la beca Doctoral. A la Universidad Nacional de La Plata, especialmente a la Facultad de Ciencias Naturales y Museo por mi formación Doctoral. Agradezco a la Facultad de Ciencias Agrarias de la Universidad Nacional de Cuyo, especialmente al Laboratorio de Biología Molecular (Adrian Ruiz Leal), donde realicé el procesamiento de las muestras vegetales para la obtención de las secuencias de ADN de cloroplasto. Finalmente agradezco a mi familia, mi madre y mis hermanos por su apoyo y confianza, siempre constante y necesaria. iii RESUMEN El género Trichocereus se encuentra representado por
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