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Asteraceae, Cardueae) UNIVERSITAT DE BARCELONA. FACULTAT DE FARMÀCIA Departament de Productes Naturals, Biologia Vegetal i Edafologia, Secció Botànica Programa de Doctorat: Biodiversitat 2009-2012 INSTITUT BOTÀNIC DE BARCELONA (IBB-CSIC-ICUB) Estudio biosistemático y evolutivo del género Echinops L. (Asteraceae, Cardueae) Memòria presentada per Ismael Sánchez Jiménez per a optar al títol de Doctor per la Universitat de Barcelona Amb el vist-i-plau de les directores de tesi: Dra. Teresa Garnatje Roca Dra. Oriane Hidalgo i del tutor de tesi: Dr. Joan Vallès Xirau Ismael Sánchez Jiménez Barcelona, 2012 ¿Por qué subir al Everest? Porque está ahí... George Mallory ÍNDICE Agradecimientos 7 Introducción 11 1. Marco taxonómico 11 1.1. La familia Asteraceae Martinov 11 1.2. La tribu Cardueae Cass. 15 1.3. La subtribu Echinopsinae (Cass.) Dumort. y el género Echinops L. 17 2. Antecedentes 23 2.1. Recuentos cromosómicos y cantidad de ADN 23 2.2. Palinología 25 2.3. Sistemática molecular 29 Objetivos de la Tesis Doctoral 33 Informe de las directoras de la Tesis 35 Síntesis y discusión de los resultados 39 Conclusiones 51 Referencias 53 Compendio de publicaciones 63 Chromosome Numbers in Three Asteraceae Tribes from Inner Mongolia (China), with Genome Size Data for Cardueae 65 Molecular systematics of Echinops L. (Asteraceae, Cynareae): A phylogeny based on ITS and trnL-trnF sequences with emphasis on sectional delimitation 83 Genome size and chromosome number in Echinops (Asteraceae, Cardueae) in the Aegean and Balkan regions: technical aspects of nuclear DNA amount assessment and genome evolution in a phylogenetic frame 99 Echinops spinosissimus Turra subsp. neumayeri (Vis.) Kožuharov (Asteraceae, Cardueae): a new record for the flora of Greece 117 Pollen study in the genus Echinops L. (Asteraceae, Cardueae) 123 ANEXO: Otros trabajos derivados de la Tesis Doctoral 139 Póster: Genome size of Echinops L. (Asteraceae, Cardueae) in the west Balkans: an attempt to elude the fresh material requirement 141 Publicación: Polyploidy and other changes at chromosomal level and in genome size: Its role in systematics and evolution exemplified by some genera of Anthemideae and Cardueae (Asteraceae) 145 AGRADECIMIENTOS Este trabajo no habría sido posible sin la colaboración y el apoyo de muchas personas que han desempeñado un papel fundamental en el desarrollo de esta tesis, desde el ámbito profesional y científico al personal y anímico. En primer lugar, tengo mucho que agradecer a mis directoras de tesis, la Dra. Teresa Garnatje y la Dra. Oriane Hidalgo, por la dedicación y empeño que han mostrado para que este trabajo salga adelante. Desde que me brindaron la oportunidad de realizar un trabajo de investigación para el Proyecto de Fin de Carrera, que tuvo su continuidad en este proyecto de tesis, hasta el apoyo científico y moral que me han dado durante estos años y, especialmente, en estos últimos meses. La labor que han realizado durante estos años es inmensurable, dirigiendo muy activamente la tesis y ayudándome en todo momento a todos los niveles, haciendo en ocasiones las tareas más espinosas en alguna de las recolecciones, o acogiéndome y dando cobijo en los momentos iniciales, y no tan iniciales, de la estancia que realicé fuera de Barcelona. La tesis tampoco habría sido posible sin la cooperación del Dr. Joan Vallès, por la colaboración y ayuda en los diferentes trabajos de estos años, así como por ejercer de tutor. Las demás personas del grupo de investigación en Biosistemática y Filogenia y Citogenética Moleculares de plantas han jugado un papel importante en la tesis, puesto que han participado en el desarrollo de 7 Estudios biosistemáticos en Echinops alguno de los trabajos que aquí se presentan, así como otras colaboraciones más allá de la tesis, además de disfrutar de la cotidianeidad de algunas de estas personas: Sònia Garcia, Jaume Pellicer, Daniel Vitales, Gemma Mas de Xaxars, Esperança Carrió, Alfredo García-Fernández, Joan Martín, Miguel Ángel Canela, y Sonja Siljak-Yakovlev, quien desde París se ha mostrado siempre interesada por el buen desarrollo de esta tesis, contribuyendo con la recolección de valiosísimos materiales. Al personal del Institut Botànic de Barcelona también le tengo que agradecer su contribución al desarrollo de esta Tesis, puesto que son éstas las personas con las que he estado conviviendo en el trabajo durante estos años, y que, en numerosas ocasiones, me han ofrecido ayuda ya sea en el laboratorio, en el ordenador u otros aspectos: Laia, Andreas, Sara, Elisabet, Encarni, Javi, Jose, Carmen, etc... Al Dr. Stephan Gleissberg le agradezco haberme permitido realizar una estancia en su laboratorio de Athens (Ohio), a pesar de que el trabajo allí realizado no haya sido incluido finalmente en esta tesis. Alastair Plant y Conny Bartholmes, también me ayudaron en el uso del laboratorio durante mi estancia en la Universidad de Ohio, bajo la dirección de una de mis directoras, Oriane, que estaba allí haciendo un PostDoc. En estos años he tenido la oportunidad de realizar varios viajes de recolección. Éstos no habrían sido posible sin la colaboración de algunas personas: Teresa Garnatje, Sonja Siljak-Yakovlev, Marija Edita Šolic y Ángel Romo. Gracias también a las personas e instituciones que me han enviado material, como Georgy Lazkov, y los/las responsables de los herbarios que me han dejado examinar numerosos pliegos, ya sea presencialmente o enviándome los materiales: B, E, ERE, G, GB, LE, MA, P, PE, W, WU, YA, y, por supuesto al personal del herbario del Instituto Botánico de Barcelona, especialmente Jordi Pallàs, por gestionarme todos estos intercambios. Son varias las personas que me han ayudado en aspectos técnicos, imprescincibles para el desarrollo del trabajo: Miquel Veny, por atender la colección de plantas vivas resultado de las recolecciones realizadas, Samuel Pyke, por las correcciones del inglés de las publicaciones, y el personal técnico de citometría de flujo y de microscopía electrónica, de los Centros Científicotécnicos y Tecnológicos de la UB, por el asesoramiento cuando ha sido necesario. Por otra parte, fuera del mundo académico, son muchas las personas con las 8 Ismael Sánchez Jiménez que he pasado buenos momentos, y que me han animado a seguir adelante con esto, la gente del Barrio del Pilar en Madrid y la que he conocido en Barcelona durante estos últimos años, en especial a la gente del Clot por el tiempo pasado y los ánimos finales que me han servido para mirar de cara el final de la tesis cuando parecía que no iba a tener fin. Es inevitable agradecer a Maria todo el tiempo que ha pasado conmigo durante casi todo el desarrollo de la esta tesis. Desde el recuerdo de los buenos momentos que hemos pasado cuando hemos vivido juntos y con la esperanza de encontrar otros en el futuro, no puedo sino agradecer y dedicar este trabajo a esta persona que ha tenido un papel tan importante en mi vida durante 4 de los últimos años. Finalmente, tengo que agradecer a mi padre y mi madre, Miguel y Ángeles, por el apoyo que me han dado todos estos años, y a Aratz, por la compañía que me hace cada día. 9 INTRODUCCIÓN 1. MARCO TAXONÓMICO 1.1. LA FAMILIA ASTERACEAE MARTINOV La familia Asteraceae (Compositae Adans.) es la de mayor extensión entre las angiospermas, comprendiendo 1.500-1.700 géneros y alrededor de 23.000- 30.000 especies (Bremer, 1994; Funk et al., 2005, 2009). Se distribuye por todo el globo salvo por el continente Antártico, aunque es especialmente diversa en las regiones tropicales y subtropicales de Norteamérica, los Andes, Brasil oriental, la región mediterránea, Asia central y el suroeste de China (Panero & Crozier, 2008). Las características diferenciales de este grupo de angiospermas son (Bremer, 1994; Funk et al., 2005): - organización de las flores en una estructura pseudoantial muy particular, el capítulo, consistente en un receptáculo donde se asientan todas las flores. Ontogénicamente se trata de un racimo cuyos ejes permanecen cortos, ensanchados y fusionados, lo cual explica la floración centrípeta de la inflorescencia (Weberling, 1992). Además, el receptáculo se encuentra rodeado de un conjunto de brácteas involucrales (Fig. 1A). 11 Estudios biosistemáticos en Echinops - fusión de las anteras del androceo, formando un anillo. Esta disposición estructural permite que durante la polinización la presentación del polen sea secundaria; esto es, el polen es depositado en el interior del anillo anterial por la dehiscencia introrsa de las anteras, empujado por la acción del estilo y posteriormente presentado en la superficie externa del fuste y ramas del mismo, que se abre exponiendo las áreas estigmáticas en la superficie adaxial (Fig. 1B). - fruto monocarpelar indehiscente, el aquenio o cipsela, generalmente coronado por un anillo de pelos o escamas procedentes de la modificación del cáliz, conocido como vilano (Figs. 1C-E). El capítulo de las asteráceas presenta un diseño adaptado para llamar la atención de los polinizadores, haciendo más visibles las partes reproductivas; la unidad de atracción del polinizador, por tanto, pasa de ser una simple flor Figura 1. Características de la familia Asteraceae Martinov. A. Estructura del capítulo © 2008 Jose L. Panero (http://www.tolweb.org/asteraceae). B. Androceo singenésico y presentación secundaria © 2010 Ismael Sánchez-Jiménez. C-E. Diversos tipos de aquenios © 2007 Luc Viatur, 2011 Philmarin y 2009 Penarc, respectivamente (http://www. wikipedia.org). 12 Ismael Sánchez Jiménez a estar constituida por un grupo de flores (Harris, 1999). La complejidad de la “flor funcional” en Asteraceae puede ser incluso mayor, puesto que también se pueden encontrar capítulos compuestos (sincefalia) de segundo y de tercer orden (Jeffrey, 2007). La estructura de la inflorescencia, además, refleja un equilibrio entre varias funciones relacionadas con la polinización, el sistema de reproducción, la dispersión, germinación y defensa de la semilla (Stuessy & Garver, 1996). Las familia Asteraceae incluye organismos fundamentalmente Figura 2. Subfamilias de Asteraceae, ilustradas por un ejemplo: A.
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