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Evolution of the Centaurea Acrolophus Subgroup

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Evolution of the Centaurea Acrolophus Subgroup Evolution of the Centaurea Acrolophus subgroup [Evolució del subgrup Acrolophus del gènere Centaurea] Andreas Hilpold ADVERTIMENT. La consulta d’aquesta tesi queda condicionada a l’acceptació de les següents condicions d'ús: La difusió d’aquesta tesi per mitjà del servei TDX (www.tdx.cat) ha estat autoritzada pels titulars dels drets de propietat intel·lectual únicament per a usos privats emmarcats en activitats d’investigació i docència. No s’autoritza la seva reproducció amb finalitats de lucre ni la seva difusió i posada a disposició des d’un lloc aliè al servei TDX. No s’autoritza la presentació del seu contingut en una finestra o marc aliè a TDX (framing). Aquesta reserva de drets afecta tant al resum de presentació de la tesi com als seus continguts. En la utilització o cita de parts de la tesi és obligat indicar el nom de la persona autora. ADVERTENCIA. La consulta de esta tesis queda condicionada a la aceptación de las siguientes condiciones de uso: La difusión de esta tesis por medio del servicio TDR (www.tdx.cat) ha sido autorizada por los titulares de los derechos de propiedad intelectual únicamente para usos privados enmarcados en actividades de investigación y docencia. No se autoriza su reproducción con finalidades de lucro ni su difusión y puesta a disposición desde un sitio ajeno al servicio TDR. No se autoriza la presentación de su contenido en una ventana o marco ajeno a TDR (framing). Esta reserva de derechos afecta tanto al resumen de presentación de la tesis como a sus contenidos. En la utilización o cita de partes de la tesis es obligado indicar el nombre de la persona autora. WARNING. On having consulted this thesis you’re accepting the following use conditions: Spreading this thesis by the TDX (www.tdx.cat) service has been authorized by the titular of the intellectual property rights only for private uses placed in investigation and teaching activities. Reproduction with lucrative aims is not authorized neither its spreading and availability from a site foreign to the TDX service. Introducing its content in a window or frame foreign to the TDX service is not authorized (framing). This rights affect to the presentation summary of the thesis as well as to its contents. In the using or citation of parts of the thesis it’s obliged to indicate the name of the author. UNIVERSITAT DE BARCELONA FACULTAT DE FARMÀCIA Departament de Productes Naturales, Biologia Vegetal i Edafologia, Secció Botànica Programa de Doctorat: Biodiversitat 2009–2012 INSTITUT BOTÀNIC DE BARCELONA (CSIC-ICUB) Evolution of the Centaurea Acrolophus subgroup [Evolució del subgrup Acrolophus del gènere Centaurea] Memòria presentada per Andreas Hilpold per a optar al títol de Doctor per la Universitat de Barcelona Amb el vist-i-plau dels directors de tesi: Dra. Núria Garcia Jacas Dra. Roser Vilatersana Lluch i del tutor de tesi: Dr. Cèsar Blanché Vergés Andreas Hilpold Barcelona, 2012 Jag frågade Jonatan, varför han måste ge sej ut på nånting som var så farligt. Han kunde ju lika gärna sitta hemma vid elden i Ryttargården och ha det bra. Men då sa Jonatan, att det fanns saker som man måste göra, även om det var farligt. "Varför då", undrade jag. "Annars är man ingen människa utan bara en liten lort" sa Jonatan. [I asked Jonathan, why he had to go away to something so dangerous. He could just sit at home by the fire in Ryttargården and have it good. But then Jonathan said it was something he must do, even if it was dangerous. ‘Why?’ I wondered. ‘Otherwise you’re not a human being but just a piece of dirt’ said Jonathan.] From Bröderna Lejonhjärta / The Brothers Lionheart, Astrid Lindgren (1973) • The concrete highway was edged with a mat of tangled, broken, dry grass, and the grass heads were heavy with oat beards to catch on a dog's coat, and foxtails to tangle in a horse's fetlocks, and clover burrs to fasten in sheep's wool; sleeping life waiting to be spread and dispersed, every seed armed with an appliance of dispersal, twisting darts and parachutes for the wind, little spears and balls of tiny thorns, and all waiting for animals and for the wind, for a man's trouser cuff or the hem of a woman's skirt, all passive but armed with appliances of activity, still, but each possessed of the anlage of movement. From: The Grapes of Wrath John Steinbeck (1939) Aquest treball ha estat possible gràcies a la concessió d’una beca predoctoral JAE del Consejo Superior de Investigaciones Científicas (CSIC) d’Espanya. La recerca ha estat parcialment finançada pels projectes de recerca del Ministerio de Ciencia y Inovación (CGL2007-60781/BOS i CGL2010-18631) i de la Generalitat de Catalunya (Ajuts a Grups de Recerca Consolidats 2009/SGR/00439), i per dues bosses de viatge del CSIC. Acknowledgements – Agradecimientos Quiero a agradecer a mis directoras de tesis Núria Garcia-Jacas y Roser Vilatersana por darme su apoyo incondicional desde el principio. Durante muchas horas de laboratorio, de campo y de discusiones cientificas me introdujieron con mucha paciencia en el mundo de la ciencia sistemática, equipándome con una base sólida de conocimientos botánicos y moleculares. A Alfonso Susanna por acompañar mi estancia en el Instituto Botánico, dándome mucha ayuda y consejos, pero también por la instrucción profunda en el mundo español. Agradezco a mi tutor Cèsar Blanché Vergés por su gran ayuda en todos los asuntos burocráticos. Muchas gracias a mis compañeros de trabajo, a Piotr Kosinsky y Konstantin Romaschenko por muchas horas de diversión deportiva, a Laia Barres que me acompañó durante todo el estudio, a Sara López por su ayuda, a Javier López que compartió mis dificultades con su grupo plantas, a María Sanz, que me mostró que vale la pena luchar para acabar una tesis. Además agradezco a Ismael Sánchez Jiménez, Noemi Montes, Almudena Galian, Jordi López, Cristina Roquet, Mercè Galbany Casals por acompañarme durante partes de mi vida laboral en Barcelona. Moltes gràcies a tots! Special thanks go to Peter Schönswetter, who joined my way into science from the beginning. In a research stay in Vienna I got introduced to phylogeographical methods and could improve my working efficiency. His working group, namely Pablo Escobar, Ruth Flatscher, Clemens Pachschwöll, Katharina Bardy, Manuela Winkler and Gerald Schneeweiss, made my stay immediately familiar. My thanks go also to Bengt Oxelman, University of Gothenburg, for integrating me into his working group and for opening my eyes for new horizons in biology. To Bernard Pfeil, of the same working group, I owe a large part of my systematic skills, but also the fundamental insight that the hard science, as dry as it may seem, can be fun. Thomas Marcussen is acknowledged for his openness to discuss whatever botanical and not botanical question emerged. Finally, only the active role of Filipe Sousa, Yann Bertrand, Elisabeth Sjöquist, Elisabeth Gondel, Anna Petri, Ulrika Rydberg, Atefeh Pirani, Zeynep Toprak Aydın, Henrik Nilson and Stephan Nylinder rendered the highly prosperous atmosphere at Botaniska possible. Tack så mycket för en ganska bra tid och jag hoppar att vi ser oss snart! Agradezco a Isabel Sanmartín que me recibió con los brazos abiertos en el Real Jardín Botánico de Madrid durante una estancia, donde pude terminar mi tesis de doctorado mientras podía aprovechar de una atmosfera científica muy productiva. Agradezco a Javier Fuertes Aguilar por la compañía en el Fomento, a Jose Luis Blanco-Pastor, Mario Fernández-Mazuecos, Isabel Liberal, Ricarda Riina, Angélica Bello y Marieta Aguilar por su compañía y muchos impulsos científicos. Estoy muy agradecido a Andrea Sánchez-Meseguer que me acompañó los últimos dos años de mi doctorado compartiendo muchas horas difíciles y felices conmigo. Además me ayudó mucho en el desarrollo de este trabajo. Ganz besonders möchte ich auch jenen danken die mich im Laufe meines Studiums tatkräftig unterstützt haben. Ein besonderer Dank geht an meine Eltern und an meine Geschwister. Als ich vor mehr als vier Jahren aufbrach um meine Doktoratsstudium in Spanien zu beginnen, war ich mir stets ihrer Unterstützung gewiss. Ein Dank auch all meinen Freunden in Südtirol, Innsbruck und Wien – viele von ihnen haben mich im Laufe der Jahre mit ihrem Besuch beehrt und vielen von ihnen verdanke ich meinerseits unvergessliche Urlaubstage. Stets gaben sie mir das Gefühl, dass es einen Platz in den Alpen gibt, an dem ich immer willkommen bin. Finally, I want also to thank the many newly gained friends who joined my way in Barcelona, sharing flats, hanging out, accompanying me for excursions by foot or by bike, namely are they, Alex, Richard, Marta, Georg, Marco, Laia, Maria, Jürgen, Magdalena, Christine, Kamil, Josep, Roberto, Estela, Ixone, Francesca, Corinna, Clothilde, Ewelina, Borja, Angela, Miriam, Sophie, Marion, Flavio, Ana Carolina, Raquel, Raphael and many more. Thank you all, muchas gracias, moltes gracies, grazie mille, vielen Dank, dziękuję bardzo, muito obrigado, merci beaucoup, donkschian. Index 1. Introduction 1 1.1 Centaurea within Cardueae 1 1.2 The genus Centaurea 1 1.2.1 Characterization, distribution and subdivision 1 1.2.2 Palynology of the genus Centaurea 4 1.2.3 Caryology of the genus Centaurea 5 1.3 The Circum-Mediterranean Clade (CMC) of the genus Centaurea 5 1.3.1 Characterization and subdivision 5 1.3.2 The Jacea-Phrygia group 6 1.4 The Centaurea group (=Acrolophus subgroup) 7 1.4.1 Distribution and ecology 7 1.4.2 Traditional, morphology based treatments and their limitations 7 1.4.3 Hybridization 9 1.4.4 Systematic treatments of the Centaurea group in light of molecular approaches 9 1.4.5 Species delimitation within the Centaurea group 9 1.4.6 Reproduction and pollination biology of the Centaurea group 10 1.4.7 Caryology of the Centaurea group 10 1.4.8 Phytochemistry of the Centaurea group 11 1.4.9 Economic importance of the Centaurea group 12 1.5 Justification for the presented work (in Spanish) 13 2.
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