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Doctoral Thesis 2017 DEFENSE STRATEGIES AGAINST Doctoral Thesis 2017 DEFENSE STRATEGIES AGAINST HERBIVORY IN SEAGRASSES Gema Hernán Martínez 2017 Doctoral Thesis 2017 Doctoral Programme of Marine Ecology DEFENSE STRATEGIES AGAINST HERBIVORY IN SEAGRASSES Gema Hernán Martínez Thesis Supervisors: Fiona Tomas Nash Jorge Terrados Muñoz Doctor by the University of the Balearic Islands A todas las gotas a cada una de las olas que han contribuido a formar este mar A mi familia v vi Agradecimientos / Acknowledgements Esta tesis no hubiese sido posible sin los impuestos de los españoles y europeos que han contribuido a pagar mi salario y el material necesario para realizar esta tesis. En particular agradezco al Govern de les Illes Balears por concederme una beca FPI financiando mi sueldo durante 4 años. Agradezco a la Universitat de les Illes Balears que ha subvencionado parte mis estancias y congresos ayudado en mi formación Gracias sinceramente a mis directores Fiona Tomas y Jorge Terrados. Gracias por confiar en mí y saber guiarme dándome libertad creativa y ayudándome en cada uno de los pasos de este aprendizaje. Gracias por aceptar los empachos de correcciones a los que os he sometido estos últimos meses y cada una de vuestras ideas y comentarios. Te agradezco Fiona que me introdujeras en el mundo de las fanerógamas marinas antes de empezar esta tesis contagiándome tu pasión por la investigación y a afrontar las adversidades. Por los madrugones y las interminables horas de trabajo de campo compartidas, por tu motivación y por la ilusión que has puesto en este proyecto, gracias por elegirme como tu primera doctoranda. He sido muy afortunada con mis dos directores de tesis, profesionales, atentos a los detalles y que siempre han tenido su puerta o Skype abierto para mí ante cualquier necesidad. Creo que teneros a los dos como directores ha sido un equilibrio perfecto, muchas gracias. Estos más de 4 años en IMEDEA me dejan muy buenos recuerdos y amistades a las que estoy agradecida por hacer de mi paso por la isla algo inolvidable. Grandes compañeros veteranos del despacho que ahora estáis repartidos por el mundo: Isma, el único no científico del despacho; Inesius, la única no-física (junto conmigo) del despacho; Barbara, mi compi de mesa siempre con palabras de ánimo y buena energía; Adri-ant, el panarra number 1; Juanman, ahora el veterano del despacho y último y desde luego no menos importante Romain por todo lo compartido que siempre recordaré con una sonrisa. Compañeros de IMEDEA que se han ido como Morten y Ceci y los todavía resilientes; Ana, Marina, Fra, Johnna, Paula, Marc, Albert, Jaume, Merit y Carlos. Ha sido un placer poder compartir con vosotros mis inicios en la isla y en IMEDEA. A los compis que han ido llegando aumentando y enriqueciendo la plantilla, Dani, Laura, Andrea, Julia, Eva, Guillem, Alex Guillermo, Edu, Vero y tantos otros, y a los cañitas, a todos, gracias por ser mi taxi a IMEDEA o a Palma en tantas ocasiones y por hacerme perder tiempo de tesis tentándome con excursiones, barbacoas y los variados eventos sociales , habéis hecho mi etapa de tesis inolvidable. vii A todos los compañeros que hacen que IMEDEA tenga tan buen ambiente, los miembros del Green-team, los del grupo de peces, los microbiólogos… A Verioskis, Evinchis y Catunis un placer haber compartido tecitos, desayunos, playas, cenitas y confidencias, seguiremos por Skype whastapp o en hologramas de aquí a unos años. A los compis que junto conmigo están pariendo sus tesis Jaume, Noe, Veriozka, Merit y Carlos por el apoyo mutuo ya sea en forma calórica (gracias Merit) o burocrática. Quiero agradecer a todos mis coautores, a mis compañeras del grupo (Las Lauras e Inés) grandes científicas! y especialmente a toda la gente que de forma voluntaria me ha apoyado en trabajo de campo o laboratorio en algún momento de esta tesis: Diego, Elisenda, Samer, Jordi, Carlitos, Ana y mi Juls!. Gracias por vuestra ayuda desinteresada, sé que en ciertos momentos sin vosotros el trabajo hubiese sido prácticamente inabarcable. Many thanks to Oregon State University for accepting me as an Intenational student, especially to the Department of Fisheries and Wildlife. Many thanks to Jen and Jer, the best co- workers, super organized and fun. It has been a pleasure to share with you three months of my thesis and discover such nice sunrises during fieldwork and such tasty doughnuts at Hatfield. Muchas gracias al resto de compis en Corvallis con quienes disfruté mi estancia. Agradezco a la Fundación del Banco Santander que subvencionó mi estancia en la Universidad Autónoma de México en la Unidad de Sistemas Arrecifales. Agradezco muy especialmente a mi tutora Susana Enríquez que me acogió preocupándose porque mi aprendizaje durante la estancia fuese máximo y me cuido durante toda mi estancia en Puerto Morelos sobrepasando sus funciones como tutora. Muchas gracias por todo. Gracias a mi profe Román por su paciencia y sus explicaciones sobre el milagro y la complejidad de la fotosíntesis. Gracias a todos los compis de allá que hicieron mi estancia inolvidable y gracias Carlos y Merit y sus familias por acogerme y mostrarme la belleza de México. Gracias Vincent entre tantísimas cosas por tu ayuda en esta tesis y en conservar mi salud mental hasta el final. Y de postre, lo mejor, muchas gracias a mi familia. A mi tía, a mi hermana, a mi sobrino, sois lo más grande y mi energía vital. A mi padre que me inculcó la pasión por comprender y observar la naturaleza. A mi madre, una luchadora, gracias a cada uno de tus consejos he llegado hasta aquí. viii List of Manuscripts included in the Doctoral Thesis: Hernán G., Ortega M.J., Henderson J., Alos J., Boyer K., Cimon S., Combes, V., Cusson M., Dennert A., Hereu C., Hessing-Lewis M., Hovel K., Jorgensen P., Kiriakopolos S., Kollars N., O’Connor M., Olsen, J., Reynols PL., Ruesink J., Terrados J., Voigt E. and F. Tomas. Latitudinal patterns of productivity and response to herbivory in a marine habitat-forming species (in preparation; Chapter 1) Hernán G., Castejón, I., Terrados J, Tomas F. Resource availability and herbivory damage do not interact to modulate plant defense strategies (in preparation; Chapter 2). Hernán G., Ramajo L., Basso L., Delgado A., Terrados J., Duarte C.M., Tomas F. (2016) Seagrass (Posidonia oceanica) seedlings in a high-CO2 world: from physiology to herbivory. Scientific Reports, 6, 38017. (Chapter 3) Hernán G, Ortega, M.J., Gándara, A.M., Castejón, I., Terrados J, Tomas F. (2017) Future warmer seas: increased stress and susceptibility to grazing in seedlings of a marine habitat- forming species. Global Change Biology, 1-14 (Chapter 4). ix x Contents ABSTRACT xiii RESUMEN xv RESUM xvii List of abbreviations and acronyms xix List of Tables xxi List of Figures xxiii 1 Introduction and Objectives 1 1.1 Herbivory: a key process regulating plant communities 1 1.1.1 Herbivory in seagrasses 2 1.2 Plant-herbivore interactions 3 1.2.1 Plant defense strategies 3 1.2.2 Plant defense theories 9 1.2.3 Effects of environmental factors in plant defenses 13 1.3 Objectives 16 2 General Methodology 19 2.1 Experimental approaches 19 2.1.1 Field experiments 19 2.1.2 Mesocosm experiments 20 2.2 Plant analyses 21 2.3 Herbivore feeding experiments 23 2.3.1 Field experiments 24 2.3.2 Laboratory experiments 25 3 Results 27 3.1 Chapter 1: Latitudinal patterns of response to herbivory in a marine habitat-forming species 27 3.1.1 Abstract 27 3.1.2 Introduction 28 3.1.3 Materials and methods 31 3.1.4 Results 36 3.1.5 Discussion 43 3.2 Chapter 2: Resource availability and herbivory damage do not interact to modulate plant defense strategies 49 3.2.1 Abstract 49 3.2.2 Introduction 50 3.2.3 Materials and methods 53 xi 3.2.4 Results 58 3.2.5 Discussion 65 3.3 Chapter 3: Seagrass (Posidonia oceanica) seedlings in a high-CO2 world: from physiology to herbivory. 69 3.3.1 Abstract 69 3.3.2 Introduction 69 3.3.3 Materials and Methods 72 3.3.4 Results 76 3.3.5 Discussion 84 3.4 Chapter 4: Future warmer seas: increased stress and susceptibility to grazing in seedlings of a marine habitat-forming species 89 3.4.1 Abstract 89 3.4.2 Introduction 90 3.4.3 Materials and methods 92 3.4.4 Results 97 3.4.5 Discussion 104 4 General Discussion 109 4.1 Resource availability and plant defenses against herbivory 109 4.2 Human-mediated environmental changes and plant defenses against herbivory 111 4.3 Knowledge gaps and further research 112 5 Conclusions 117 6 References 119 7 Appendix 164 7.1 Supplementary Material Chapter 1: Latitudinal patterns of response to herbivory in a marine habitat-forming species 164 7.2 Supplementary Material Chapter 2: Resource availability and herbivory damage do not interact to modulate plant defense strategies 170 7.3 Supplementary Material Chapter 3: Seagrass (Posidonia oceanica) seedlings in a high- CO2 world: from physiology to herbivory 172 7.4 Supplementary Material Chapter 4: Future warmer seas: increased stress and susceptibility to grazing in seedlings of a marine habitat-forming species 174 xii ABSTRACT Herbivory is a key ecological process that regulates the composition and structure of plant communities and determines the energy transferred from primary producers to upper trophic levels. Plants have evolved a suite of defense strategies to avoid or resist herbivory. Tolerance strategies reduce the impact of herbivory in plant fitness (e.g., increased belowground reserves), and resistance strategies reduce preference or performance of the herbivore (e.g., low nutritional quality, high fiber content).
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