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Caulerpa Taxifolia (Chlorphyta: Caulerpaceae) and Established

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Caulerpa Taxifolia (Chlorphyta: Caulerpaceae) and Established Tesis Doctoral Ecología de Caulerpales: Fauna y Biomarcadores Autor: Antonio Box Centeno Directora: Dra. Salud Deudero Company Programa doctorado Ciencias Marinas (Instituto Mediterráneo de Estudios Avanzados) Universidad Islas Baleares 22-Julio-2008 TESIS DOCTORAL Ecología de Caulerpales: Fauna y Biomarcadores Tesis doctoral presentada por Antonio Box Centeno para optar al titulo de doctor del programa en Ciencias Marinas de la Universidad de las Islas Baleares, bajo la dirección de la Dra. Salud Deudero Company Antonio Box Centeno Palma, 30 de Mayo de 2008 Directora de la Tesis Doctoral El interesado - 3 - Índice Índice de la presente tesis doctoral ÍNDICE …………………………………………………………………………………. 5 AGRADECIMIENTOS ………………………………………………………………… 7 ABREVIACIONES …………………………………………………………………….. 9 CAPÍTULO 1: INTRODUCCIÓN GENERAL ………………………………………… 11 1.1 Introducción general ………………………………………………… 13 1.2 Especies invasoras ……………………………………………………. 15 1.2.1 Definición de especie invasora y problemática ……. 15 1.2.2 Especies invasoras en el Mediterráneo y Baleares …. 21 1.3 Praderas de Posidonia oceanica , Caulerpales e invertebrados ………………………………………………………………. 23 1.4 Caulerpenina y biomarcadores de estrés oxidativo ………….. 29 1.4.1 Metabolitos secundarios, Caulerpenina ……………... 29 1.4.2 Estrés oxidativo …………………………………………….. 29 CAPÍTULO 2 : CAMBIOS EN LAS COMUNIDADES DE MOLUSCOS …………… 35 2.1 Introducción al capítulo …………………………………………….. 37 2.2 A mollusc community associated with invasive Caulerpa racemosa in the Western Mediterranean shallow seagrass beds . 39 2.3 How different is the mollusc community between invasive Caulerpa taxifolia (Chlorphyta: Caulerpaceae) and established Caulerpa prolifera in shallow seagrass beds of the Western Mediterranean ? …………………………………………………………. 63 2.4 Resumen de las comunidades de moluscos …………............. 87 CAPÍTULO 3 : CAMBIOS EN LAS COMUNIDADES DE DECÁPODOS ............... 91 3.1 Introducción al capítulo .............................................................. 93 3.2 Changes in decapod crustacean faunistic composition associated with the seagrass Posidonia oceanica (Delile) induced by the presence of invasive Caulerpa species ................ 95 3.3 Resumen de las comunidades de decápodos ......................... 123 CAPÍTULO 4 : CAMBIOS EN LAS COMUNIDADES DE POLIQUETOS ................ 125 4.1 Introducción al capítulo ............................................................... 127 4.2 Seagrass polychaete assemblages changes after invasion by Caulerpa racemosa var cylindracea : community structure and - 5 - Índice trophic guilds ........................................................................................ 129 4.3 Community structure and trophic guilds: Differences in seagrass polychaete asemblage between invasive Caulerpa taxifolia and established C. prolifera .................................................. 155 4.4 Resumen de las comunidades de poliquetos ............................ 183 CAPÍTULO 5 CAMBIOS EN LAS COMUNIDADES DE INVERTEBRADOS. NUEVAS ESPECIES Y CITAS ................................................................................ 187 5.1 Introducción al capítulo ............................................................... 189 5.2 Changes in the faunistic communities associated with the presence of green algae Caulerpa .................................................. 191 5.3 A new species Nuuanu (Amphipoda: Gammaridea: Melitidae) from shallow sandy bottoms of the Balearic Islands (W Mediterranean) .................................................................................... 207 5.4 Ocurrence of Automate branchialis (Holthuis & Gottlieb, 1959)(Decapoda, Alpheidae) in the Balearic Islands (Western Mediterranean) .................................................................................... 225 5.5 Resumen de cambios en las comunidades de inverebrados . 233 CAPÍTULO 6 : CAULERPENINA Y DEFENSAS ANTIOXIDANTES ......................... 235 6.1 Introducción al capítulo ............................................................... 237 6.2 Seasonality of caulerpenyne contents in the native Caulerpa prolifera and the invasive Caulerpa taxifolia and Caulerpa racemosa var cylindracea in the Western Mediterranean ............ 241 6.3 Enzimatic antioxidant response of a labrid fish ( Coris julis ) liver to environmental caulerpenyne ........................................................ 261 6.4 Reciprocal effects of caulerpenyne and intense herbivorism on the antioxidant response of Bittium reticulatum and Caulerpa taxifolia ................................................................................ 279 6.5 Antioxidant response and caulerpenyne production of Caulerpa taxifolia (Vahl) C. Agardh epiphyted by the invasive algae Lophocladia lallemandii (Montagne) .................................. 297 6.6 Resumen de caulerpenina y defensas antioxidantes ............... 313 CAPÍTULO 7 RECAPITULACIÓN DE RESULTADOS .............................................. 319 REFERENCIAS BIBLIOGRÁFICAS ........................................................................ 331 - 6 - Agradecimientos Agradecimientos Nunca pensé que hacer una tesis doctoral implicase tanto trabajo, horas de lupa, tiempo de ordenador, ratos delante de un espectofotómetro o el HPLC, horas y más horas. En el momento de presentar mi tesis doctoral no me puedo olvidar de todas aquellas personas que han puesto su tiempo a mi disposición, enseñándome y explicándome nuevos conceptos, ayudándome y dándome los medios para poder desarrollar mis ideas e inquietudes. En primer lugar quiero agradecer a la Dra Salud Deudero el ofrecerme la posibilidad de trabajar dentro de su equipo y siempre buscar la forma de que pudiese continuar adelante con mi tesis. Además quiero agradecer su forma de incentivar y permitir el desarrollo de mis ideas en la presente tesis doctoral. Agradecer la ayuda del Laboratorio de Biología Marina de la Universidad de las Islas Baleares y todas las personas que durante más o menos tiempo han desarrollado prácticas o se han iniciado en la investigación, ya que muchas de ellas han colaborado activamente en la separación de las muestras, su procesado y salidas de campo. He de dar gracias especialmente a Piluca Sarrirera, Andreu Blanco, Toni Frau, Pep Alòs, y Cristina Alonso entre otras muchas persona por su tiempo dedicación y ayuda. A la Catedrática de la Universidad de las Islas Baleares Isabel Moreno toda su paciencia, consejos en momentos difíciles y ser la persona que me permtió entrar dentro del Laboratorio de Biología Marina. La presente tesis doctoral está enmarcada dentro de dos proyectos de Plan Nacional, el proyecto CAULEXPAN (REN-2002-00701/MAR), MACROALGAS INVASORAS (CTM2005-01434/MAR). Ambos proyectos han suministrado los fondos necesarios para el desarrollo de la presente investigación. Por ello quiero agradecer a la Dr. Nuria Marbà (IP Caulexpan) como el Dr. Jorge Terrados (IP Macroalgas invasoras) el permitirme trabajar, aprender y colaborar activamente con ellos. Al Dr. Antoni Sureda su colaboración directa conmigo a la hora de desarrollar y poner a punto las metodologías necesarias para evaluar la concentración de caulerpenina en las tres caulerpales y en la realización de los trabajos ecotoxicológicos para interpretar el efecto de la caulerpenina sobre invertebrados, vertebrados y algas. Al Dr. Daniel Martín, investigador de CEAB gracias al cual aprendí a clasificar los poliquetos, me proporcionó la bibliografía necesaria, me dió unas clases intensivas y - 7 - Agradecimientos particulares de clasificación de poliquetos y que continua ayudándome en la escritura de nuevas publicaciones. Al Dr. Pere Abellò, investigador de ICM por su ayuda a la hora de clasificar todos los decápodos que encontré en las muestras de Caulerpa y Posidonia , me proporcionó bibliografía y continua ayudándome durante la escritura de nuevas publicaciones. Al Dr. Antoni Pons, profesor titular de la UIB, sus ideas, su tiempo, su interés, las facilidades para trabajar con los miembros de su equipo y la accesibilidad y facilidades para utilizar toda la maquinaria necesaria para poder abarcar todos los temas bioquímicos. Al Dr. Damià Jaume, investigador del IMEDEA el cual me ha puesto en contacto con coleguas suyos para la clasificación de otros grupos taxonómicos y me ha ayudado a clasificar tanto anfípodos como isópodos. Al Dr. Guillem Pons, profesor de la UIB experto en moluscos, su ayuda a la hora de clasificar los “bichos” de las muestras. Agradecerle también la oportunidad de realizar mi primera conferencia de invertebrados en Caulerpa . Al Dr. Phillipe Amade por venir hasta Mallorca y poner a punto la técnica de cuantificación de caulerpenina además deproporcionarnos caulerpenina pura con la cual pudimos establecer la recta patrón. Agradercer especialmente al Dr. Guillem Mateu-Vicens, Silvia Tejada, Felip Cirer y Teresa Marcos por su paciente lectura de los manuscritos y de la tesis, sus comentarios y aportaciones. Agradecer a mis padres todo su apoyo incondicional durante mis años de doctorando, sus ánimos y ayuda en momentos duros y de dudas. Quiero agradecer también a mi futura esposa, Paqui, el apoyo y ánimos que me ha dado en los momentos de desanimo y que ha sacrificado muchas cosas viniendo a vivir conmigo. Quiero acordarme también de todas aquellas personas que han compartido parte de su vida conmigo haciendola más feliz y que ahora no están, muchas gracias a todos por todo lo que me habeis aportado. Muchas gracias a todos - 8 - Abreviaciones Abreviaciones usadas ANOSIM Analisis of Similarities,
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