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Efectos De La Contaminación Por Fertilizantes Sobre Pelophylax Perezi (Seoane, 1885)

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Efectos De La Contaminación Por Fertilizantes Sobre Pelophylax Perezi (Seoane, 1885) Universidad de Murcia Facultad de Biología Departamento de Zoología y Antropología Física Aspectos relevantes en la conservación de anfibios en la Región de Murcia: efectos de la contaminación por fertilizantes sobre Pelophylax perezi (Seoane, 1885) Memoria presentada para optar al grado de Doctor en Biología por el Licenciado en Biología Andrés Egea Serrano Directores: Dra. Mar Torralva Forero (Universidad de Murcia) Dr. Miguel Tejedo Madueño (Estación Biológica de Doñana-CSIC) A mi familia y, muy especialmente, a mis padres, Paco y María With magic, you can turn a frog into a prince. With science, you can turn a frog into a Ph.D. and you still have the frog you started with. Terry Pratchett, Ian Stewart & Jack Cohen. 2002. The Science of Discworld . Ebury Press, Londres. ÍNDICE Agradecimientos i Resumen general (versión inglesa) v Resumen general (versión española) xiii Estructura de la presente Tesis Doctoral xxi BLOQUE I. INTRODUCCIÓN 1 Capítulo 1. Introducción y objetivos 3 Capítulo 2. Área de estudio, descripción de la especie estudiada y sinopsis metodológica 65 BLOQUE II. ANÁLISIS DE LOS EFECTOS DE LOS COMPUESTOS NITROGENADOS EN PELOPHYLAX PEREZI EN EXPERIMENTOS DE LABORATORIO 89 Capítulo 3. Estimación de las concentraciones letales medias de tres compuestos nitrogenados para larvas de rana común, Pelophylax perezi (Seoane, 1885) 91 Capítulo 4. Divergencia poblacional en el impacto de tres compuestos nitrogenados y su combinación sobre larvas de la rana Pelophylax perezi (Seoane, 1885) 111 Capítulo 5. Estimación del impacto de tres compuestos nitrogenados y su combinación sobre el nivel de inactividad y el uso del hábitat de larvas de Pelophylax perezi (Seoane, 1885) 145 Capítulo 6. Análisis de la evitación de los fertilizantes nitrogenados en la columna de agua por parte de juveniles de rana común, Pelophylax perezi (Seoane, 1885), en condiciones de laboratorio 171 Índice BLOQUE III. ANÁLISIS DE LOS EFECTOS DE LOS COMPUESTOS NITROGENADOS EN PELOPHYLAX PEREZI EN EXPERIMENTOS DE MESOCOSMOS Y CAMPO 189 Capítulo 7. Efectos de la contaminación por nitrógeno sobre la supervivencia, la morfología y el crecimiento de larvas de rana común, Pelophylax perezi (Seoane, 1885): una aproximación jerárquica a las condiciones naturales 191 Capítulo 8. Análisis de los efectos de la contaminación por nitrógeno sobre las habilidades natatorias de larvas de Pelophylax perezi (Seoane, 1885) a través de ensayos representativos de las condiciones naturales 241 BLOQUE IV. ANÁLISIS DE LOS EFECTOS DE LA CONTAMINACIÓN SOBRE LOS ANFIBIOS 285 Capítulo 9. ¿Están los anfibios realmente amenazados por los compuestos químicos? Una revisión meta-analítica 287 BLOQUE V. CONCLUSIONES GENERALES 353 Capítulo 10. Conclusiones (versión inglesa) 355 Capítulo 10. Conclusiones (versión española) 361 ÍNDICE Agradecimientos i General Abstract (English version) v Resumen general (versión española) xiii Estructura de la presente Tesis Doctoral xxi BLOQUE I. INTRODUCCIÓN 1 Capítulo 1. Introducción y objetivos 3 Capítulo 2. Área de estudio, descripción de la especie estudiada y sinopsis metodológica 65 BLOQUE II. ANÁLISIS DE LOS EFECTOS DE LOS COMPUESTOS NITROGENADOS EN PELOPHYLAX PEREZI EN EXPERIMENTOS DE LABORATORIO 89 Capítulo 3. Estimating mean letal concentrations of three nitrogenous compounds for the Iberian water frog, Pelophylax perezi (Seoane, 1885), larvae 91 Capítulo 4. Populational divergence in the impact of three nitrogenous compounds and their combination on larvae of the frog Pelophylax perezi (Seoane, 1885) 111 Capítulo 5. Examining the impact of three nitrogenous compounds and their combination on inactivity level and habitat use of larvae of Pelophylax perezi (Seoane, 1885) 145 Capítulo 6. Analysis of the avoidance of nitrogen fertilizers in the water column by juvenile Iberian water frog, Pelophylax perezi (Seoane, 1885), in laboratory conditions 171 Índice BLOQUE III. ANÁLISIS DE LOS EFECTOS DE LOS COMPUESTOS NITROGENADOS EN PELOPHYLAX PEREZI EN EXPERIMENTOS DE MESOCOSMOS Y CAMPO 189 Capítulo 7. Effects of nitrogenous pollution on survival, morphology and growth of larvae of the Iberian waterfrog, Pelophylax perezi (Seoane, 1885): a hierarchical approach to natural conditions 191 Capítulo 8. Analysis of the effects of nitrogenous pollution on swimming performance of larvae of Pelophylax perezi (Seoane, 1885) through assays representative of natural conditions 241 BLOQUE IV. ANÁLISIS DE LOS EFECTOS DE LA CONTAMINACIÓN SOBRE LOS ANFIBIOS 285 Capítulo 9. Are amphibians actually threatened by chemicals? A meta- analytic review 287 BLOQUE V. CONCLUSIONES GENERALES 353 Capítulo 10. Conclusions (English version) 355 Capítulo 10. Conclusiones (versión española) 361 AGRADECIMIENTOS La realización de un proyecto de investigación, con independencia de sus objetivos, es una tarea compleja, por gratificante que pueda revelarse. Los numerosos obstáculos, conspicuos o sutiles, que aparecen durante el desarrollo de una investigación sólo pueden ser salvados gracias a la contribución de un ejército de personas, y mi caso no es una excepción. Así, debo agradecer en primer lugar a mis directores, Miguel Tejedo y Mar Torralva, su apoyo logístico e, indudablemente, intelectual. Las discusiones sobre las cuestiones planteadas directamente en la presente Tesis Doctoral, así como sobre otras más indirectas, pero no por ello menos interesantes, han resultado enormemente estimulantes para mí. Sin duda alguna, durante este periplo predoctoral he aprendido mucho de ellos. Por otra parte, deseo mostrar a todos los miembros del Departamento de Zoología y Antropología Física de la Universidad de Murcia mi agradecimiento por su apoyo, ya que sin él los experimentos realizados no pudieron haber sido ejecutados. Asimismo, la Universidad de Murcia facilitó el espacio necesario para poder instalar los mesocosmos utilizados para la realización de algunos de los experimentos presentados en esta memoria. Como el aspecto pecuniario de la investigación no deja de ser asunto baladí, no quiero dejar de agradecer a la Caja de Ahorros del Mediterráneo su apoyo, ya que fui beneficiario de una de las becas predoctorales financiadas a través del programa de obras sociales establecido por esta entidad. Agradezco a Gema Parra, Enrique García-Muñoz, Juan Diego y Francisco José Guerrero todas las atenciones que tuvieron para conmigo durante mi estancia en su laboratorio en la Universidad de Jaén. Asimismo, estoy en deuda con Anssi Laurila, i Agradecimientos Katja Räsänen, Sandra Hangartner, Emma Dahl, Germán Orizaola, María Quintela y, especialmente, con Alex Richter-Boix por ampliar mis horizontes profesionales y personales durante mi estancia en la Universidad de Uppsala. Confío en que mi paso por sus laboratorios no haya supuesto grandes estropicios y que fruto de estas estancias se deriven futuras fructíferas colaboraciones. Deseo expresar a través de estas líneas mi agradecimiento a Geoffrey Smith, Anne Lise Mandrillon, Manu Ortiz-Santaliestra y Albert Montori por facilitarme datos inéditos o detalles de los ya publicados que enriquecieron la matriz de datos confeccionada para la realización de meta-análisis. Asimismo, agradezco a Domingo Campillo su apoyo al autorizarme a utilizar las instalaciones de la estación de tratamiento de agua potable de la Contraparada, perteneciente a la Empresa Municipal de Aguas y Saneamiento de Murcia, S.A., para determinar las concentraciones de iones nitrogenados en mis mesocosmos y localidades de campo. De modo especial agradezco a Isabel Hurtado que se tomara la molestia de compartir su tiempo conmigo e instruirme en la aplicación de técnicas fotoespectrométricas y cromatográficas. Otro grupo de personas cuya ayuda me resultó indispensable son la fish people , excelentes compañeros en el laboratorio y mejores amigos, a pesar de que mis repetidos intentos de corromperlos para que se iniciaran en el mundo de los anfibios se vieron recompensados con el más glorioso de los fracasos. Así, entre otros motivos, agradezco a David Verdiell su ayuda en el campo, y Ana Ruiz sus bocadillos de tortilla, intendencia con la que tan amablemente a veces tuvo a bien abastecerme. A ellos, y naturalmente también a Raquel Moreno, Asun Andreu y Antonio Lacunza, muchas gracias por tener la paciencia de escucharme en esos momentos de tribulación y desasosiego. ii Agradecimientos Naturalmente, y aunque a veces llegué a olvidarlo, también hay vida fuera del laboratorio. Así, agradezco a Antonio Bastida, a Jose Antonio de Maya y, de modo muy especial, a Pedro Miñano todas las charlas y aventuras que hemos tenido tras los anfibios (y lo que no son anfibios). Muchas gracias por todo lo que he aprendido y por todo lo que nos hemos reído. Asimismo, he contraído una profunda deuda con Lucrecia Acosta por su auxilio, ya que gracias a su conversación, epistolar o verbal, el autor de estas líneas pudo mantener la escasa cordura de la que siempre ha hecho gala. Por último, pero no menos importante, quiero agradecer a mi hermana y, muy especialmente, a mis padres su paciencia. Ellos han sido capaces de doctorarse summa cum laude en el difícil arte de soportar las excentricidades de un doctorando, amén de proporcionar su valiosa ayuda en campos de naturaleza mucho más prácticos. iii GENERAL ABSTRACT (English version) Current extinction rates due to the direct and indirect consequences of human intervention on natural systems are several orders of magnitude higher than those recorded throughout the historical evolution of the Earth. Consequently, minimizing the loss of global biodiversity
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