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Thesis Ovv 02 Algunas consecuencias ecológicas y evolutivas del movimiento animal para las interacciones bióticas TESIS DOCTORAL Oriol Verdeny Vilalta Granada, 2013 Departamento de Ecología Funcional y Evolutiva Departamento de Ecología Estación Experimental de Zonas Áridas Facultad de Ciencias CONSEJO SUPERIOR DE INVESTIGACIONES CIENTÍFICAS UNIVERSIDAD DE GRANADA Oriol Verdeny Vilalta Estación Experimental de Zonas Áridas Consejo Superior de Investigaciones Científicas Carretera de Sacramento, s/n, Cañada de San Urbano E‐04120, Almería, Spain [email protected] Portada diseñada y realizada por Assumpta Vilalta Guart Este trabaJo ha sido financiado por una beca de Formación de Personal Investigador (FPI, ref: BES‐2008‐004515), el proyecto del Ministerio de Ciencia e Innovación/FEDER (CGL2010‐18602) y del Ministerio de Agricultura, Alimentación y Medio Ambiente/FEDER (OAPN20/2008). Algunas consecuencias ecológicas y evolutivas del movimiento animal para las interacciones bióticas Memoria presentada por Oriol Verdeny Vilalta para optar al Grado de Doctor por la Universidad de Granada El Doctorando Oriol Verdeny Vilalta Granada, marzo de 2013 Dr. Jordi Moya Laraño, Científico Titular de la Estación Experimental de Zonas Áridas‐CSIC CERTIFICA Que los trabajos de investigación realizados en la Memoria de Tesis Doctoral: “Algunas consecuencias ecológicas y evolutivas del movimiento animal para las interacciones bióticas”, son aptos para ser presentados por el Ldo. Oriol Verdeny Vilalta ante el Tribunal que en su día se designe, para aspirar al Grado de Doctor en Ciencias Biológicas por la Universidad de Granada. Y para que así conste, en cumplimiento de las disposiciones vigentes extendemos el presente certificado a 1 de marzo de 2013 en Almería. VºBº Director Dr. Jordi Moya Laraño El doctorando Oriol Verdeny Vilalta y el director de la tesis Jordi Moya Laraño garantizamos, al firmar esta tesis doctoral, que el trabajo ha sido realizado por el doctorando bajo la dirección de los directores de la tesis y hasta donde nuestro conocimiento alcanza, en la realización del trabajo, se han respetado los derechos de otros autores a ser citados, cuando se han utilizado sus resultados o publicaciones. En Almería a 1 de marzo de 2013 Director/es de la Tesis Doctorando Fdo.: Fdo.: Als meus pares, al meu germà, i a la meva petita Ana Índice Índice .............................................................................................................................. 1 Resumen ......................................................................................................................... 3 Introducción general ............................................................................................... 7 Introducción ....................................................................................................... 9 Justificación y objetivos .................................................................................... 17 Bibliografía .......................................................................................................... 21 Chapter 1 ‐ Seeking for water while avoiding predators: moisture gradients can affect centipede‐springtail predator‐prey interactions ........................................................................................... 29 Chapter 2 ‐ Spatial heterogeneity in the availability of water and basal resources determines spatial aggregations and possibly indirect effects in soil macrofauna ................................................ 61 Chapter 3 ‐ Foraging paths in complex environments: relative roles of architectural complexity and intrinsic stimulus strength on flies foraging for fruits ....................................................................... 113 Chapter 4 ‐ Food web allometric constraints, foraging theory and foraging mode explain the evolution of egg size ...................... 139 Chapter 5 ‐ Climate change and eco‐evolutionary dynamics in food webs ......................................................................................................... 177 Discusión integradora .............................................................................................. 275 Discusión .............................................................................................................. 276 Bibliografía .......................................................................................................... 282 Conclusiones ................................................................................................................ 287 Agradecimientos ........................................................................................................ 291 1 2 Resumen 3 4 Resumen En el mundo animal, la evolución ha dotado a distintas especies con diversas formas de dispersión, lo que les permite satisfacer sus necesidades mas básicas, como por ejemplo encontrar recursos, escapar de los depredadores o aparearse. Además, distintos patrones de movimiento de los animales conllevarán una redistribución de las poblaciones en el espacio, lo que afectará a las tasas de encuentro y a las posibles interacciones bióticas que se pueden establecer. Por este motivo las tasas de encuentro están, en última instancia, relacionadas con la eficacia biológica de los animales. Dado que las interacciones están necesariamente localizadas en el espacio, el conocimiento del movimiento de los animales y sus tasas de encuentro pueden mejorar nuestro conocimiento de las interacciones bióticas así como qué condiciones llevan a la estabilidad de las poblaciones que interaccionan. Además, la movilidad de los animales juega un papel importante en la evolución de determinados rasgos, ya que determinados fenotipos pueden determinar la eficacia de dispersión y las tasas de encuentro, así como el potencial de interacción. El objetivo de la presente tesis ha sido estudiar el papel que tienen distintos factores abióticos ‐haciendo hincapié sobre aquellos factores que están siendo afectados por el cambio climático‐ sobre las tasas de encuentro de invertebrados, así como algunas de las consecuencias ecológicas y evolutivas que pueden tener dichas tasas de encuentro. Para ello en esta tesis se desarrollan cinco objetivos específicos: (1) estudiar la importancia relativa de la distribución y disponibilidad hídrica en las tasas de encuentro y de depredación de un depredador y una presa típicos de bosques caducifolios, (2) determinar, en una red trófica de descomponedores de bosques caducifolios, los efectos de la disponibilidad de agua y de recursos basales en los patrones de agregaciones espaciales, así como sus consecuencias en las tasas de depredación, (3) investigar los roles de la complejidad estructural de la vegetación y de la fuerza de atracción de un recurso en la geometría de los movimientos del díptero Rhagoletis pomonella, (4) investigar las consecuencias de la movilidad en la evolución del tamaño y número de los huevos en depredadores intragremiales (5) estudiar las consecuencias de un incremento en la temperatura sobre las dinámicas eco‐evolutivas en interacciones tri‐tróficas. Los resultados de esta 5 tesis muestran la importancia de la disponibilidad hídrica y la productividad basal a la que va normalmente asociada el agua en suelos (i.e. hongos) en determinar las tasas de encuentro entre animales, la estructura de la red trófica, y los posibles efectos indirectos que los depredadores ejercen sobre las presas. También hemos visto cómo la estructura de la vegetación determina los movimientos de un insecto que forrajea a la vez que modifica el radio de percepción del insecto. Finalmente los resultados nos han mostrado que la movilidad de los animales juega un importante papel en la evolución de ciertos rasgos fenotípicos como son el tamaño y el número de los huevos, así como en las dinámicas eco‐evolutivas de una red tri‐trófica. 6 Introducción general 7 8 Introducción En el mundo animal, la evolución ha dotado a distintas especies con diversas formas de dispersión para satisfacer sus necesidades mas básicas, como por ejemplo encontrar recursos, escapar de los depredadores o aparearse (Bell 1990, Moya‐Laraño et al. 2002, Bartumeus et al. 2003, Nilsson and Bengtsson 2004). La dispersión de los animales también permite la recolonización de ciertas áreas en las que subpoblaciones no se encontraban presentes o se habían extinguido, lo que a nivel global afecta a las dinámicas de poblaciones e incrementa la persistencia de las poblaciones (Bascompte and Solé 1995, Thompson et al. 2012), especialmente en poblaciones que se encuentran en hábitats fragmentados (Zollner and Lima 1999). Además, la movilidad de las especies también afecta al flujo de material genético entre distintas áreas y por lo tanto a la diversidad genética y a su mantenimiento (Lundberg and Moberg 2003, Brown et al. 2008, Amos et al. 2012), y es un factor clave que afecta a las invasiones biológicas y a la propagación de enfermedades o plagas (Fevre et al. 2006, Nathan 2008, Stoddard et al. 2013). La capacidad de movimiento es una característica fundamental que determina las tasas de encuentro de los organismos y sus interacciones bióticas potenciales. Por este motivo las tasas de encuentro también están en última instancia relacionadas con la eficacia biológica de los animales. Por lo tanto, distintos patrones de movimiento como son la velocidad de los animales, la sinuosidad de los recorridos efectuados por los mismos o la atracción hacia ciertas áreas, conllevará redistribuciones de las poblaciones que afectarán
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