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Diapositiva 1 IMPLICACIONES DE LA CAPACIDAD DE 2011 MOVIMIENTO EN LAS ARAÑAS (ARANEAE) PARA LA EVOLUCIÓN ADAPTATIVA DEL TAMAÑO OCTORAL Y LA FORMA SEGÚN EL SEXO, HÁBITAT D Y MODO DE VIDA ESIS T ÁRQUEZ M 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 ORCOBADO C UADALUPE M. GUADALUPE CORCOBADO MÁRQUEZ TESIS DOCTORAL 2011 M. G Implicaciones de la capacidad de movimiento en las arañas (Araneae) para la evolución adaptativa del tamaño y la forma según el sexo, hábitat y modo de vida TESIS DOCTORAL M. Guadalupe Corcobado Márquez Granada, 2011 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 M. Guadalupe Corcobado Márquez 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] Los dibujos de la portada son obra de Mercedes Corcobado Márquez La imagen de la página 77 ha sido cedida por Dennis Kuntel. Derechos de autor: Dennis Kunkel Microscopy, Inc. (www.denniskunkel.com). Las fotos de las páginas 53, 97, 163 y la contraportada han sido tomadas y cedidas por Eva de Mas. Este trabajo ha sido financiado por una beca de Formación de Personal Investigador (FPI, ref: BES-2005-9234) y dos proyectos del Ministerio de Ciencia e Innovación/FEDER (CGL2004-03153 y CGL2007-60520) Implicaciones de la capacidad de movimiento en las arañas (Araneae) para la evolución adaptativa del tamaño y la forma según el sexo, hábitat y modo de vida Memoria presentada por M. Guadalupe Corcobado Márquez para optar al Grado de Doctora por la Universidad de Granada La Doctoranda M. Guadalupe Corcobado Márquez Granada, mayo2011 Dr. Jordi Moya-Laraño, Científico Titular de la Estación Experimental de Zonas Áridas-CSIC y el Dr. Miguel Ángel Rodríguez-Gironés Arbolí, Científico Titular de la Estación Experimental de Zonas Áridas-CSIC CERTIFICAN Que los trabajos de investigación realizados en la Memoria de Tesis Doctoral: “Implicaciones de la capacidad de movimiento en las arañas (Araneae) para la evolución adaptativa del tamaño y la forma según el sexo, hábitat y modo de vida”, son aptos para ser presentados por la Lda. M. Guadalupe Corcobado Márquez ante el Tribunal que en su día se designe, para aspirar al Grado de Doctora en Ciencias Biológicas por la Universidad de Granada. Y para que así conste, en cumplimiento de las disposiciones vigentes extendemos el presente certificado a 16 de mayo de 2011 en Almería. VºBº Director VºBº Director Dr. Jordi Moya-Laraño Dr. Miguel Ángel Rodríguez-Gironés Arbolí A mis padres Genaro y Carmen, y a mis tres hermanas Mari, Conchi y Merce, como recompensa por todo el tiempo que esta tesis les ha robado. Desde hace tiempo, intento cambiar cantidad por calidad en el tiempo que compartimos A Miguel, por su infinita paciencia, y por poner su energía cuando la mía se agotaba La ignorancia afirma o niega rotundamente; la ciencia duda. Voltaire (1694-1778) Filósofo y escritor francés. Índice Índice ÍNDICE............................................................................................................................................................1 RESUMEN......................................................................................................................................................3 INTRODUCCIÓN GENERAL.................................................................................................................11 Introducción................................................................................................................................13 Las arañas como sistema de estudio.................................................................................25 Justificación y Objetivos.........................................................................................................28 Bibliografía...................................................................................................................................36 MANSUCRITOS..........................................................................................................................................51 Manuscrito I: Morphological evolution of spiders predicted by pendulum mechanics.....................................................................................................53 Manuscrito II: Silk elasticity as a potential constraint on spider body size...77 Manuscrito III: Introducing the refined gravity hypothesis of extreme sexual size dimorphism..........................................................................97 Manuscrito IV: Sociality level explains dispersal ability in spiders.................135 Manuscrito V: Sprint vs cruise speed: Implications for natural and sexual selection......................................................................................................163 Manuscrito VI: Scramble competition by male long-distance mobility in a burrowing wolf spider..........................................................................187 DISCUSIÓN INTEGRADORA..............................................................................................................221 Resultados y Discusión........................................................................................................223 Bibliografía................................................................................................................................235 CONCLUSIONES / CONCLUSIONS..................................................................................................241 Conclusiones.............................................................................................................................243 Conclusions...............................................................................................................................245 AGRADECIMIENTOS............................................................................................................................247 1 2 Resumen RESUMEN 3 Resumen 4 Resumen Resumen La habilidad de un organismo para moverse desempeña un papel fundamental en muchas de las actividades que debe realizar para cumplir sus funciones vitales. De este modo, podemos decir que la capacidad de movimiento puede incrementar la eficacia biológica de un individuo vía selección natural, aumentando la supervivencia y/o fecundidad de los organismos, o vía selección sexual, incrementando su éxito a la hora de obtener pareja. Cabría esperar, por tanto, que a lo largo de la evolución se hayan seleccionado tamaños y formas adaptadas a maximizar la capacidad de movimiento dependiendo del modo de vida de los organismos. Así, en taxones donde ambos sexos adoptan distinto modo de vida una vez alcanzado el estado adulto, como es el caso de muchas especies de arañas, la importancia de desarrollar morfologías adaptadas al movimiento va a depender del rol sexual y va a variar entre machos y hembras, originando patrones de dimorfismo sexual. En particular, los desplazamientos suelen aumentar la tasa de depredación. Como los costes del desplazamiento van a condicionar la elección entre dispersión y filopatria a la que los individuos deben enfrentarse, se deduce que la capacidad de locomoción de un individuo debe influir en su tendencia a la dispersión. La mayoría de los estudios muestran una estrecha relación entre tamaño y/o forma y capacidad de locomoción, aunque la mayoría de los trabajos han utilizado la velocidad de escape como índice de capacidad de locomoción, mientras que otros aspectos tales como velocidad de crucero apenas han sido investigados. Además, existe un número considerable de trabajos que han analizado cómo la capacidad de movimiento se relaciona con la supervivencia (selección natural), mientras que el papel de las habilidades locomotoras de los organismos sobre su éxito reproductivo (selección sexual) se ha estudiado mucho más raramente. Por último, existen modos de locomoción exclusivos de algunos grupos taxonómicos, como es el puenteo en las arañas –un tipo de locomoción aérea mediado por seda muy común en arañas que viven en la vegetación–, que han sido totalmente obviados en los estudio de ecología 5 Resumen evolutiva. Todo ello a pesar de estar probablemente muy extendidos y de desempeñar por tanto un papel fundamental en la evolución de los rasgos de las especies que los emplean. A la vista de los antecedentes expuestos anteriormente, a lo largo de esta tesis se han estudiado algunas de las implicaciones ecológicas y/o evolutivas de la capacidad de movimiento, relacionadas tanto con la selección natural como con la selección sexual. En particular, en este trabajo se evalúan implicaciones de la capacidad de movimiento en la selección de las morfologías más adecuadas dependientes del tipo de hábitat o modo de vida, la evolución del dimorfismo sexual en el tamaño (SSD, del inglés Sexual Size Dimorphism), la selección sexual por orden de llegada y la evolución de la sociabilidad en arañas. En el manuscrito I de esta tesis, a través de un estudio comparativo que incluyó una muestra de amplia diversidad filogenética y un amplio rango de tamaños, se estudió si la forma de las arañas podría haber evolucionado de acuerdo a las predicciones de la mecánica pendular, teniendo en cuenta que, desde un punto de vista biomecánico, el puenteo
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