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Crustacea: Ostracoda) De Pozas Temporales Heterocypris bosniaca (Petkowski et al., 2000): Ecología y ontogenia de un ostrácodo (Crustacea: Ostracoda) de pozas temporales. ESIS OCTORAL T D Josep Antoni Aguilar Alberola Departament de Microbiologia i Ecologia Universitat de València Programa de doctorat en Biodiversitat i Biologia Evolutiva Heterocypris bosniaca (Petkowski et al., 2000): Ecología y ontogenia de un ostrácodo (Crustacea: Ostracoda) de pozas temporales. Tesis doctoral presentada por Josep Antoni Aguilar Alberola 2013 Dirigida por Francesc Mesquita Joanes Imagen de cubierta: Vista lateral de la fase eclosionadora de Heterocypris bosniaca. Más detalles en el capítulo V. Tesis titulada "Heterocypris bosniaca (Petkowski et al., 2000): Ecología y ontogenia de un ostrácodo (Crustacea: Ostracoda) de pozas temporales" presentada por JOSEP ANTONI AGUILAR ALBEROLA para optar al grado de Doctor en Ciencias Biológicas por la Universitat de València. Firmado: Josep Antoni Aguilar Alberola Tesis dirigida por el Doctor en Ciencias Biológicas por la Universitat de València, FRANCESC MESQUITA JOANES. Firmado: F. Mesquita i Joanes Profesor Titular de Ecología Universitat de València A Laura, Paco, i la meua família Resumen Los ostrácodos son un grupo de pequeños crustáceos con amplia distribución mundial, cuyo cuerpo está protegido por dos valvas laterales que suelen preservarse con facilidad en el sedimento. En el presente trabajo se muestra la primera cita del ostrácodo Heterocypris bosniaca Petkowski, Scharf y Keyser, 2000 para la Península Ibérica. Se trata de una especie de cipridoideo muy poco conocida que habita pozas de aguas temporales. Se descubrió el año 2000 en Bosnia y desde entonces solo se ha reportado su presencia en Israel (2004) y en Valencia (presente trabajo). En primer lugar se lleva a cabo una descripción del área de estudio. Tras conocer varias pozas con la presencia de la especie se realizó un seguimiento semanal para conocer el ciclo de vida y la dinámica poblacional. A continuación se estudia la supervivencia y otros aspectos ecológicos en respuesta a diferentes tratamientos (desecación, temperatura, contaminantes) controlados en el laboratorio para poder conocer mejor la ecología de las poblaciones naturales. Finalmente se presentan en detalle los cambios en la morfología externa a lo largo de la ontogenia y en los procesos de eclosión. Los datos obtenidos indican que H. bosniaca tiene una estrategia de vida adaptada a tolerar el elevado estrés de ambientes acuáticos temporales de ciclo irregular, donde se observa que puede completar varias generaciones en un año. Su ciclo vital está fuertemente influenciado por la temperatura: la velocidad de crecimiento aumenta con la temperatura, aunque no siempre cumplen la regla de alcanzar mayor tamaño corporal a temperaturas más bajas. La especie muestra una gran resistencia a los contaminantes comparado con otros crustáceos, lo cual puede ofrecerle ventajas en la colonización de nuevos ambientes. Se ha conseguido describir la morfología de todos los estadíos postembrionarios, incluyendo el poco conocido prenauplio, esencial en el proceso de eclosión. 7 Abstract The ostracods are a group of small crustaceans with worldwide distribution, whose body is protected by two lateral valves that are easily preserved in the sediment. In this paper we present the first record of the ostracod Heterocypris bosniaca Petkowski, Scharf and Keyser, 2000 for the Iberian Peninsula. It is a poorly known cipridoidean species inhabiting temporary pools. It was discovered in 2000 in Bosnia and since then its presence has been reported only in Israel (2004) and Valencia (this work). Firstly, we describe the study area; after discovering several pools with the presence of the species, these were monitored weekly to investigate its life cycle and population dynamics. Then, we tested the survival and other ecological aspects of the species in response to different treatments (desiccation, temperature, pollutants) under laboratory controlled conditions in order to better understand the ecology of natural populations. Finally, the changes suffered in individual external morphology through the ontogenetic and hatching processes are presented in detail. The data obtained indicate that the life strategy of H. bosniaca is adapted to tolerate the high stress of temporary aquatic environments with irregular hydrological cycles, and it is able to complete several generations in one year. Its life cycle is strongly influenced by temperature: the growth rate increases with temperature, but not always follows the rule of attaining larger body size at lower temperatures. The species shows a great resistance to pollutants compared to other crustaceans, which may offer advantages in the colonization of new environments. In addition, we were able to fully describe the morphology of all postembryonic stages, including the neglected prenauplius, essential in the hatching process. 8 Índice Introducción general ...............................................................................................11 Listado de artículos ...................................................................................................11 Ecosistemas acuáticos temporales ............................................................................12 Introducción a los ostrácodos ...................................................................................17 Justificación de la temática y objetivos ....................................................................25 Capítulo I On the presence of parthenogenetic populations of Heterocypris bosniaca Petkowski et al., 2000 (Crustacea: Ostracoda) in temporary rock pools of the eastern Iberian Peninsula, with insights into the ecology and life history of the species ...................................................................................... 33 Capítulo II Population dynamics and tolerance to desiccation in a crustacean ostracod adapted to life in small ephemeral water bodies .................................... 49 Capítulo III Breaking the temperature-size rule: Thermal effects on growth, development and fecundity of a crustacean from temporary waters .................... 71 Capítulo IV Acute toxicity tests with cadmium, lead, sodium dodecyl sulfate, and Bacillus thuringiensis on a temporary pond ostracod .................................... 93 Capítulo V Ontogeny of Heterocypris bosniaca (Ostracoda: Cyprididae): Description of postembryonic instars and rediscovery of the neglected A-9 stage .................................................................................................................... 113 Resumen global ......................................................................................................153 Resumen ampliado .................................................................................................153 Discusiones y perspectivas .....................................................................................157 Conclusiones generales .........................................................................................163 Anexos ......................................................................................................................167 Agradecimientos .....................................................................................................169 9 Introducción general Listado de artículos Esta Tesis Doctoral ha sido escrita en la modalidad de compendio de trabajos previamente publicados o aceptados para su publicación. Consta de las siguientes publicaciones: I. Aguilar-Alberola, J.A. & Mezquita, F. 2008. On the presence of parthenogenetic populations of Heterocypris bosniaca Petkowski et al., 2000 (Crustacea: Ostracoda) in temporary rock pools of the eastern Iberian Peninsula, with insights into the ecology and life history of the species. Bulletin de la Société des naturalistes luxembourgeois 109: 149-158. II. Aguilar-Alberola, J.A. & Mesquita-Joanes, F. 2011. Population dynamics and tolerance to desiccation in a crustacean ostracod adapted to life in small ephemeral water bodies. Limnologica 41: 348-355. III. Aguilar-Alberola, J.A. & Mesquita-Joanes, F. (en revisión, 2013). Breaking the temperature-size rule: Thermal effects on growth, development and fecundity of a crustacean from temporary waters. Journal of Thermal Biology. IV. Aguilar-Alberola, J.A. & Mesquita-Joanes, F. 2012. Acute toxicity tests with cadmium, lead, sodium dodecyl sulfate, and Bacillus thuringiensis on a temporary pond ostracod. International Review of Hydrobiology 97: 375-388. V. Aguilar-Alberola, J.A. & Mesquita-Joanes, F. 2013. Ontogeny of Heterocypris bosniaca (Ostracoda: Cyprididae): Description of postembryonic instars and rediscovery of the neglected A-9 stage. Journal of Crustacean Biology 33: 348-371. Los artículos no se presentan en un orden cronológico de publicación sino en un orden lógico para una mejor comprensión del trabajo global. Los artículos forman los principales capítulos de la Tesis y por lo tanto se les ha adaptado el formato para que sea uniforme y así ofrecer una lectura más cómoda. También se proporciona una copia electrónica de los artículos con el formato original de cada revista. 11 Introducción general Ecosistemas acuáticos temporales Los ecosistemas acuáticos temporales pueden encontrase casi en cualquier parte del mundo. Proporcionan un hábitat muy distinto del que ofrecen los sistemas permanentes puesto que su ciclo hidrológico no es continuo en el tiempo, y por lo tanto la mayoría de especies que los habitan están
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