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Copepoda, Siphonostomatoida, Asterocheridae) Laboratorio de Biología Marina, Universidad de Sevilla, REVISIÓN DEL GÉNERO ASTEROCHERES BOECK 1860 (COPEPODA, SIPHONOSTOMATOIDA, ASTEROCHERIDAE) Memoria presentada para optar al título de Doctora Biología por la Universidad de Sevilla Mª EUGENIA BANDERA GARCÍA SEVILLA 2015 DÑA MERCEDES CONRADI BARRENA, PROFESORA TITULAR DEL DEPARTAMENTO DE ZOOLOGÍA DE LA UNIVERSIDAD DE SEVILLA HACE CONSTAR: Que esta memoria, titulada “Revisión del género Asterocheres Boeck, 1860 (Copepoda, Siphonostomatoida, Asterocheridae)”, presentada por Dña. María Eugenia Bandera García, resume su trabajo de Tesis Doctoral y, considerando que reúne todos los requisitos legales y las condiciones de originalidad y rigor científico, autoriza su presentación y defensa para optar al grado de Doctora en Biología por la Universidad de Sevilla. Sevilla, Abril, 2015 La directora Fdo Mercedes Conradi Barrena A la memoria de mi padre, A mi madre “…..Allí donde me he equivocado, o donde mi trabajo ha resultado imperfecto, y cuando he sido ardientemente criticado, incluso cuando me han adulado, todo lo cual me mortifica, mi mayor consuelo ha sido decirme a mí mismo miles de veces que-he trabajado tan duramente y tan bien como he sido capaz, y nadie puede hacer más que eso-…..” Charles Darwin (Autobiografía, 1876) AGRADECIMIENTOS Desde que terminé la carrera y tenía muy claro que quería hacer la tesis, hasta llegar a este momento, he recorrido un camino muy largo, a veces cuesta arriba, a veces cuesta abajo, la mayoría de las veces disfrutando de estos animalitos diminutos, otras veces con mucho esfuerzo, con paciencia, con desesperación….Sin embargo, el esfuerzo y la paciencia, no han sido solo míos sino también de otras personas a las que tengo que agradecer que yo hoy esté escribiendo la última página de esta memoria. En primer lugar, mi más profundo y sincero agradecimiento es para Mercedes, mi directora, a la que yo nunca he visto como una jefa sino como una amiga. Sin su ayuda, su esfuerzo y su infinita paciencia conmigo, yo nunca hubiera terminado esta tesis. Sentir tu apoyo “a pesar de todo” ha sido fundamental para llegar hasta el final. Contigo, Mer, espero seguir compartiendo esta pasión por los mini-crustáceos y disfrutar, sin prisas, de las charlas taxonómicas, las disecciones y del relajante mundo de pasar a tinta… Nada de esto hubiera sido posible sin el apoyo y la compresión de mi familia. Quiero agradecer a mis padres que me apoyaran cuando les dije que quería hacer la tesis. Aunque mi padre no haya visto el proceso, el pensar que pueda sentirse orgulloso de mi trabajo es lo que más me ha animado en los momentos de mayor desanimo. Los que más han padecido esta tesis han sido mi madre y Sergio, mi marido. Ellos han compartido conmigo mis momentos de euforia y mis alegrías, y han soportado mis momentos de desánimo, mi desesperación y mis lágrimas … y a pesar de todo han seguido apoyándome. Me siento muy afortunada de teneros en mi vida. A mi hijo, Alejandro, que en estos últimos meses ha perdido la paciencia y que no comprende porque algo que me gusta tanto me pone de tan mal humor. A mi hermano que soportaba mis charlas, siempre sobre el mismo tema, en los inicios. A mis tíos y primos que siempre me preguntan cuándo voy a terminar, porque no es la primera tesis en la familia, pero si la que ha durado más años. A los que se fueron con la duda de si esto llegaría a su fin algún día, y que seguramente me estarán ayudando desde donde estén. A Fani por estar siempre ahí, apoyándome y animándome y por ser una de las pocas personas en el mundo que comprende lo que siento, ¿por qué será? En este largo camino, me he cruzado con muchas personas, y a todas tengo algo que agradecerle. Principalmente a José Carlos que me permitió entrar en su grupo de investigación, con overbooking, cuando terminé la carrera, gracias por haberme abierto las puertas en aquel momento y gracias por haberme vuelto a acoger en esta parte final. Seguramente tus clases de Biología Marina hayan tenido mucha culpa de que yo quisiera dedicarme a esto. A Isabel Alfonso, con quien compartí mis inicios en este mundo, compañera de muestreos de esas otras tesis que vinieron antes que ésta, le agradezco su apoyo y compresión en aquellos primeros años. A Carlos María, siempre dispuesto a ayudar en cualquier cosa que haga falta, le agradezco sus consejos para que la redacción en español fuera lo más correcta posible. A Emilio su ayuda en la edición, a Isa esa claridad de ideas que tiene. A José “guerrita”, tu apellido no te hace justicia, ¡con lo pacífico que tú eres! Gracias por tu interés en mi trabajo durante todos estos años, siempre preguntado qué tal va y animándome. Al final, ya ves, ¡te ha tocado ser secretario! A Rony, gracias por haberme enseñado tu técnica de disección, poniendo una pieza en cada porta y haciendo un sándwich con los cubres y tu técnica para pasar los dibujos a tinta pensando en los copépodos como animales telescópicos. Gracias por mostrarte siempre dispuesto a ayudarme. A mis amigos, que no entienden muy bien de que va el asunto, pero que siempre preguntan cuándo voy a terminar. Especialmente a Úrsula, siempre mostrándome su cariño y su apoyo en cualquier cosa que yo decida hacer en la vida. Agradezco a los museos que me han prestado el material para la realización de la tesis, especialmente a Miranda Lowe, conservadora del Natural History Museum de Londres; a Dirk Platvoet, conservador del Zoological Museum de Amsterdam; a Jørgen Olesen, conservador del Zoologisk Museum of the University de Copenague; y a Lutz Bachmann y Ase Wilhelmsen, conservadores del Natural History Museum de la Universidad de Oslo. A todos ellos les agradezco su paciencia y la flexibilidad en las fechas de devolución del material. A Joaquín, el guardián de mi expediente. Siempre dispuesto a solucionar todas las dudas sobre los papeleos de tercer ciclo y siempre tan amable. ¿De verdad que he rellenado todos los anexos posibles del reglamento de doctorado? Y finalmente, ¿por qué no?, a los cientos de copépodos anónimos que donaron sus cuerpos para que yo pudiera realizar esta memoria. Creo que en la vida las cosas pasan cuando tienen que pasar, todo tiene su momento y su lugar. Seguro que hay algún motivo para que yo haya tardado tanto tiempo en leer la tesis. Quizás necesitaba recorrer este camino tan largo y aprender no sólo sobre los copépodos, sino también aprender de la vida, de la gente y sobretodo de mi misma. Ahora que parece que las piezas encajan… será que ha llegado el momento ♪♫♪ when all the parts of the puzzle start to look like they fit, then I must remember there´ll be days like this ♪♫♪ (Van Morrison, 1995). ÍNDICE CAPÍTULO 1: INTRODUCCIÓN Y OBJETIVOS 1.1. Los copépodos y su gran diversidad ……………………….…….3 1.2. El orden exclusivamente simbiótico: los Siphonostomatoida…......7 1.3. La familia Asterocheridae……………...………………………...22 1.4. El género Asterocheres.……………….…………………………...24 1.5. Objetivos………………………………………………………..28 1.6. Bibliografía……………………………………………………...28 CAPÍTULO 2: MATERIAL Y MÉTODOS……………………….………39 CAPÍTULO 3: RESULTADOS 3.1. Artículo I; Asterocheres hirsutus, a new species of parasitic copepod (Siphonostomatoida: Asterocheridae) associated with an Antarctic hexactinellid sponge………………………………………………………...51 3.2. Artículo II; The copepods associated with the coral Astroides calycularis (Scleractinia, Dendrophyllidae) in the Strait of Gibraltar……………………………………………………………………63 3.3. Artículo III; Two new asterocherid species (Siphonostomatoida: Asterocheridae) from Madeira and the Canary Islands (eastern Atlantic)…....85 3.4. Artículo IV; Proposal of new genus for Asterocheres mucronipes Stock, 1960 (Copepoda, Siphonostomatoida, Asterocheridae), an associate of the scleractinian coral Astroides calycularis (Pallas, 1766) in the Strait of Gibraltar.....................................................................................................................103 3.5. Artículo V; Some poorly known Asterocheres species (Siphonostomatoida: Asterocheridae) deposited in the Natural History Museum of London ………………………………………………………125 3.6. Artículo VI; Redescription of Asterocheres suberitis Giesbrecht, 1897 and A. tenerus (Hansen, 1923) (Copepoda: Siphonostomatoida), including notes on A. abyssi (Hansen, 1923) and A. intermedius (Hansen, 1923)……………………………………………………………………...141 3.7. Artículo VII; Two copepod species largely confused: Asterocheres echinicola (Norman, 1868) and A. violaceus (Claus, 1889). Taxonomical implications………………………………………………………….……155 3.8. Artículo VIII; Asterocherids (Copepoda: Siphonostomatoida) associated with marine invertebrates in the Strait of Gibraltar………..……173 3.9. Artículo IX; Redescription of five Asterocheres species (Copepoda: Siphonostomatoida) and a description of a new species discovered in the collections of the Zoological Museum of Amsterdam………………..……193 3.10. Artículo X; A new of Asterocheres (Copepoda, Siphonostomatoida) with a redescription of A. complexus Stock, 1960 and A. sarsi Bandera & Conradi, 2009…………………………………………………………..…217 3.11 Artículo XI; Redescription of two Asterocheres species (Copepoda: Siphonostomatoida): A. corneliae Schirl, 1973 and A. boecki (Brady, 1880), and a proposal of new genus for A. fastigatus Kim, 2010………………………....237 CAPÍTULO 4: DISCUSIÓN GENERAL………………………………..271 CAPÍTULO 5: CONCLUSIONES……………………………………………..339 RESUMEN Los copépodos son un grupo de diminutos crustáceos acuáticos con una amplia diversidad de estructuras y hábitos así como una gran capacidad de adaptación. Aunque estos pequeños crustáceos son conocidos como
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