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Ruditapes Hybrids EIDO Escola Internacional de Doutoramento TESE DE DOUTORAMENTO Caracterización citogenética de bivalvos veneroideos y algunos de sus parásitos Cytogenetic characterization of veneroid bivalves and some of their parasites Daniel García Souto 2017 Mención internacional Aquéllos que me busquen, sabrán que he existido. Los demás no tienen necesidad de saberlo. Gustav Mahler AGRADECIMIENTOS Estimado lector anónimo, sea Ud. consciente de que, más allá de la loable función de calzar algún que otro mueble cojo, esta tesis es la consecución, el pináculo quizás, de una etapa de considerable esfuerzo en mi vida. Pero ante todo, sepa que se encuentra ante el trabajo de muchas personas. Esta es, en gran parte la (enésima) tesis de Juanjo, mi director y de Paloma, mi social manager. Ambos han tenido a bien aguantarme un buen rato y, siendo francos, han contribuido tanto (o más) que uno mismo al presente manuscrito. Creo, sinceramente que me habéis ayudado a madurar como investigador y como persona. Espero haber estado a la altura de vuestras expectativas. A ambos muchas gracias. Tras las presentes páginas subyace además el trabajo colaborativo de todo un departamento cuyos integrantes, incluso de forma inconsciente, han contribuido a hacer de mi estancia en el bucólico CUVIlete más cómoda, más agradable y, en definitiva, mejor. Gracias pues a todos los que están y a los que en su momento estuvieron. Esta es también la tesis de los Erasmus-party de la Universidad de Portsmouth (en estricto orden de aparición Neil, Jack, Vesa y Auriel) y de los ocupas de turno del grado en Biología (Jonathan, Pablo, Sonia, Yaiza, Sandra, Susana, Gonzalo y Agustín). En cierta medida todos habéis puesto vuestro granito de arena y me he reído mucho a vuestra costa he disfrutado mucho de vuestra compañía. Espero que hayáis podido aprender de mi la mitad de lo que yo de vosotros. A todos gracias. A la buena gente de Croacia, Brankica, Miro, Tanja Martina y tantos otros, que me brindaron un invierno un poco más largo de lo normal y una buena cantidad de conocimientos en aquello de la biología molecular. Fue realmente un acierto el recurrir a vosotros. Por el buen trato y lo bien que nos lo pasamos, hvala za sve. Gracias igualmente al grupo Chromevol al completo, Chema, Rodri, Vicky, Javier, Lu I “The Mighty” y tantos otros por el invierno que me salté en Miami. Mi estancia en Florida no podría haber sido más placentera con vuestra compañía. Mil gracias por la oportunidad. Cómo olvidar la impagable compañía del Lisensiado, Sr. Magíster y Dottore Rodríguez-Seijo. Gracias por las sobremesas en cafetería y tus sabios consejos. El fuego purificador asolará este lugar y de sus cenizas erigiremos la Universidad de la Calle. Quisiera invertir unas líneas en agradecer, además, a la Dra. Pelos, la Verdadera y Única Presidenta de la AGKM, por sus rescates a última hora y los viajes de taxi personalizados. Ha sido un alivio el tenerte ahí. Gracias Esther. Gran mérito yace también, de forma indiscutible, en el buen hacer de la Dra. Debbenneddetti y del buen (cuasi) Dr. Don Jesús (ante la limitación de espacio el autor se abstiene de recrear sus apellidos de forma íntegra… Espero lo sepan entender…). Vuestra compañía, afecto y apoyo incondicional a lo largo de estos años me han incentivado a continuar hasta este punto. Sin vosotros no lo habría logrado. Por todo, gracias Kela, gracias Gusu. Gracias también a mi familia, especialmente a mis padres y hermanos. No soy la persona más sociable del planeta y os ha tocado, más para mal que para bien, el aguantarme. Sin vuestra ayuda nada de esto hubiera sido posible. Gracias Chu. Algún día sabré compensarte como es debido, señorita. A todos, gracias. This work was partly funded by grants from Xunta de Galicia and Fondos FEDER “Unha maneira de facer Europa” (08MMA023310PR; Grupos de Referencia Competitiva, 2010/80; Grupos con Potencial Crecimiento, GPC2013-011; Axudas do programa de consolidación e estruturación de unidades de investigacións competitivas do SUG: ED431C 2016-037). Daniel García Souto was beneficiary of grants from Xunta de Galicia (PRE/2012/175) and Spanish Ministry of Education (FPU: AP-12/05040). Daniel García Souto research stays in Zagreb (Croatia, 2015) and Miami (USA, 2016) were funded by research stay grants of the Spanish Ministry of Education FPU program (EST14/00151, EST15/00486). INDEX / ÍNDICE SUMMARY ................................................................................................................................................ 1 RESUMEN ................................................................................................................................................. 6 INTRODUCTION ................................................................................................................................... 11 OBJETIVES ............................................................................................................................................. 19 VENERIDAE ........................................................................................................................................... 21 Divergent evolutionary behavior of H3 histone gene and rDNA clusters in venerid clams Molecular Cytogenetics 8:40 (2015) ......................................................................................... 23 Resolving the taxonomic status of Chamelea gallina and C. striatula (Veneridae, Bivalvia): A combined molecular cytogenetic and phylogenetic approach BioMed Research International (accepted 2017 04 03) .......................................................... 33 MACTRIDAE ......................................................................................................................................... 56 Molecular cytogenetics in trough shells (Mactridae, Bivalvia): Divergent GC-rich heterochromatin content Genes 7: 47 (2016) ...................................................................................................................... 57 Methylation profile of a satellite DNA constituting the intercalary CG-rich heterochromatin of the cut trough shell Spisula subtruncata (Bivalvia, Mactridae) Scientific Reports (under review, 2017 03 31) ......................................................................... 67 DONACIDAE ......................................................................................................................................... 106 Are pericentric inversions reorganizing wedge shell genomes? ......................................................... 107 TELLINIDAE ........................................................................................................................................ 126 Karyotype differentiation in tellin shells (Bivalvia, Tellinidae) .......................................................... 127 DIGENEA ............................................................................................................................................... 143 Molecular cytogenetics in digenean parasites: Linked and unlinked major and 5S rDNAs, B chromosomes and karyotype diversification Cytogenetics and Genome Research 147: 195-207 (2015) .................................................... 144 DISCUSIÓN GENERAL ..................................................................................................................... 157 Números cromosómicos y cariotipos ................................................................................................. 159 Heterocromatina constitutiva, DNA satélites y metilación ................................................................. 161 Mapas físicos de familias multigénicas .............................................................................................. 163 CONCLUSIONS .................................................................................................................................. 169 REFERENCES .................................................................................................................................... 171 ANNEX I Curriculum vitae ................................................................................................................... 184 SUMMARY Summary Bivalves are a group of exclusively aquatic mollusks characterized by presenting a laterally compressed body whose soft parts are covered by a shell (Gosling 2015). More than 8500 species are currently accepted (World Register of Marine Species, WoRMS, http://www.marinespecies.org/) and they are distributed worldwide inhabiting marine, brackish and, to a lesser extent, freshwater ecosystems. The class Bivalvia includes a considerable number of species of great economic interest that have been traditionally subjected to extensive exploitation on commercial purposes. Their ecological role is of high relevance as they constitute both the basis of trophic networks and the physical support for the development of numerous benthonic animal species, both vertebrates and invertebrates. Bivalve mollusks also act as intermediary hosts in the life cycle of many digenean trematodes, parasitic metazoans that may have a huge impact on bivalve populations, both natural and cultured. Although the classification of bivalves is a rather controversial subject and recent phylogenetic analyses (Bieler et al. 2014) propose six monophyletic lineages within Bivalvia, WoRMS still utilizes four subclasses, Protobranchia, Pteriomorphia, Paleoheterodonta and Heterodonta. The subclass Heterodonta includes more than two thirds of the extant bivalves,
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