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Unravelling Mechanisms of Reproductive Isolation Between Two Sister Species of Iberian Voles

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Unravelling Mechanisms of Reproductive Isolation Between Two Sister Species of Iberian Voles UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Unravelling mechanisms of reproductive isolation between two sister species of Iberian voles Doutoramento em Biologia Biologia Evolutiva Margarida Alexandra de Sousa Carvalho Tavares Duarte Tese orientada por: Doutora Cristiane Bastos-Silveira Professora Doutora Maria da Luz Mathias Documento especialmente elaborado para a obtenção do grau de Doutor 2016 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS Unravelling mechanisms of reproductive isolation between two sister species of Iberian voles Doutoramento em Biologia Biologia Evolutiva Margarida Alexandra de Sousa Carvalho Tavares Duarte Tese orientada por: Doutora Cristiane Bastos-Silveira Professora Doutora Maria da Luz Mathias Júri: Presidente: • Professor Doutor Pedro Miguel Alfaia Barcia Ré Vogais: • Doutor Gerald Heckel • Professor Doutor Carlos Manuel Martins Santos Fonseca • Professor Doutor António Paulo Pereira de Mira • Doutora Susana Sá Couto Quelhas Lima Mainen • Professora Doutora Maria da Luz da Costa Pereira Mathias • Professor Doutor Octávio Fernando de Sousa Salgueiro Godinho Paulo Documento especialmente elaborado para a obtenção do grau de Doutor Fundação para a Ciência e Tecnologia (SFRH/BD/70646/2010; PTDC/BIA- BEC/103729/2008; UID/AMB/50017); FCT/MEC; FEDER 2016 The Ph.D. candidate was awarded with studentship SFRH/BD/70646/2010, financed by Fundação para a Ciência e Tecnologia (FCT). The present Ph.D. project was developed under PTDC/BIA-BEC/103729/2008 ”Understanding the origin and persistence of Microtus species in the Iberian Peninsula: the genetical, behavior and ecological forces” (FCT), coordinated by Cristiane Bastos-Silveira. It was also supported by Centro de Estudos de Ambiente e Mar through National (UID/AMB/50017 and FCT/MEC) and European (FEDER, within the PT2020 Partnership Agreement and Compete 2020) funds. The present Ph.D. dissertation should be cited as: Duarte MA (2016) Unravelling mechanisms of reproductive isolation between two sister species of Iberian voles. Dissertation, Faculdade de Ciências, Universidade de Lisboa, Lisboa. “Success is not final, failure is not fatal: it is the courage to continue that counts.” Winston Churchill Preliminary notes According to the Post‐graduate Studies Regulation (Diário da República, 2.ª série, N.º 57, published on the 23th of March, 2015), the present dissertation includes papers published in ISI-indexed scientific journals and manuscripts in preparation for submission. These papers are included in Chapters 2, 3, 4 and 5, and were formatted according to the dissertation style, except for specific requirements made by the scientific journals, such as references and measurement units. The Ph.D. candidate is presented as the first author since it was responsible for the scientific design, data collection, analysis of the results and writing of the papers, under the supervision of Cristiane Bastos-Silveira (primary supervisor) and Professor Maria da Luz Mathias (secondary supervisor). Lisbon, 25th of July, 2016 Margarida Alexandra de Sousa Carvalho Tavares Duarte I II Table of contents Preliminary notes ............................................................................................................... I Acknowledgments ............................................................................................................ III Resumo .............................................................................................................................. IX Abstract ............................................................................................................................ XV Chapter 1 - General Introduction .................................................................................... 1 1.1 The genus Microtus ............................................................................................................... 3 1.1.1 Microtus voles of the Iberian Peninsula ............................................................... 5 1.2 Iberian sister species ........................................................................................................... 6 1.2.1 The Lusitanian pine vole ........................................................................................... 8 1.2.2 The Mediterranean pine vole ................................................................................ 10 1.2.3 Mating system ............................................................................................................. 12 1.2.4 Reproductive isolation barriers ............................................................................. 13 1.3 Aims and hypotheses ......................................................................................................... 16 1.4 References ............................................................................................................................. 18 Chapter 2 - Candidate genes related to odour cues communication ........................ 29 2.1 Olfactory receptors ............................................................................................................. 31 2.1.1 Abstract ......................................................................................................................... 31 2.1.2 Keywords ...................................................................................................................... 32 2.1.3 Introduction ................................................................................................................. 32 2.1.4 Materials and methods ............................................................................................ 34 2.1.5 Results and discussion ............................................................................................. 36 2.1.6 Acknowledgments ..................................................................................................... 43 2.1.7 References .................................................................................................................... 43 2.1.8 Supplementary material .......................................................................................... 51 2.2 Major histocompatibility complex I and II ................................................................. 60 2.2.1 Abstract ......................................................................................................................... 60 III 2.2.2 Keywords ...................................................................................................................... 60 2.2.3 Introduction ................................................................................................................. 61 2.2.4 Materials and methods ............................................................................................ 63 2.2.5 Results and discussion ............................................................................................. 66 2.2.6 Acknowledgements ................................................................................................... 69 2.2.7 References .................................................................................................................... 69 Chapter 3 - Urinary protein expression ........................................................................ 75 3.1 Urinary protein expression .............................................................................................. 77 3.1.1 Abstract ......................................................................................................................... 77 3.1.2 Keywords ...................................................................................................................... 77 3.1.3 Introduction ................................................................................................................. 78 3.1.4 Materials and methods ............................................................................................ 80 3.1.5 Results and discussion ............................................................................................. 82 3.1.6 Acknowledgments ..................................................................................................... 86 3.1.7 References .................................................................................................................... 86 Chapter 4 - Partner preference ..................................................................................... 89 4.1 Partner preference in an artificial syntopic environment .................................... 91 4.1.1 Abstract ......................................................................................................................... 91 4.1.2 Keywords ...................................................................................................................... 92 4.1.3 Introduction ................................................................................................................. 92 4.1.4 Materials and methods ............................................................................................ 93 4.1.5 Results ........................................................................................................................... 94 4.1.6 Discussion ..................................................................................................................... 96 4.1.7 Compliance with ethical standards ..................................................................... 99 4.1.8 References ...................................................................................................................
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