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Genetic Divergence, Reproductive Isolation and the Early Stages of Speciation

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Genetic Divergence, Reproductive Isolation and the Early Stages of Speciation Genetic divergence, reproductive isolation and the early stages of speciation Silje Hogner Dissertation presented for the degree of Philosophiae Doctor Natural History Museum Faculty of Mathematics and Natural Sciences University of Oslo © Silje Hogner, 2013 Series of dissertations submitted to the Faculty of Mathematics and Natural Sciences, University of Oslo No. 1305 ISSN 1501-7710 All rights reserved. No part of this publication may be reproduced or transmitted, in any form or by any means, without permission. Cover: Inger Sandved Anfinsen. Printed in Norway: AIT Oslo AS. Produced in co-operation with Akademika publishing. The thesis is produced by Akademika publishing merely in connection with the thesis defence. Kindly direct all inquiries regarding the thesis to the copyright holder or the unit which grants the doctorate. Supervisor Co-supervisor Arild Johnsen Jan T. Lifjeld Associate Professor Professor Natural History Museum Natural History Museum University of Oslo University of Oslo Norway Norway Adjudicating committee 1st opponent Jacob Höglund Professor Department of Ecology and Genetics Population Biology and Conservation Biology Uppsala University Sweden 2nd opponent Laura Kvist Docent/Phd Department of Biology University of Oulu Finland Administrative leader Øystein Wiig Professor Natural History Museum University of Oslo Norway Acknowledgements The work of this PhD thesis was undertaken at the Natural History Museum, University of Oslo, and would not have been possible to complete without the help of several people. First, I would like to thank my supervisor Arild Johnsen. It has been a privilege to have you as my supervisor and to work with you. Your enthusiasm, knowledge and support have been a great inspiration. Many thanks also to Jan T. Lifjeld for being my co-supervisor. To both of you, thank you for inspiring me to be a better scientist and for always taking the time to discuss my work. I would like to thank Terje Laskemoen, Gunnhild M. Marthinsen, Lars Erik Johannessen, Melissah Rowe, Jostein Gohli, Even Stensrud, Oddmund Kleven, Trond Øigarden, Becky Cramer, Taiwo Crossby Omotoriogun and Jarl Andreas Anmarkrud, also known as the “bird group”. Thank you for providing such a friendly, dynamic and supportive research environment. I have enjoyed the numerous discussions we have had during coffee-breaks and lunches. A special thanks to Terje, for help with sperm analyses and statistics. Lars Erik thank you for helping me with all the formatting, and Gunnhild thank you for valuable help in the lab. Melissah, thank you for helping me improve my writing, for all the fruitful and helpful discussions we had during the final stages of my PhD and for always encouraging me to continue. It was a pleasure sharing an office with you. Kjersti S. Kvie, thank you for being a great friend and a great colleague. Thank you for always having time for conversation and support when things may have seemed too chaotic and unmanageable. Eirik Rindal, thank you for valuable help with statistics. Thanks also to Vegar Bakkestuen for help with mapping of geographical distributions. I am also grateful to all my colleagues at NHM for making my PhD a memorable time. This thesis would not have been possible to carry out without field work and extensive sampling of both blood- and sperm samples. Thanks to all who were involved in the sampling of both birds and moths and a special thanks to all of you making it possible for me to do field work in Trysil, Karmøy, Heimdalen, Bulgaria, France and Czech Republic. I had a great time collecting samples in all of those places, and it will not be forgotten. I also want thank my supervisor during my master thesis, Glenn-Peter Sætre. Thank you for helping me with the flycatcher paper, for making it possible for me to have a second field trip to Bulgaria catching semicollared flycatcher and for inspiring me to start working on a PhD. Tore Oldeide Elgvin and Stein Are Sæther, thank you for all the discussions we had when I was struggling with the flycatcher paper. Jo Skeie Hermansen, thank you for taking time off your PhD to help me catch birds in Bulgaria. To all my co-authors (Jiri Porkert, Tamer Albayrak, Bekir Kabasakal, Leif Aarvik, Václav Pavel, Bohumír Chutný, Javier García, Marie-Christine Eybert, Ekaterina Matsyna, Thomas Borge and Torbjørn Bruvik), thank you for sharing your data with me and for helping me improve our manuscripts. Marius and Natalia, thank you for help taking care of my horse, especially during the last final weeks of my writing process. Keeping him would not have been possible without your help. Finally a big thanks to my family. Eirik, thank you for letting me use your apartment while you were working abroad. Mom and dad, thank you for always believing in me, never giving up and always encouraging me to continue. Thank you for taking interest in my work, no matter how abstract it might seem to you. Oslo, December 2012 Silje Hogner Contents List of papers ......................................................................................................................... 1 1 Abstract ............................................................................................................................... 2 2 General introduction ........................................................................................................... 4 2.1 Main aims .............................................................................................................. 7 3 General methods ................................................................................................................. 8 3.1 Study taxa ......................................................................................................................... 8 3.2 Field work and sampling ................................................................................................ 12 3.3 DNA extraction, PCR and sequencing ........................................................................... 13 3.4 Sperm analysis ............................................................................................................... 13 4 Summary of publications .................................................................................................. 14 4.1 Paper I: Deep sympatric mitochondrial divergence without reproductive isolation in the common redstart Phoenicurus phoenicurus ....................................................... 14 4.2 Paper II: Deep sympatric mtDNA divergence in the autumnal moth (Epirrita autumnata) .................................................................................................................... 15 4.3 Paper III: Rapid sperm evolution in the bluethroat (Luscinia svecica) subspecies complex ........................................................................................................................ 17 4.4 Paper IV: Increased divergence but reduced variation on the Z chromosome relative to autosomes in Ficedula flycatchers: differential introgression or the faster-Z effect? .............................................................................................................. 17 5 General discussion.......................................................................................................... .. 19 5.1 Speciation in reverse ...................................................................................................... 19 5.2 Early speciation/rapid evolution of phenotypic traits..................................................... 20 5.3 Recent speciation, Z chromosome evolution and the faster-Z hypothesis ..................... 21 5.4 DNA barcoding .............................................................................................................. 22 5.5 Genetic divergence and the process of speciation .......................................................... 23 5.6 Speciation and sexual selection ...................................................................................... 24 6 Conclusions ...................................................................................................................... 26 7 Future prospects ................................................................................................................ 27 8 Literature cited .................................................................................................................. 29 Individual papers I-IV List of papers I Silje Hogner, Terje Laskemoen, Jan T. Lifjeld, Jiri Porkert, Oddmund Kleven, Tamer Albayrak, Bekir Kabasakal and Arild Johnsen (2012). Deep sympatric mitochondrial divergence without reproductive isolation in the common redstart Phoenicurus phoenicurus. Ecology and Evolution 2: 2974–2988. II Kjersti S. Kvie, Silje Hogner, Leif Aarvik, Jan T. Lifjeld, and Arild Johnsen (2013). Deep sympatric mtDNA divergence in the autumnal moth (Epirrita autumnata). Ecology and Evolution 3: 126–144. III Silje Hogner, Terje Laskemoen, Jan T. Lifjeld, Václav Pavel, Bohumír Chutný, Javier García, Marie-Christine Eybert, Ekaterina Matsyna and Arild Johnsen. Rapid sperm evolution in the bluethroat (Luscinia svecica) subspecies complex. Manuscript to be submitted to Behavioral Ecology and Sociobiology. IV Silje Hogner, Stein A. Sæther, Thomas Borge, Torbjørn Bruvik, Arild Johnsen and Glenn-Peter Sætre (2012). Increased divergence but reduced variation on the Z chromosome relative to autosomes in Ficedula flycatchers: differential introgression or the faster-Z effect? Ecology and Evolution 2: 379–396.
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