High levels of realized dispersal and phenotypic plasticity in the polymorphic squat lobster Munida gregaria (Decapoda: Anomura: Munididae) in Patagonia and New Zealand - a multi-marker genetic and morphological approach Dissertation in partial fulfillment of the requirements for Doktor der Naturwissenschaften -Dr. rer. Nat.- in the Faculty of Biology and Chemistry (Fachbereich 2) University of Bremen, Germany Chen Wang 2016 Prüfungsausschuss 1. Gutachter: Prof. Dr. Thomas Brey Alfred-Wegener-Institut, Helmholtz-Zentrum für Polar- und Meeresforschung Funktionelle Ökologie – Bremerhaven 2. Gutachter: Prof. Dr. Martin Zimmer Leibniz-Zentrum für Marine Tropenökologie Ökologie 1. Prüfer: Prof. Dr. Kai Bischof Universität Bremen Marine Botanik 2. Prüfer: Dr. Christoph Held Alfred-Wegener-Institut, Helmholtz-Zentrum für Polar- und Meeresforschung Funktionelle Ökologie - Bremerhaven For my wife Yameng and our son Ziyuan Content Table of Contents ERKLÄRUNG ....................................................................................................................................... I ACKNOWLEDGEMENTS .................................................................................................................. II SUMMARY ......................................................................................................................................... V ZUSAMMENFASSUNG ................................................................................................................... VII ABBREVIATIONS ..............................................................................................................................IX 1. INTRODUCTION ........................................................................................................................ 1 1.1 Theory of speciation ......................................................................................................................... 1 1.2 Molecular markers in realizing speciation ..................................................................................... 4 1.3 Genetic divergence of widespread marine species ......................................................................... 7 1.4 The study case Munida gregaria ...................................................................................................... 9 2. AIMS, QUESTIONS AND OUTLINE OF THE THESIS ......................................................... 17 2.1 Aims and questions of the thesis .................................................................................................... 17 2.2 Outline of the thesis ........................................................................................................................ 19 3. LIST OF PUBLICATIONS AND DECLARATION OF CONTRIBUTION ............................. 21 4. ISOLATION AND CHARACTERIZATION OF ELEVEN MICROSATELLITE MARKERS FROM THE SQUAT LOBSTER MUNIDA GREGARIA (DECAPODA: MUNIDIDAE) AROUND THE FALKLAND ISLANDS/ISLAS MALVINAS ..................................................................................... 23 4.1 Abstract ........................................................................................................................................... 23 4.2 Methods and Results ...................................................................................................................... 23 4.3 Acknowledgments ........................................................................................................................... 24 5. DISCRETE PHENOTYPES ARE NOT UNDERPINNED BY GENOME-WIDE GENETIC DIFFERENTIATION IN THE SQUAT LOBSTER MUNIDA GREGARIA (CRUSTACEA: DECAPODA: MUNIDIDAE): A MULTI-MARKER STUDY COVERING THE PATAGONIAN SHELF ............ 27 5.1 Abstract ........................................................................................................................................... 27 5.2 Background ..................................................................................................................................... 28 5.3 Methods ........................................................................................................................................... 32 5.4 Results ............................................................................................................................................. 36 5.5 Discussion ........................................................................................................................................ 42 5.6 Conclusions ..................................................................................................................................... 45 5.7 Acknowledgments ........................................................................................................................... 45 5.8 Additional files ................................................................................................................................ 46 6. CROSSING THE IMPASSABLE: MITOCHONDRIAL AND MICROSATELLITE MARKERS CONFIRMED HIGH LEVEL OF REALIZED DISPERSAL AND GENE FLOW IN THE TRANSOCEANIC SQUAT LOBSTER MUNIDA GREGARIA (DECAPODA: MUNIDIDAE) ACROSS THE EAST PACIFIC BARRIER ....................................................................................................... 51 6.1 Abstract ........................................................................................................................................... 51 6.2 Introduction .................................................................................................................................... 51 6.3 Materials and Methods .................................................................................................................. 54 6.4 Results ............................................................................................................................................. 60 6.5 Discussion ........................................................................................................................................ 67 6.6 Conclusion ....................................................................................................................................... 71 Content 6.7 Additional files ................................................................................................................................ 71 7. SYNTHESIS .................................................................................................................................. 75 7.1 The power of combined molecular markers in evolutionary studies ......................................... 75 7.2 M. gregaria represents a single transoceanic species with discrete phenotypes ........................ 76 7.3 Possible mechanisms underlying the discrete phenotypes .......................................................... 78 7.4 Gene flow across the EPB .............................................................................................................. 81 7.5 Outstanding diversity in small lakes vs the lack of that in the ocean ........................................ 83 7.6 Concluding remarks ....................................................................................................................... 86 7.7 Future perspectives ........................................................................................................................ 87 REFERENCES ................................................................................................................................... 89 Erklärung Erklärung Chen Wang Borriesstr. 20 27570 Bremerhaven Erklärung gemäß §6(5) der Promotionsordnung der Universität Bremen für die mathematischen, natur- und ingenieurwissenschaftlichen Fachbereiche vom 14. März 2007 Hiermit erkläre ich, Chen Wang, dass ich die Doktorarbeit mit dem Titel: “High levels of realized dispersal and phenotypic plasticity in the polymorphic squat lobster Munida gregaria (Decapoda: Anomura: Munididae) in Patagonia and New Zealand - a multi-marker genetic and morphological approach” selbstständig verfasst und geschrieben habe und außer den angegebenen Quellen keine weiteren Hilfsmittel verwendet habe. --------------------------------------------- I Acknowledgements Acknowledgements I would like to thank so many people that helped me along over my PhD time, to some I am especially grateful. First of all, I would like to express my deepest gratitude to my PhD supervisor Dr. Christoph Held for giving me the opportunity to study in his group in the past few years and to be involved in this very exciting project. His guidance and constant inspiration teach me what a scientist should be. His knowledge and enthusiasm for new ideas make me admire and motivated. Learning from him I obtained skills in the molecular field, improved my English in speaking and writing, built the ability of developing ideas independently. I would like to thank Prof. Dr. Thomas Brey, Dr. Christoph Held, Dr. Uwe John and Dr. Bank Beszteri for their valuable suggestions during my regular PhD committee meetings. I am thankful to my colleague and friend Shobhit Agrawal for his help in data analyses and developing article ideas, as well as to Meike Seefeldt for reviewing my thesis. I would like to thank my committee members, especially Prof. Dr. Thomas Brey and Prof. Dr. Martin Zimmer for reviewing this thesis. I would like to give my special thanks to Dr. Jürgen Laudien for his great help in collecting samples in Chile. I am very grateful to Dr. Vreni Häussermann and