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Université De La Méditerranée, France UNIVERSITÉ DE LA MÉDITERRANÉE, FRANCE Centre d'Océanologie de Marseille Unité Mixte de Recherche CNRS 6540 - DIMAR Diversité, Évolution et Écologie Fonctionnelle Marine UNIVERSITÉ DE GDANSK, POLOGNE Institute of Oceanography Department of Marine Ecosystems Functioning THÈSE DE DOCTORAT Spécialité Biosciences de l'Environnement présentée par Katarzyna TARNOWSKA Genetic structure and physiological variation of a widespread European lagoon specialist Cerastoderma glaucum (Bivalvia) living in extreme environmental conditions Soutenue le 25 mars 2010 Jury : Anne Chenuil, Chargée de recherche, CNRS, Marseille (Directrice de thèse) Jean-Pierre Féral, Directeur de recherche, CNRS, Marseille (Directeur de thèse) Maciej Wo łowicz, Professeur, Université de Gda Lsk, Gda Lsk, Pologne (Directeur de thèse) Sophie Arnaud-Haond, Chargée de recherche, IFREMER, Brest (Rapporteur) Herman Hummel, Professeur, Netherlands Institute of Ecology, Yerseke, Pays-Bas (Rapporteur) Philippe Borsa, Chargé de recherche, IRD, Montpellier (Examinateur) Didier Forcioli, Maitre de conférence, Université de Nice Sophia Antipolis , Nice (Examinateur) Jerzy Sell, Professeur, Université de Gda Lsk, Pologne (Président, Examinateur) TO MY PARENTS 1 ACKNOWLEDGEMENTS First of all, I would like to thank the Director of the Department of Marine Ecosystems Functioning at the University of Gda Lsk, Prof. Maciej Wołowicz and the Director of DIMAR (“Diversité, évolution et écologie fonctionnelle marine”) laboratory at the Université de la Méditerranée Dr Jean-Pierre Féral for accepting me as a PhD student in Your research teams and enabling me to work on this PhD project. I would like to thank my supervisors: Dr Anne Chenuil and Dr Jean-Pierre Féral and Prof. Maciej Wołowicz for sharing Your valuable experience and all the support and encouragement You gave me. I appreciate Your contagious excitement for my work and the criticism at the moments, when it was needed. Special thanks to Dr Anne Chenuil for a lot of patience and help while showing me the fascinating world of molecular biology. I would like to thank: • Dr A. Rogers from the Zoological Society of London for sharing microsatellite primers sequences, which enabled me to enrich my work a lot, • all the people who contributed to samples collection: E. Egea, J.-P. Féral, H. Hummel, J. Jansen, B. Kelemen , T. Kevrekidis, A. Kiewro, M. Krakau, S. Krupi Lski, R. Lasota, M. Machado, C. Marschal, A. E. Mogias , G. Sara, R. Sussarellu, J. Zaouali, H. Zibrowius, • J. Jansen, H. Hummel, O. Van Hoesel from The Netherlands Institute of Ecology (NIOO-KNAW) in Yerseke for helping me when I was investigating physiological parameters of C. glaucum from the North Sea, • all the people who helped me somehow at the different stages of my work, especially: R. Nikula, A. Green, J. Drzazgowski, K. Jaworski, K. Bradtke and the staff of the research vessel "Oceanograf II", • all the colleagues from the laboratories both in Poland and in France, especially: Kenza, Emilie, Didier, Karolina, Magda, Jean-Baptiste, Justyna, Michał, Pierre-Alexandre, Iwona, Adam, Defne, Karin, Rafał. You gave me a lot of help and advice and You also brought some light and smile to my long days at work, • R. Lasota, D. Rush, A. Putteeraj, J. Zieli Lski and J. Dunder for editorial and language corrections and suggestions. 2 I would like to acknowledge the provisions of the French government studentship, which enabled me to work on this Polish-French PhD project and the financial support from: the Ministry of Science and Higher Education in Poland (project number: N304 047 32/2162) and the University of Gda Lsk in Poland (project number: BW/13A0-5-0089-6). This research was performed in the frame of MarBEF (Marine Biodiversity and Ecosystem Functioning EU Network of Excellence) GBIRM (Genetic Biodiversity Responsive Mode) Project. MarBEF is funded by the European Union within the “Sustainable development, global change and ecosystems” RTD Programme (Contract no. GOCE-CT-2003-505446 2004-2009). I am very grateful for the possibility to participate in this project and for all the fruitful meetings. I would also like to thank: • my Mum and Dad. You always let me follow my own path in life. Thank you for your endless love, patience and support. You taught me to work hard and not to be afraid to reach far, • my friends, You went with me through all ups and downs. If it hadn’t been for your support, I wouldn’t have made it, • Fabien, for giving me a lot of support and motivation in the last, the most difficult part of this PhD, • many others who are not mentioned here, but helped me a lot. 3 CONTENTS 1 Introduction ..................................................................................................................... 8 1.1 Goals of the study .................................................................................................... 8 1.2 Outline of the thesis ................................................................................................. 9 2 Lagoons ........................................................................................................................... 9 2.1 Lagoon habitats ....................................................................................................... 9 2.2 Colonization and gene flow in lagoon habitats ..................................................... 10 3 Basic information on Cerastoderma glaucum .............................................................. 13 3.1 Distribution and ecological preferences ................................................................ 13 3.2 Life history ............................................................................................................. 14 3.3 Diet ......................................................................................................................... 15 3.4 On taxonomy, divergence and fossil record of C. glaucum ................................... 15 3.5 Distinguishing between C. glaucum and C. edule ................................................. 18 Chapter I: Studies on morphometry and physiology of Cerastoderma glaucum ................ 22 4 Interest in studying physiological responses of C. glaucum ......................................... 22 5 Materials and methods .................................................................................................. 24 5.1 Sampling and hydrological conditions of sampling sites ...................................... 24 5.1.1 Hydrological conditions in the Gulf of Gda Lsk .............................................. 26 5.1.2 Hydrological conditions in the Lake Veere .................................................... 27 5.1.3 Hydrological conditions in the Berre Lagoon ................................................ 29 5.2 Morphometric and physiological analysis ............................................................. 31 6 Results ........................................................................................................................... 33 7 Discussion ..................................................................................................................... 38 7.1 Environmental conditions ...................................................................................... 38 7.2 Morphometric parameters and condition index .................................................... 39 7.3 Reproduction .......................................................................................................... 40 7.4 Gross biochemical composition ............................................................................. 41 7.5 Respiration ............................................................................................................. 43 8 Conclusions ................................................................................................................... 44 Chapter II: Phylogeography of Cerastoderma glaucum ...................................................... 45 9 Theoretical background ................................................................................................ 45 9.1 Factors influencing genetic structures in marine species ..................................... 45 9.1.1 Role of glaciations in genetic structures of marine species ............................ 45 9.1.2 Dispersal modes and barriers in marine species; the case of C. glaucum ..... 46 9.2 Different properties of genetic markers applied .................................................... 49 9.3 Theoretical context of methods applied to analyze genetic data ........................... 53 9.3.1 Genetic diversity within populations .............................................................. 53 9.3.2 F-statistics ....................................................................................................... 53 9.3.3 AMOVA ........................................................................................................... 56 9.3.4 Multidimensional scaling (MDS) .................................................................... 57 9.3.5 Mantel test ....................................................................................................... 57 9.3.6 Detecting inbreeding ....................................................................................... 58 9.3.7 Neutrality tests ................................................................................................ 59 9.3.8 Selection detection- F ST -outlier method ........................................................
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