DISTRIBUTION AND EVOLUTION OF ATLANTO- MEDITERRANEAN SPONGES FROM SHALLOW-WATER AND DEEP-SEA CORAL ECOSYSTEMS: A molecular, morphological and biochemical approach Reveillaud J., 2011. Distribution and evolution of Atlanto-Mediterranean sponges from shallow-water and deep-sea coral ecosystems: A molecular, morphological and biochemical approach. PhD thesis, Ghent University, Belgium. Publically defended on 1st June, 2011 The work presented in this dissertation was funded by an IWT fellowship (Flemish agency for Innovation by Science and Technology) and a Special Research Fund (BOF) from Ghent University. The research leading to this thesis has received funding from the European Community's Six and Seventh Framework Programme under the the HERMES project, EC contract No. GOCE-CT-2005-511234 (FP6) and the HERMIONE project, grant agreement No 226354 (FP7). UNIVERSITEIT GENT Marine Biology Research Group Campus Sterre –S8 Krijgslaan 281 B-9000 Ghent Belgium Cover design: Clara Cornu and Julie Reveillaud Layout: Clara Cornu and Julie Reveillaud Printed by: DCL, Zelzate, Belgium ISBN: 978-94-9069-574-3 Cover: Hexadella topsenti sp. nov, species described in this dissertation, photography by Thierry Pérez (front); deep-sea coral reefs in Santa Maria Di Leuca, Ionian Sea, under- water video with ROV Victor, Copyright IFREMER (back). DISTRIBUTION AND EVOLUTION OF ATLANTO- MEDITERRANEAN SPONGES FROM SHALLOW-WATER AND DEEP-SEA CORAL ECOSYSTEMS: A molecular, morphological and biochemical approach Julie Reveillaud Promotor Prof. Dr. Ann Vanreusel Co-promotors Dr. Rob Van Soest Dr. Sofie Derycke Academic year 2010-2011 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor in Science (Biology) Members of the reading committee Dr. Dirk Erpenbeck Ludwig-Maximilians-University Munich, Munich, Germany Prof. Dr. Tom Moens Ghent University, Ghent, Belgium Prof. Dr. Olivier De Clerck Ghent University, Ghent, Belgium Members of the examination committee Prof. Dr. Dominique Adriaens, Chairman Ghent University, Ghent, Belgium Prof. Dr. Ann Vanreusel, Promotor Ghent University, Ghent, Belgium Dr. Rob Van Soest, Co-Promotor Zoological Museum of Amsterdam, University of Amsterdam, Amsterdam, The Netherlands Dr. Sofie Derycke, Co-Promotor Ghent University, Ghent, Belgium Prof. Em. Dr. Jean-Pierre Henriet Ghent University, Ghent, Belgium Dr. Dirk Erpenbeck Ludwig-Maximilians-University Munich, Munich, Germany Prof. Dr. Tom Moens Ghent University, Ghent, Belgium Prof. Dr. Olivier De Clerck Ghent University, Ghent, Belgium Prof. Dr. Magda Vincx Ghent University, Ghent, Belgium Table of contents SUMMARY 1 CHAPTER I General introduction 9 CHAPTER II The distribution of scleractinian corals in the Bay of 51 Biscay, NE Atlantic CHAPTER III Species boundaries and phylogenetic relationships 79 between Atlanto-Mediterranean shallow-water and deep-sea coral associated Hexadella species (Porifera, Ianthellidae) CHAPTER IV Relevance of an integrative approach for taxonomic 105 revision in sponge taxa: case study of the shallow- water Atlanto-Mediterranean Hexadella (Porifera, Ianthellidae) CHAPTER V Phylogenetic relationships among NE Atlantic 143 Plocamionida Topsent (1927) (Porifera, Poecilosclerida): under-estimated diversity in reef ecosystems CHAPTER VI General discussion 165 ADDENDUM I Appendix to CHAPTER II 197 ADDENDUM II Appendix to CHAPTER V 211 ADDENDUM III Appendix to CHAPTER V 215 CITED LITERATURE 223 PUBLICATIONS 255 CURRICULUM VITAE 257 ACKNOWLEDGMENTS Felix qui potuit rerum cognoscere causas, wrote the poet Virgil: happy is the person who has been able to learn the causes of things. I have been indebted to many experts, colleagues, friends and family for the great help and support they provided in the preparation of this PhD, and for making this long journey - a fascinating experience. At first I would like to sincerely thank the members of the jury committee for watching out the quality of this work: Prof. Dr. Ann Vanreusel, Dr. Rob Van Soest, Dr. Sofie Derycke, Dr. Dirk Erpenbeck, Prof. Em. Dr. Jean-Pierre Henriet, Prof. Dr. Magda Vincx, Prof. Dr. Tom Moens, Prof. Dr. Olivier De Clerck, Prof. Dr. Dominique Adriaens. I am really grateful for your valuable and constructive criticism, which substantially improved the final version of this dissertation. Then, I would like to warmly thank Ann Vanreusel, my PhD supervisor. I thank you Ann for leaving your door open; first to a project far from nematodes and then to all kind of discussions, from science to topics far beyond. I am very grateful for your advices, for your guidance through the difficult idea that there is NO absolute truth (even, or particularly in science), for your exigency, and for your elegance while solving any scientific issues. I learned lot from it! So many thanks for the careful and precious corrections of the many chapters of my thesis, especially during the last redaction weeks. I am very indebted for your positive welcome to my initiatives, from within the country to far outside. I guess it is the combination of this freedom and support that made this PhD a really happy experience and that helped me to become an independent researcher. Enfin MERCI Ann pour votre amitié, votre generosité et pour tous les bons moments partagés en conference et à l‘étranger. This brings me to thank Rob Van Soest, from the Zoological Museum of the University of Amsterdam. It has been a honour to have you Rob as co-promotor of this PhD; thank you for accepting this responsibility. I am greatly indebted for the many sponge samples that you provided for this work as well as for your invaluable help on sponge taxonomy. Thank you for the time and generous advices you offered me in Amsterdam and by email the last years; they have been very important for my scientific progress and for the completion of this thesis. I am extremely grateful to Sofie, who provided inestimable and constructive advices for this PhD. Scientific problems seemed to magically disappear from discussing it with you, Sofie. I know the time it took from you and I am sincerely indebted for all your HELP. Science aside, it has been a pleasure to share an office and so many 'baby' conversations with you... Bedankt Sofie, for your precious friendship and for your empathy. Thanks for the keys of your place on a busy lab summer in Ghent as well as for the very nice dinners at your place with your loved one (s). This leads me to thank Bart, for making us discover the culture heritage of Belgium, especially the remarkable trappist Westvleteren. There is a 'Bart-Sofie-' print on our house in Brussel.... I deeply thank Jean-Pierre Henriet, who is at the beginning of all this adventure. Your true enthusiasm for science has been a great stimulus to me, Jean-Pierre. I thank you for introducing me to the fascinating world of deep-sea research, and for your great support (personal and financial) for our outreach initiative. It eventually turned into a different project but a really nice documentary. I thank you for closely following my PhD. I am really grateful to Magda Vincx, who really early told me how motivating it has been to have her children during her PhD. Thanks for sharing with me these few words and some more thoughts about the shallow vs. deep-sea marine systems. I am very indebted to Dirk Erpenbeck for his careful and timely advices on phylogenetic analyses. I thank you Dirk for such a kind invitation in Gottingen, for your willingness to look at my dataset and for your precious help all these years by email. I also would like to thank Gert Wörheide for his welcome in his lab and for the very useful explanations and tips, together with Bastian Bentlage, about the nuclear introns. I‘ll come now back a bit in time, when I just entered the Marine Biology Section. I am very thankful to Thomas Remerie for all his help and discussions during the first two years of my PhD. You helped me to go on the right road, Thomas, which I didn't forget! Flanders and Wallonia can continue to disagree; Belgium will always have the sweetest figure, Miss Rappé. Thank you Karen for your warm welcome in Belgium. Merci pour ton soutient, et pour ton amitié. It has been a pleasure to be the neighbour of the sweet Karen-Alexander family in Brussel, and then in Ghent. Katja, Chouchou, thanks for your daily smile, and for the many talks about science, baby, baby, baby...Thanks for reading and providing useful comments on my discussion. We still have to figure out what sponges and nematodes may have in common? I thank you for your warm support, your concern, and for all your friendship. Take care of the-so-cute Robyn‘ s family… Ulrijke, thank you Mevrouw for your help at these chaotic last months of my PhD. It was really nice to follow your steps and to use ‗LIST‘. From then on, all went very fine…. So veel bedankt, for your patience to speak with me ‗heel traag‘ in Nederlands; these were really sweet and pleasant private lessons… Merci Guy, merci Annick for all your help these last years. Although you both always answered ‗It s my job‘, I believe that you do it in a uniquely kind and efficient way. I really enjoyed the discussions about politics, about the Belgian anarchy without any government, and life in general with both of you. Vous êtes des petites perles pour le laboratoire de Biologie Marine… Isolde, thanks for your help with all the administrative papers. I wish there is an Isolde everywhere to make one‘s life much easier... Danke wel Annelien, for your admirable work in the lab. I am very grateful for all your help with the samples and the complex organisation of the many lab products. So many thanks for your patience, your efforts, and numerous trials to get some DNA amplification out of the most difficult samples.