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Saskia Van Gaever FACULTY OF SCIENCES BIODIVERSITY , DISTRIBUTION PATTERNS AND TROPHIC POSITION OF MEIOBENTHOS ASSOCIATED WITH REDUCED ENVIRONMENTS AT CONTINENTAL MARGINS . BIODIVERSITEIT , DISTRIBUTIEPATRONEN EN TROFISCHE POSITIE VAN MEIOBENTHOS GEASSOCIEERD MET GEREDUCEERDE MILIEUS OP CONTINENTALE RANDEN . by/door SASKIA VAN GAEVER Promotor: Prof. Dr. Ann Vanreusel Academic year 2007 – 2008 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor in Science (Biology) Nature composes some of her loveliest poems for the microscope and the telescope. Theodore Roszak, Where the Wasteland Ends, 1972 MEMBERS OF THE READING COMMITTEE Prof. Dr. ANN VANREUSEL, Promotor (Ghent University) Dr. KARINE OLU (IFREMER, Brest, France) Dr. MARLEEN DE TROCH (Ghent University) Prof. Dr. TOM MOENS (Ghent University) MEMBERS OF THE EXAMINATION COMMITTEE Prof. Dr. WIM VYVERMAN, Chairman (Ghent University) Prof. Dr. ANN VANREUSEL, Promotor (Ghent University) Dr. KARINE OLU (IFREMER, Brest, France) Dr. LEON MOODLEY (NIOO, Yerseke, The Netherlands) Prof. Dr. ANNE WILLEMS (Ghent University) Dr. MARLEEN DE TROCH (Ghent University) Prof. Dr. TOM MOENS (Ghent University) Prof. Dr. MAGDA VINCX (Ghent University) VOORWOORD De onmetelijke oceaan en het wonderlijke leven daarin heeft me al sinds jongsafaan gefascineerd. Nooit had ik durven dromen dat ik ooit zou deelnemen aan wekenlange campagnes op oceanografische onderzoeksschepen... Als eerste in de rij wil ik dan ook Prof. Dr. Ann Vanreusel, mijn promotor, oprecht bedanken voor het vertrouwen in mij, en voor al de kansen die ze me heeft geboden. Ik herinner me nog levendig onze gezamenlijke ACES- workshop in Oban en het charmante huisje van Peter Lamont op dat idyllische eilandje. De talrijke congressen, symposia en workshops die daarop volgden waren elk een verrijking voor mijn wetenschappelijke visie. Maar ook de verschillende staalnamecampagnes waren ieder op zich een ervaring om nooit meer te vergeten. Ontelbare keren heb ik tijdens mijn werkshift de zonsopgang en -ondergang op zee gezien, en dat schonk me een onbeschrijflijk gevoel van rust en vrijheid. Prof. Dr. Magda Vincx wil ik hierbij mijn dank betuigen voor al de faciliteiten op het labo waarvan ik onbeperkt gebruik mocht maken. Mede dankzij haar leidinggevende functie als diensthoofd stond zij borg voor een gemoedelijke sfeer en gedisciplineerde samenwerking. I warmly thank Prof. Dr. Wim Vyverman, Dr. Karine Olu, Dr. Leon Moodley, Prof. Dr. Anne Willems, Dr. Marleen De Troch, Prof. Dr. Tom Moens and Prof. Dr. Magda Vincx for accepting to be in my thesis committee. Je voudrais cordialement remercier Dr. Karine Olu, Joëlle Galéron, Marie-Claire Fabri, Jean-Claude Caprais, Yann Garnier et tous les autres gens d’Ifremer et de Genavir qui m’ont accompagnié sur les campagnes ‘Biozaire’ et ‘Vicking’. Les passionantes discussions scientifiques, danser et swinguer sur les houles de la mer, plonger au milieu de l’Atlantic, la tradition des apéros; tout ça étaient des experiences impossible à oublier. I would like to express my sincere gratitude to Dr. Leon Moodley for the introduction and excellent guidance in the stable isotope and fatty acid research, for the motivating discussions, for the large amount of analyses (also thanks to Marco Houtekamer!) and for the support during the writing and publishing. I am also grateful to Prof. Dr. Jack Middelburg for his feedback and useful comments. Dankzij Prof. Dr. Anne Willems en Renata Coopman heb ik ook kennisgemaakt met het moleculair werk, en ik wil hun hierbij bedanken voor hun grote inzet om mij verder te helpen. Mijn welgemeende dank ook aan Dr. Marleen De Troch en Prof. Dr. Tom Moens voor hun kritisch naleeswerk en de nuttige suggesties ter verbetering van dit proefschrift, voor alle hulp bij het opzoeken van literatuur en niet in het minst ook voor de vele ‘ouder’-gesprekjes gedurende de laatse jaren. Zeven jaar lang was ik assistent op het labo Mariene Biologie, maar ze waren als in een oogopslag voorbij. De job was enorm afwisselend en boeiend, er waren altijd wel nieuwe, leuke vooruitzichten. Ook de aangename groep (ex-)collega’s zorgden ervoor dat er nooit een moment van verveling de kop opstak. Dank jullie allemaal voor de gierende lachbuien en geanimeerde discussies tijdens de koffie- en lunchpauzes, voor de ambiance tijdens de feestjes, en voor alle bereidwillige hulp die jullie mij steeds boden. Een speciale merci aan Sofie Vandendriessche, Annelies, Maarten en Eveline voor al de ‘leute’ die we reeds tijdens het studentenleven en nadien ook in onze job en daarbuiten samen beleefden.... Zoveel leuke herinneringen, het zijn er teveel om op te noemen: de suikerkick in de plantentuin, met olie besmeurde vogels gaan redden op het strand, werken en genieten op de Zeeleeuw en in de zodiac, sudderen in de sauna ... En er zullen zeker nog talrijke uitstapjes/feestjes met de partners en kindjes erbij volgen. Ook een welgemeende dankjewel aan Sofie Derycke, Annick Verween en Annick Van Kenhove voor de ontspannende zwembeurten, de vele gezellige kletsmomenten, kadootjes en kaartjes, en zoveel meer... Naast Sofie wil ik eveneens ook Thomas en Julie bedanken voor de ontspannen, moleculaire bureausfeer. We hebben samen veel plezier gehad maar er was zeker ook tijd voor de nodige peptalk. En ondertussen ken ik wel alle geheimen van het moleculair platform... Een grote merci aan Isolde, Tim, Guy, Dirk, Bart en Daniëlle voor het oplossen van die andere ‘kleine’ probleempjes, de hulp bij het praktisch werk en de toffe onderonsjes. Thanks to the always enthusiastic Italian student Francesca Pasotti for her excellent job in processing a lot of meiofauna samples, and Prof. Dr. Roberto Danovaro for his help in the writing process. Een grote dankbare knuffel aan Debby, maar ook Kristof en hun twee kleintjes Tristan en Talina! We zullen elkaar nooit uit het oog verliezen! Merci aan alle oude en nieuwe vrienden van ‘BC Fight’ en compagnie: Franky, Geert en Sofie, Lars en Mieke, Katrien en Wouter, Alexander, Bob en Joke, Hendrik en Eva, Quinten en Eveline, ... Vijftien jaar lang badmintonplezier en uitbundige Ardennen-weekendjes waren ideaal om telkens opnieuw de batterijen op te laden. Ik sta al te popelen voor ons eerstkomende weekendje binnenkort. Dankjewel aan alle (ex-)badmintonners van de dinsdagavond in het GUSB: Annelies, Sofie en Gert, Maarten, David, Wendy, Tom, Peter, Jan, Kris, Ivan, ... Onze sportieve afspraken zijn ooit van start gegaan na een gezamenlijke Belgica-campagne met David, Tom en Peter. Jullie hebben me de voorbije maanden zelden gezien, maar vanaf oktober mep ik er weer op los. Mijn grootouders, schoonouders, Nele en Geert, Jeroen en Berre, Kobe, Wannes en Nel wil ik bedanken voor alle hartelijke familiale momenten en voor hun oprechte interesse in mijn werk. Dimitri, merci om altijd en overal voor me klaar te staan! Mama en papa, duizendmaal dank voor jullie onuitputtelijke steun, hulp en ongeveinsde interesse in mijn wetenschappelijke loopbaan! Jullie schonken mij een liefdevolle, onbezorgde thuis, en hebben me steeds de vrijheid gelaten om te kiezen voor hetgene waar mijn hart naar uitging. Deze verwezenlijking is zeker ook jullie verdienste! Ik blijf jullie grootste fan… Bees, jij hebt deze reis samen met mij gemaakt en was daarbij altijd even enthousiast, geduldig en begripvol. Dit boek is een gepast einde van de voorbije zeven jaren op de Mariene Biologie. Maar mijn twee grootste meesterstukken zijn ongetwijfeld onze kindjes en oogappels: Brent en Jade. Zij schonken mij dagelijks een enorm plezier en maakten mijn geluk alleen maar completer. Die rijkende handjes, onschuldige zoentjes, knuffels en gekkebekken waren en zijn voor mij van onschatbare waarde... Merci drie lieve schatten van me! Finally I would like to thank all other friends, colleagues and scientists who supported and worked with me during this period. Saskia, Oostakker, september 2008 TABLE OF CONTENTS SUMMARY – SAMENVATTING _______________________________________________________ i SUMMARY i SAMENVATTING v CHAPTER 1 General introduction, aims and thesis outline____________________________ 3 1. The deep-sea topography 3 2. Hotspot Ecosystem Research on the Margins of European Seas (HERMES) 4 3. BIOZAIRE Project 5 4. Reduced marine environments 6 5. Seep environments in the deep sea 6 5.1. Global distribution and general features 6 5.2. Geochemistry of seeps 9 5.2.1. Brine seeps 9 5.2.2. Hydrocarbon seeps 10 5.3. Geological structures formed by seeps 11 5.3.1. Mud volcanoes 11 5.3.2. Pockmarks 12 6. Chemosynthetic benthic communities 16 6.1. Nutritional strategies of seep fauna 17 6.2. Metazoan seep community: a focus on meiofauna 18 6.3. Seep nematode diversity 19 6.4. Adaptations and tolerance of nematofauna 20 7. Aims and thesis outline 23 CHAPTER 2 The macro- and micro-scale patchiness of meiobenthos associated with the Darwin Mounds (North-East Atlantic) _____________________________________ 29 INTRODUCTION 31 MATERIAL AND METHODS 32 1. Study area 32 2. Sampling and treatment of samples 34 3. Data analyses 36 RESULTS 37 1. Variability related to the mounds 37 1.1. Sediment properties 37 1.2. Horizontal distribution on the mound 37 1.3. Vertical distribution within the sediment 40 1.4. Multivariate analyses of the nematode communities 41 2. Variability related to xenophyophores 42 DISCUSSION 44 1. Meiobenthic community 44 2. Nematode diversity 45 3. Impact of large topographic structure 46 4. Impact of small biogenic structure 47 CONCLUSION 48 ACKNOWLEDGEMENTS 48 CHAPTER 3 Deep-sea habitat heterogeneity
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