Thomas Vanagt

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Thomas Vanagt FACULTEIT WETENSCHAPPEN The role of swash in the ecology of Ecuadorian sandy beach macrofauna, with special reference to the surfing gastropod Olivella semistriata De rol van swash in de ecologie van macrofauna op Ecuadoriaanse zandstranden, met speciale aandacht voor de surfende gastropode Olivella semistriata Thomas Vanagt Promotor: Prof. Dr. Magda Vincx Co-promotor: Dr. Steven Degraer Academic year 2006 – 2007 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor in Science (Biology) Members of the reading committee Prof. Dr. Magda Vincx, promotor (UGent, Belgium) Dr. Steven Degraer, co-promotor (UGent, Belgium) Prof. Dr. Mariano Lastra (University of Vigo, Spain) Prof. Dr. Jean-Pierre Maelfait (UGent, Belgium) Members of the examination committee Prof. Dr. Wim Vyverman, Chairman (UGent, Belgium) Prof. Dr. Magda Vincx, promotor (UGent, Belgium) Dr. Steven Degraer, co-promotor (UGent, Belgium) Prof. Dr. Mariano Lastra (University of Vigo, Spain) Prof. Dr. Jean-Pierre Maelfait (UGent, Belgium) Dr. Dries Bonte (UGent, Belgium) Prof. Dr. Maria Pilar Cornejo (ESPOL, Ecuador) Dr. Nancy Fockedey (VLIZ, Belgium) Prof. Dr. Ann Vanreusel (UGent, Belgium) ‘Treat the Earth well: it was not given to you by your parents, it was loaned to you by your children. We do not inherit the Earth from our ancestors, we borrow it from our children’ Native-American saying TABLE OF CONTENTS Dankwoord Samenvatting I Summary VII Resumen XII PART I INTRODUCTION AND AIMS 1 1.1 Introduction 3 1.1.1 Sandy beaches 3 1.1.2 Macrofauna 12 1.1.3 Survival strategies of macrofauna on sandy beaches 20 1.1.4 The sandy beach gastropod Olivella semistriata 25 1.1.5 Framework of this thesis 27 1.2 Aims and thesis outline 28 MACROFAUNA ON ECUADORIAN SANDY BEACHES: THE ROLE OF SWASH PART II 31 AND MORPHODYNAMICS Changes in macrofaunal assemblage structure Chapter 1 along a morphodynamic sandy beach gradient 33 in the eastern tropical Pacific Abstract 34 2.1.1 Introduction 35 2.1.2 Materials and Methods 37 2.1.3 Results 43 2.1.4 Discussion 63 2.1.5 Conclusions 72 Swash zonation patterns of two surfers on exposed tropical Chapter 2 73 sandy beaches Abstract 74 2.2.1 Introduction 75 2.2.2 Materials and Methods 77 2.2.3 Results 82 2.2.4 Discussion 88 2.2.5 Conclusions 94 Size distribution of the surfing gastropod Olivella semistriata Chapter 3 95 along a cross-shore gradient in an Ecuadorian sandy beach Abstract 96 2.3.1 Introduction 97 2.3.2 Material and Methods 98 2.3.3 Results 101 2.3.4 Discussion 108 2.3.5 Conclusions 112 TABLE OF CONTENTS SOME ASPECTS OF THE BIOLOGY OF THE GASTROPOD OLIVELLA PART III 113 SEMISTRIATA Is the burrowing performance of a sandy beach surfing Chapter 4 115 gastropod limiting for its macroscale distribution? Abstract 116 3.4.1 Introduction 117 3.4.2 Material and Methods 118 3.4.3 Results 121 3.4.4 Discussion 129 3.4.5 Conclusions 133 Can sandy beach molluscs show an endogenous controlled Chapter 5 circatidal migration behaviour? Hints from a swash rig 135 experiment Abstract 136 3.5.1 Introduction 137 3.5.2 Material and Methods 139 3.5.3 Results 144 3.5.4 Discussion 146 3.5.5 Conclusions 149 Surfing on sandy beaches: an efficient strategy to flourish in a Chapter 6 151 highly dynamic environment? Abstract 152 3.6.1 Introduction 153 3.6.2 Material and Methods 155 3.6.3 Results 148 3.6.4 Theoretical models 168 3.6.5 Discussion 173 3.6.5 Conclusions 180 Appendix 1 181 Appendix 2 182 PART IV GENERAL DISCUSSION 185 4.1 Sandy beaches in Ecuador 187 4.2 Distribution of macrofauna in Ecuador 189 4.2.1 Temporal effects 190 4.2.2 Sandy beach macrofauna and ENSO 191 4.2.3 Spatial distribution 192 4.2.4 Explaining the spatial patterns 195 4.3 Biology of Olivella semistriata 199 4.3.1 Distribution of the species 199 4.3.2 Feeding habits 203 4.3.3 Surfing behaviour 204 4.4 Open questions 207 4.4.1 Studying macrofauna on exposed beaches 207 4.4.2 Macrofauna and ENSO 207 4.4.3 Olivella semistriata 207 TABLE OF CONTENTS PART V ADDENDA 209 Macrofaunal community structure and zonation of an Ecuadorian Addendum 1 211 sandy beach (bay of Valdivia) Critical note 228 Spatial and temporal patterns in the sandy beach surf zone Addendum 2 231 benthos of the tropical East Pacific. ENSO and sandy beach macrobenthos of the tropical East Addendum 3 255 Pacific: some speculations Reference List 267 DANKWOORD Dankwoord Dit doctoraat is het resultaat van vier jaar wetenschappelijk onderzoek aan de Sectie Mariene Biologie van de Universiteit Gent, in samewerking met de polytechnische universiteit van Guayaquil (ESPOL) in Ecuador. Het schrijven van een doctoraat, maar vooral het voorafgaande onderzoek, is een proces dat niet alleen de inzet van de doctorandus vergt, maar evenzeer de hulp, steun en motivatie van een hele reeks mensen, die ik bij deze dan ook graag in de bloemetjes wil zetten. Allereerst wil ik mijn promotor, Prof. Dr. Magda Vincx, bedanken om mij de kans te geven dit project te starten. Hoewel het een weg was met vele obstakels, is ze mij altijd blijven steunen. Haar rol in het effenen van de paden in Ecuador was zonder meer cruciaal, en ik heb mij er regelmatig over verbaasd hoe de naam La doctora Vincx deuren kon openen die anders zeker gesloten zouden blijven. De dagelijkse begeleiding lag in goeie handen van Dr. Steven Degraer, mijn co-promotor. Van het begin tot het einde van mijn doctoraat kon ik altijd rekenen op zijn wetenschappelijke creativiteit en zijn kracht om mensen te motiveren. Daarnaast speelde hij ook de soms ondankbare rol van ‘advocaat van de duivel’ bij het doornemen van mijn schrijfsels. Tenslotte wil ik Steven ook bedanken voor de twee korte trips die we samen in Ecuador gemaakt hebben, en die cruciaal gebleken zijn in de onderzoekslijn die ik uiteindelijk gevolgd heb. When in Ecuador, I could always count on the full support of the Oceanography and Marine Sciences departement of ESPOL and the aquaculture institute CENAIM in San Pedro. In ESPOL, I was always welcome in the lab of Dr. Maria Pilar de Cornejo. Dr. Jorge Calderon, Dr. Sam Stern, Ing. Andrés Pedrazzolli and Dr. Stanislaus Sonnenholzner guaranteed me total freedom in my work at CENAIM, even if many last-minute changes had to be arranged. I know it always seemed weird to the people of CENAIM that someone could do scientific research with nothing but a spade, a bucket and a frame, but I hope this PhD proves it can. Of course, I also want to thank my Ecuadorian colleagues, Sonnia, Luis, Veronica, Alba and Mare, for the happy times in Ecuador, and their help in translating my abstracts, summaries and talks into Spanish! Special thanks go to Galo, without whom I would never have survived the many road trips and sampling campaigns. DANKWOORD Uiteraard bepaalt de omgeving waarin je werkt voor een groot stuk de werkvreugde. Ik wil dan ook heel graag mijn collega’s en ex-collega’s van de Sectie Mariene Biologie bedanken voor een creëen van een gemoedelijke maar wetenschappelijk stimulerende sfeer. Ik denk dat ik bij zowat elke collega wel eens (of soms vele malen) ben langsgeweest met één of andere vraag. In het bijzonder zou ik graag Maarten, Marleen en Nancy vermelden voor hun hulp bij allerhande problemen die ik op mijn weg ben tegengekomen. De – voor mij althans – ingewikkelde wiskundige modellen van hoofdstuk zes werden opgebouwd door Bea. Mijn bureaugenoten, Jan, Veronique en Veronique wil ik bedanken voor de vele leuke babbels tussen het werk door. Ook mijn collega-strandjutters, Tom en Jeroen, verdienen een plaatsje in dit dankwoord, voor de vaak levendige discussies over beach indices en staalnameprotocols. De soms lange dagen werden een stuk lichter gemaakt door de aangename lunchpauzes met de resto-gangers. Voor allerhande praktische beslommeringen kon ik altijd terecht bij Isolde, Annick en Guy. Voor de verwerking van de sedimentstalen en hulp bij het bepalen van omgevingsvariabelen kon ik dan weer rekenen op Danielle, Bart en Dirk. Jurgen en Yves zorgden ervoor dat ik de swashomgeving kon nabootsen in het laboratorium door de swash rig te ontwerpen en te bouwen. Veel van het zware staalnamewerk en het uitsorteren van de stalen is gebeurd door mijn thesisstudenten, Eveline, Pieter, An, Lien en Leen, waarvoor mijn oprechte dank. Jullie verblijven in Ecuador zorgden voor een aangename afwisseling, en ik hou er heel mooie herinneringen aan over! Bij de verwerking van de soms moeilijk interpreteerbare gegevens heb ik veel hulp gehad aan de bezoekjes bij Frederik Hendrickx en Stijn Vansteelandt alsook aan de mailconversaties met Andrew Short en Dave Schoeman. De juryleden van mijn doctoraat hebben dankzij hun opmerkingen en suggesties geleid tot een beter eindproduct. In het dagelijks leven hebben een hele reeks mensen, elk op hun manier, een steentje bijgedragen tot dit werk, vaak zonder het te beseffen. Daarom, merci ‘foute vrienden’, ‘fotovrienden’, ‘biologievrienden’, ‘schaatsvrienden’, familie en schoonfamilie. In Ecuador, I had great nights out, and lots of diverting chats with mis amigos/-as Ecuatorianos/-as. DANKWOORD Mijn eerste stappen in de wondere wereld van de biologie heb ik gezet samen met mijn moeder en Oom Willem. De ervaringen die ik tijdens onze vele uitstapjes in binnen- en buitenland heb opgedaan, koester ik voor de rest van mijn leven. Mam, het is dankzij jouw steun en inzet dat ik heb kunnen en mogen doen wat ik graag doe. Vrijheid en zelfstandigheid zijn misschien wel de mooiste dingen die je in een opvoeding kan meekrijgen.
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