Subtidal Rocky Reef Communities Galapagos
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_____________________________________________________________________________________ Small-scale Differences in Tropical Subtidal Rocky Reef Communities of Floreana Island, Galápagos Annika Krutwa Doctoral thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Natural Sciences (Dr. rer. nat.) to Faculty 2 (Biology and Chemistry), University of Bremen Bremen, 30 July 2014 Charles Darwin Foundation The present study was conducted at the Charles Darwin Foundation in Puerto Ayora, Galápagos (Ecuador) and compiled at the Leibniz Center for Tropical Marine Ecology (ZMT) GmbH (Germany). This thesis was mainly funded by the German Academic Exchange Service (DAAD) through a research grant for doctoral candidates and the Charles Darwin Foundation. Additional financial support was provided by the ZMT and the German Federal Ministry of Education and Research (BMBF, Grant No. ECU 10/A01). 1. Reviewer: Prof. Dr. Matthias Wolff Leibniz Center for Tropical Marine Ecology (ZMT) GmbH 2. Reviewer: Prof. Dr. Kai Bischof University of Bremen 1. Examiner: Prof. Dr. Thomas Brey Alfred Wegener Institute for Polar and Marine Research (AWI) 2. Examiner: Dr. Claire Reymond Leibniz Center for Tropical Marine Ecology (ZMT) GmbH Erklärung Gemäß §6 der Promotionsordnung der Universität Bremen für die mathematischen, natur- und ingenieurwissenschaftlichen Fachbereiche vom 14. März 2007 erkläre ich, dass die Arbeit mit dem Titel: “Small-scale differences in tropical subtidal rocky reef communities of Floreana Island, Galápagos” 1) ohne unerlaubte Hilfe selbstständig verfasst und geschrieben wurde; 2) keine anderen als die angegeben Quellen und Hilfsmittel benutzt wurden; 3) die den benutzten Werken wörtlich oder inhaltlich entnommenen Stellen als solche kenntlich gemacht wurden; 4) es sich bei den von mir abgegeben Arbeiten um 3 identische Exemplare handelt. Bremen, ____________________________________________ Annika Krutwa II “...What impressionable youngster could resist the images of a mystical moonscape on earth, inhabited by giant tortoises large enough to ride, prehistoric iguanas that share a sea filled with fishes, sea lions and whales, and spreading above it all, a blue equatorial sky alive with the sailing silhouettes and sounds of ocean-going birds on the wing?..” Paul Humann (1993) about Galápagos Table of contents List of Tables ............................................................................................................................. III List of Figures ............................................................................................................................. V Abstract ..................................................................................................................................... VII Zusammenfassung ...................................................................................................................... IX Resumen ..................................................................................................................................... XI Introduction ..................................................................................................................... 1 Chapter 1: Pronounced small-scale differences between Galápagos subtidal rocky reef communities exposed to upwelling and non-upwelling conditions ................... 11 Abstract ...................................................................................................................................... 11 Introduction ................................................................................................................................ 12 Materials and Methods ............................................................................................................... 14 Results ........................................................................................................................................ 17 Discussion .................................................................................................................................. 21 Conclusion ................................................................................................................................. 26 Acknowledgments ...................................................................................................................... 27 References .................................................................................................................................. 27 Appendix .................................................................................................................................... 34 Chapter 2: Natural succession patterns on a tropical rocky reef (Galápagos, Ecuador) under upwelling and non-upwelling conditions ........................................ 41 Abstract ...................................................................................................................................... 41 Introduction ................................................................................................................................ 42 Material and methods ................................................................................................................. 45 Results ........................................................................................................................................ 47 Discussion .................................................................................................................................. 56 Conclusions and Outlook ........................................................................................................... 59 Acknowledgements .................................................................................................................... 60 References .................................................................................................................................. 61 Appendix .................................................................................................................................... 67 Chapter 3: Grazing effects on tropical rocky reef communities exposed to upwelling and non-upwelling conditions in Galápagos (Ecuador) ........................... 69 Abstract ...................................................................................................................................... 69 Introduction ................................................................................................................................ 70 Material and methods ................................................................................................................. 72 Results ........................................................................................................................................ 74 Discussion .................................................................................................................................. 79 Conclusions ................................................................................................................................ 82 Acknowledgements .................................................................................................................... 83 References .................................................................................................................................. 83 Appendix .................................................................................................................................... 89 I Chapter 4: First insights into the food web structure of a tropical subtidal rocky reef community (Galápagos, Ecuador) using the stable isotope approach .............. 91 Abstract ...................................................................................................................................... 91 Introduction ................................................................................................................................ 92 Material and Methods ................................................................................................................. 94 Results ........................................................................................................................................ 95 Discussion .................................................................................................................................. 96 Conclusions .............................................................................................................................. 101 Acknowledgements .................................................................................................................. 101 References ................................................................................................................................ 101 Synthesis ....................................................................................................................... 109 References .................................................................................................................... 121 Acknowledgements ...................................................................................................... 131 II List of Tables Chapter 1 Table 1. Water temperature statistics (in °C). ................................................................ 18 Table 2. Means of univariate measures of fish, mobile macroinvertebrate and fish community structure at LB and TC. ..............................................................................