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Little Things Become Big Little Things Become Big Drivers and impacts of benthic cyanobacterial blooms on coral reefs Dissertation zur Erlangung des akademischen Grades eines Doktors der Naturwissenschaften ‐ Dr. rer. nat. ‐ Am Fachbereich 2 (Biologie/ Chemie) der Universität Bremen Vorgelegt von Hannah Juliane Brocke Bremen 2014 I II 1. Gutachter: Prof. Dr. Kai Bischof 2. Gutachter: Associate Prof. Maggy M. Nugues Hannah Juliane Brocke1,2,3, Dipl. Biol. 1Max Planck Institute for Marine Microbiology (MPI Bremen) Celsiusstr. 1, 28359 Bremen, Germany 2Leibniz Center for Tropical Marine Ecology (ZMT) Fahrenheitstr. 6, 28359 Bremen, Germany 3 Laboratoire d’Excellence Corail, CRIOBE – USR 3278, CNRS-EPHE-UPVD, 58 avenue Paul Alduy, 66860 Perpignan Cedex, France III IV Dedicated to all optimists who search for solutions and don’t give up trying. “In the long history of humankind (and animal kind, too) those who learned to collaborate and improvise most effectively have prevailed.” Charles Darwin “Every [coral reef] ecosystem I studied is unrecognizably different from when I started. I have a son who is 30, and I used to take him snorkeling on the reefs of Jamaica to show him all the beautiful corals there. I have a daughter who is 17. I can't show her anything but seaweed.” Jeremy Jackson V TABLE OF CONTENTS Table of contents Declaration ................................................................................................................................... IX Acknowledgements ...................................................................................................................... X Zusammenfassung ........................................................................................................................ XIV 1. Introduction .............................................................................................................................. 1 1.1. Overview of cyanobacteria and their life in microbial mats 1.1.1. Cyanobacteria ................................................................................................ 2 1.1.2. Microbial mats – benthic cyanobacteria and their associates ....................... 3 1.2. Coral reefs and the phase shift to benthic algae and cyanobacteria dominated reefs 1.2.1. Coral reef – a highly productive ecosystem in a marine desert ..................... 5 1.2.2. Degradation of coral reefs and the revolution of microbes ........................... 6 1.3. State of the art – benthic cyanobacterial mats on coral reefs 1.3.1. Expansion and causes of cyanobacterial mats ............................................... 7 1.3.2. Ecological relevance of benthic cyanobacteria dominating the reef ............. 9 1.4. Aims and objectives of PhD thesis ............................................................................... 11 1.5. Overview of manuscripts and authors contributions .................................................. 14 2. Manuscript I .............................................................................................................................. 19 3. Manuscript II ............................................................................................................................. 53 4. Manuscript III ............................................................................................................................ 85 5. Manuscript IV ........................................................................................................................... 109 6. Manuscript V ............................................................................................................................ 149 VI TABLE OF CONTENTS 7. Manuscript VI ........................................................................................................................... 181 8. Manuscript VII .......................................................................................................................... 187 9. Synopsis ...... ............................................................................................................................. 193 9.1. Divers and causes of benthic cyanobacterial blooms .................................................. 194 9.2. Impacts of cyanobacterial mats on the reef ecosystem .............................................. 198 9.3. High cyanobacterial mat abundances – a future scenario? ........................................ 200 9.4. Perspectives for future research ................................................................................. 202 10. Literature ................................................................................................................................ 207 VII DECLARATION VIII DECLARATION Declaration Name: Hannah Juliane Brocke Anschrift: St.-Jürgen-Str. 18, 28203 Bremen Erklärung gemäss §6 Abs. 5 der Promotionsordnung der Universität Bremen für die mathematischen, natur- und ingenieurwissenschaftlichen Fachbereiche. Hiermit erkläre ich, dass ich die Doktorarbeit mit dem Titel: Little Things Become Big – Drivers and impacts of benthic cyanobacterial blooms on coral reefs – 1. Selbstständig verfasst und geschrieben habe, 2. keine anderen als die von mir im Text angegebenen Quellen und Hilfsmittel benutzt, 3. die den benutzten Werken wörtlich und inhaltlich entnommenen Stellen als solche kenntlich gemacht habe, und 4. dass es sich bei den von mir abgegebenen Arbeiten um drei identische Exemplare handelt. Ort/Datum Unterschrift IX ACKNOWLEDGMENTS Acknowledgements Vielen herzlichen Dank Kai Bischof for being my Doktorvater and 1st Gutachter. Thank you for always taking time, listening carefully and giving friendly advises. Those were very important to me. Merci beaucoup Maggy Nugues for being my 2nd Gutachter and more importantly my project leader, giving me this PhD position and putting me on track to unravel the mystery of benthic cyanobacteria. Heel hartelijk bedankt Dirk de Beer for being my examiner and having me in your group. Thank you for your helpful advises, idea contributions and giving me all this nice equipment for this project. Vielen Dank Claudio Richter for being my examiner and finding great interest in my work. I am glad I have you in my committee. Thank you very much Peter Mumby and Rosanna Griffith-Mumby for helping us in turbulent times; and you, Peter, for all the great project discussions. Also a big thank you to all FORCE members. It was a great pleasure to meet you all and be part of your group. Heel hartelijk bedankt Petra Visser for all the great discussions and your support. You are definitively a paragon of a great scientist to me. Hartstikke bedankt Mark Vermeij for last minute advises, ideas, support and great discussions. Mascha danki Carlos and Oscar for spending many hours fixing the boat motor with me, giving me advise on broken cars or the latest self-build equipment. I feel that during my PhD project I also received half a mechanics education from you two. A big thank you also to Kristin Marhaver as well as to Ashwin Engelen, Forest Rohwer, Stuart Sandin for fruitful discussions and motivational feedback. Heel erg bedankt Paul Stokkermans for finding solutions for all the administrative problems I was facing on Curaçao. Visa, boat papers, licenses, shipments, sample export… even calling ministers was no problem for you. Vielen Dank Benjamin Müller for a helpful hand and discussions in the field. I also thanks all other people from Carmabi, Art & Nature Inn, especially Gerardine Kuijpers and Shirah, as well as Diveversity who made my stay a great experience on Curaçao. D`akujem moc Lubos for all the great discussions, especially in the first month of my PhD. Thank you also for having a great trust in me that I can understand your mathematics and programs. Danny Ionescu for all your help. I really appreciated that you always took time and gave me הרבה תודה quick, fast and great advices! Raeid Abed for teaching me a lot of writing skills, sharing the same fascination towards مكل ركشلا cyanobacteria, and discussing world politics. I very much enjoyed working with you and going for lunch. Vielen Dank Miriam Weber for your great motivation and taking all the emergency calls from the field. Thank you very much Timothy G. Ferdelman for your time and very helpful advises. X ACKNOWLEDGMENTS Thank you very much Duygu Sevgi Sevilgen, Stefan Heusler, Mar Fernández-Méndez, Saar Sztejrenszus, Mina Bizic, Jon Graf, Anna Behrendt, Judith Klatt, Arjun Chennu,… and all other MPI PhD students for being my PhD buddies. Thank you for showing me programs, DOMS use, calculations, measurements etc., and being there during all ups and downs. Many thanks also to Laura Piepgras, Artur Fink for being in my committee and Ines Stuhldreier for helping me with the abstract translations. Vielen Dank Frank Wenzhoefer, Felix Janssen and Moritz Holtappels for teaching me to work with sediment chambers, the self-made permeability apparatus and accompanied calculations. Vielen Dank Anja Kamp und Peter Stief for the very nice lunch breaks and helpful advises. Vielen Dank liebe TAs, especially from the Microsensor group, but also from the Biogeochemistry and Deep-Sea Ecology and Technology group. Thank you for your time teaching me how to build and work with microsensors, or teaching me fancy machines, such as the SEM, or less fancy machines, such as the CMS analyzer. By mentioning the last, I can also not forget to thanks the IT department for the time- consuming troubleshooting, and last but not least, a big thank you to the MPI workshop, especially to
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