Resin collection and use in stingless bees Wie stachellose Bienen Pflanzenharze sammeln und nutzen Doctoral Thesis for a doctoral degree at the Graduate School of Life Sciences, Julius-Maximilians-Universität Würzburg Section: Integrative Biology Sara Diana Leonhardt born in Stuttgart Würzburg 2010 The Bee His labor is a chant, His idleness a tune; Oh, for the bee’s experience Of clovers and of noon! Emily Dickinson – Poems XV Submitted on/ eingereicht am: ……………………………………………………………... Members of PhD student’s committee/ Promotionskomitee: Chairperson: ……………………….................. Primary Supervisor: PD Dr. Nico Blüthgen (Würzburg) Supervisor (second): Prof. Dr. James Nieh (San Diego) Supervisor (third): Prof. Dr. Jürgen Tautz (Würzburg) Date of Public Defense/Verteidigung am: …………………………………….. Date of Receipt of Certificates/ Zeugnis erhalten am: …………………….. 2 Affidavit/Eidesstattliche Erklärung according §4 Abs. 3 Ziff. 3, 5 und 8 of the „Promotionsordnung der Julius-Maximilians-Universität Würzburg“ I hereby declare that my thesis entitled “Resin collection and use in stingless bees” is the result of my own work. I did not receive any help or support from commercial consultants. All sources and / or materials applied are listed and specified in the thesis. Furthermore, I verify that this thesis has not yet been submitted as part of another examination process neither in identical nor in similar form. Würzburg ……………………………………………………………………………... 3 Acknowledgements/ Danksagung So long, and thanks for all the bees… About ten years ago, while still being a scholar at the “Gymnasium”, I was, for the first time, faced with the analysis of mass-spectra, in my chemistry class. At that precise moment, I was hoping for never having to do something like this again in my whole life. It is, however, due to my great and very motivating chemistry-teacher, Frank Pahnke, that I discovered my passion for chemical ecology and that, years later, I spent a huge part of my PhD research with “Pikologie”, to say: analyzing and interpreting mass spectra. For sure, I would not have had such a fabulous time of tropical bee research without a great number of very special people: Nico Blüthgen once saved me from studying frogs and got me intrigued by the tiny smelly stingless bees. Moreover, on one of our many downpour-walks in the beautiful rainforest of Borneo, he started to support my crazy idea of studying resin collection in these bees. He has since then been a fabulous and incredibly encouraging advisor who was always there and open for discussing new as well as old ideas, for sharing a class of wine when (once again) a paper got rejected and for listening no matter what the subject was. Thomas Schmitt helped me not only to survive endless hours of data analysis, but also one of the most horrible thunderstorm flights in Borneo. His optimism was always contagious and encouraging, his knowledge on chemical ecology and his willingness to fight the self-doubt of a young scientist apparently endless. He is not listed as a formal advisor of my PhD thesis, but he is the officially missing one I would never want to miss. Jürgen Tautz and James Nieh both agreed to supervise my PhD project. They strongly encouraged my often ambitious plans and contributed with fruitful ideas, long discussions and very helpful comments. Linda-Maria Jung, Sonja Zeilhöfer and Sven Form all had their load of stingless bee research. They strongly contributed to the overall success of this project and were great company in Borneo, Würzburg and Freiburg. Gunnar Knobloch, Sonja Zeilhöfer and Sven Kurz were the ones working behind the curtain by spending endless hours analyzing mass-spectra. Without their help, I would most likely have needed an additional year to finish the bee “Pikologie”. Florian Menzel is a little bit of everything: the best lab-made I could think of, the one person who knew statistic solutions when I had longtime given up, co-founder of “Thunderstorm” and a fabulous Tiong-made, a co-fighter against the dark sides of Danum, more than a colleague and a really good friend. 4 Robert Junker brought me to terms when I started complaining about things that were hardly worth complaining. He was always up for a bottle of wine, long lasting discussions, helpful comments or just a disarming laughter (Aloha). Mareike Wurdak, Jan Buellesbach and all the people from the Freiburg Lab were great company and help whenever I came to the Breisgau to do some lab work in Freiburg. In Freiburg, Wolf and Renate Lindner provided me with the coziest place to stay I could think of – as well as with great discussions, open fires and extremely tasteful wines. My time in Borneo would not have been as great and successful as it was without a whole lot of people: the company of Martin Stenflo, Kerstin Poltz, Florian Menzel, Wong Siew Te and Waipak Ng made life in Danum and Sepilok an unforgettable event and long lasting memory. Arthur Chung, Robert Ong, Rory Walsh, Glen Reynolds and particularly Chey Vun Khen supported me and my project in every possible way. Mike (Bernadus Bala Ola) was the best research assistant, Bahasa Melayu teacher, Karaoke singer, badminton player, birder and forest companion I could have wished for. Meeting Jonathan Tan and his family was a special event and gave me insight not only into the life of a fascinating Chinese Malaysian family, but also into the unknown and distrusted world of oil palm plantations. Shoko Sakai, Takao Itioka and all researchers from the Japanese Lab in LHN enabled me to expand my work from Sabah to Sarawak and meet even more incredibly helpful and interesting people. I finally have to thank the British Royal Society as well as the Economic Planning Unit (EPU) in Kuala Lumpur for giving me permission to do fieldwork in Malaysia. Terima kasih banya. When I expanded my studies to Australia, I would not have gone far without the help of Tim Heard who knew not only every stingless bee species, but also every stingless bee keeper and provided me with a place to stay, bee specimens en masse, the first stingless bee honey I ever tasted and lots of advice. Janine and Russell Zabel, Peter Davenport, Ben Oldroyd, Bob Lawn as well as Lewis and Charly Roberts kindly allowed me to collect bees from their backyard hives and assisted me in every possible way. Helen Wallace did not only provide funding for part of the trip, but also knew where to find bees and trees, how to cook a great green Curry and how to find the Roberts brothers, rendering my work in Australia above all a great adventure. Including stingless bees from the New World into my work was only possible thanks to Stefan Jarau and Christian Reichle who shared their connections and knowledge on Costa Rican stingless bees and stingless bee people (“abajeros”) with me. In Costa Rica, Eduardo 5 Herrera and his family provided me with a second home, a second family, a bike, easy accessible bee hives, great company and even more people I am glad to have met. Back in Würzburg, Dieter Mahsberg is one of the first people to mention, because I would not have started this dissertation without his help and encouragement! Funding for my dissertation was provided by the Graduate School of Life Sciences (GSLS) of Würzburg. Here, Gabriele Blum-Oehler, Liane Lichtlein, Stefan Schröder-Köhne, Susanne Fischer and Rose Liebert helped me finding a way through the maze of German Academic red tape. Xaver Kober was always there for me, tolerated all my moods, supported all my ideas, waited for me when I spent months away in the field, and, most of all, gave me his unconditional support and his love! Ulrike Kober immediately accepted me in her family and supported me with food, clothes and shelter, encouraging words, open arms and an always open ear. There is no one else like my Mama – Ursula Leonhardt! A best friend, a great women and a foothold I could always and ever count on. Ich habe Dich wahnsinnig lieb! My Papa, Hans-Jörg Leonhardt, is a biologist himself and without doubt one of my earliest reasons to become one as well. He enthusiastically taught me the name of all kind of birds and bees and – together with my Mama – got me intrigued by the beauty of nature at a time where I could barely walk or talk. ‘Blessed’ with two occasionally stubborn daughters, he might not always have been the luckiest man, but he is the best father I can imagine! My sister and best friend, Lena Leonhardt, was the one who accompanied me on my trip to Australia. She managed to be a great field assistant and companion, while holding a film camera on her right arm in plaster. Thanks to her, this trip and life in general became even more special. And there is no one else I’d rather call. Ganz besonders möchte ich meiner Oma, Ilse Meyreiß, danken, die trotz der zahlreichen Krisen in ihrem Leben und ihren nun 84 Jahren immer für mich da war. Brauchte ich Ruhe, gab und gibt es keinen schöneren Platz auf dieser Erde, als das Aulatal! Thanks to Katharina Schwarz, Steffen Wick, Claudia Blüm, Elizabeth Derryberry, Dylan Burge and Lena Leonhardt I have learned that true friends are rare, but one of the most wonderful things in this world. And… last but not least I have to thank my for-legged antidote, Miko, just for seeing the world the way he does. 6 Thesis is based on the following manuscripts: (1) Leonhardt SD, Zeilhofer S, Blüthgen N, Schmitt T (under review, Chemical Senses) Stingless bees use terpenes as olfactory cues to find resin sources.