DISSERTATION Titel der Dissertation Cues and Signals Used For Orientation and Communication in the Superorganismic Meliponine Bees (Hymenoptera, APIDAE, Meliponini) angestrebter akademischer Grad Doktor/in der Naturwissenschaften (Dr. rer.nat.) Verfasserin / Verfasser: Dirk Louis Schorkopf Matrikel-Nummer: 9607465 Dissertationsgebiet (lt. Stu- A 091 439 Dr.-Studium der Naturwissenschaften Zoologie (Stzw) dienblatt): Betreuerin / Betreuer: Univ.-Prof. Dr. Friedrich G. Barth Wien, im Jänner 2010 Formular Nr.: A.04 Cues and Signals Used For Orientation and Communication in the Superorganismic Meliponine Bees (Hymenoptera, APIDAE, Meliponini) a dissertation by Dirk Louis P. Schorkopf © Dirk Louis P. Schorkopf University of Vienna February 2010 Cover illustration (if provided with this copy) shows the surface of a Trigona spinipes nest. Illustration under the title was kindly presented by Martina Schorkopf. Dedicated to all my teachers not all of them professors not all of them human beings Thanking them for their guidance by providing signals and cues in so many ways * * Cited from: Sextus Empiricus (“adversos mathematicos), cited in Hermmann, D. (1957): Die Fragmente der Vorsokratiker. Hamburg: Rowohlt. CONTENTS Preface __1 I. General introduction __9 II. Identification of signals and cues inducing defensive and aggressive behaviour on the intra‐ and intersuperorganismic level (J Exp Biol 2009) _27 III. A pheromone to coordinate foraging and the exploitation of highly profitable food sources (Proc R Soc B 2007) 75 IV. A new view of the importance of pheromone paths in meliponines OR Revisiting a 50 year old experiment (J Exp 95 Biol, submitted 2009) V. Some more aspects of meliponine communication and orientation in coauthored papers 121 VI. Unpublished data and observations 167 VII. Concluding remarks and summary 185 References 203 Preface “Besides the dissertation committee nobody will read a thesis anyway” This is what I constantly hear from close or distant friends in research – some of them supervising thesis‐writing students themselves. Although I fear that there is some truth in the saying, I nevertheless believe that a thesis represents a good opportunity to revisit the work done and published during several long years of total dedication to a subject of scientific interest. For the readers, however few, I briefly list what can be expected in this thesis. Chapter I is a general introduction and provides some selected background information relevant to the research interests of this thesis. It also provides a list of a few important definitions as well as the main questions asked. All readers already familiar with the field of research of this thesis may skip this chapter, except those who wish to have some idea about the particular concepts I had in mind when working for and writing up the thesis. Chapters II and III consist of papers already published. For those who already know the content, some notes and the appendix may provide more information. Chapter IV presents a paper currently under revision. 1 Chapter V summarizes and comments coauthored papers which appeared while I was working for my thesis and are strongly related to it. Chapter VI contains some unpublished data and observations considering chapters II to IV. Chapter VII finally presents the answers to the questions asked at the beginning (chapter I), some general conclusions and a summary, followed by the references. ACKNOWLEDGEMENTS „Der beste Dank ist, wenn alles gut geht“1 [„Best acknowledgement of gratitude is achieved when everything turns out well“] This thesis would not have been realized without the help and support by a numerous number of people worldwide. Main financial support came from a FWF2 project (P17530) granted to my thesis supervisor Friedrich G. Barth. Professor Barth recruited me for the latter project in the end of 2004, presented me unsolicited with a brand new GC‐FID on my birthday in 2005 and renewed my contract no less than 3 times 1 From: Haruki Murakami (2006) Kafka am Strand. München: Random House. [German translation by Ursula Gräfe of “Umibe No Kafuka” published in 2002 by Shinchosha, Tokyo] 2 „Förderung Wissenschaftlicher Forschung”, Austria 2 before the project had to finally end in the beginning of 2008. Rarely somebody was able to drive me to the boarders of my resources and abilities as he did. He also critically read this thesis and significantly (Pchi‐square test < 0.001) improved the style and logical thread of my hitherto written manuscripts. Without him I wouldn’t be where I am now. My truly loving and understanding wife supported me in so many ways and provided the vitamins, catalysts and motivation needed to survive many difficult and challenging events such as unemployment and a marathon. My chummy colleagues Michael Hrncir, Stefan Jarau and Veronika Schmidt contributed greatly to the success of my career. Michael would would not even desert me when I needed him to work wiw with not less than six hives simultan‐ eously (when I write simultaneously, I really mean simultaneously). Stefan introduced me to the, sometimes quite mythical world of chemical ecology and gener‐ously ously shared his knowledge. Veronika was our most mostmaccomplished organization talent, did share knowledge, humorous tales and food. Austria gained one of its best scientific journalists, but lost one of ist most promising bee researchers. Clemens Florian Schaber was an ever‐helpful‐never‐failing‐ fellow, gradually developing or unveiling himself as a true friend in either of the departments3 we both have survived since the beginning of our careers. Linde Morawetz helped me to realize a considerable number of pheromone path experiments, some of them requiring a great deal of enthusiasm and dedi‐ cation, not to mention a seemingly unbreakable loyalty to some of my ventures and last minute experiments. For chemistry I highly acknowledge both Robert Twele and Wittko Francke. Without their expertise my work would never have reached its scientific impact. In addition I want to thank for their hospitality during my stay in Hamburg. Izabel Cristina Casanova Turatti and Norberto P. Lopes most 3 Firstly “Neurobiology division of the Institute of Zoology”, followed by “Department of Neurobiology and Behavioural Sciences”, then “Department of Neurobiology and Cognition Research”; today called “Department of Neurobiology”. 3 generously entrusted me their GC‐MS in Ribeirão Preto (muito obrigado por a confianca!). Ronaldo Zucchi always welcomed us to stay in his department and to use his facilities. A person whom we could always rely on and whom we prize. I dare say: no Austrian bee researcher in Ribeirão Preto in the last decade without his help... To realize the experiments at Piracicaba I thank Roberto Antonio Zucchi for introducing me to José Maurício S. Bento, who made his laboratory facilities available to us. I am very grateful for the opportunities of research at the ESALQ in Piracicaba. Sidnei Mateus and Weyder Santana introduced me to several species of meliponine bees and also showed me how to handle them. Many colonies important for this thesis were only available due to S. Mateus’ generosity and commitment. True Brazilian Hospitality was granted by Fabio Nascimento and his wife Ivelize during my stay at Aracaju. In addition I want to thank Fabio for constantly helping me to realize my ambitious research plans whilst staying at the University Federal de Sergipe and for patiently waiting for the results to be published (by my honour – they will!). Without the bee keeper Erinaldo’s generous offer to borrow 12 of his most precious colonies, however, I would not have gone far. I had some of the most wonderful insights to the behaviour of meliponine bees whilst working with his colonies. Paulo Nogueira Neto permitted me to work at his Fazenda Aretuzina. His discussions at lunch and dinner and his generous hospitality will never be forgotten. Antonio Gilberto da Silva, Izaura Pereira Nunes, Paulo Hernesto Nunes and Artur Luiz Souto were of great support. Vera Imperatriz Fonseca and Christiano Menezes established the contacts for my works at the Fazenda. They also kindly permitted me to work with some of their colonies at the end of my last stay in Ribeirão Preto. Luci Rolandi Bego supported me with access to meliponine colonies and laboratory facilities. Lionel Conçalves provided some rare literature and advice. Remy Vandame and Daniel Sánchez Guillén kindly enabled me to work at ECOSUR in Tapachula, Mexico. Leopoldo Cruz‐ López generously allowed me to use the GC‐EAD at his laboratory. Daniel Guillén Sanchez presented us with a colony 4 of Scaptotrigona mexicana. Guillermo Ibarra borrowed his cold source lights during my stay. In addition I want to thank Miguel Guzmán‐Diaz, Adolfo León, Omar Arguello, Bernhard Kraus and the students at the laboratory of “Abejas de Chiapas”. In Mexico I also want to thank José Javier Quezada Euán, José Humberto Moo Valle, Jorge Àngel González Acereto and José Chavier de Araujo Freitas. At the Department of Neurobiology I thank Maria Wieser, Margit Kainerstorfer, Elisabeth Fritz‐Palanck, Lisa Fenk, Rudi Machan, Andrzej Szpetkowski, Miro Dragasev, Walter Witek, Simon Tragust, Thomas Hoinkes, Christian Klopsch and Rainer Müllan. For support and advice during difficult times I kindly thank all of my family, Elisabeth Zimmermann, Werner Cekal, Britta Grillitsch, Antonia Vieira, Judith Reinhard and Martina Jordan. Helping hands in Brazil to realize experiments or to keep them going were Ana Carolina Roselino, Angel Roberto Barchuk, Solange Santos, Sofia Mc Cabe, Túlio Nunes, Fabio Nascimento, Sergio Andena, Silvio Morato, Marcelo dos Santos Pádua Diniz, Newton Cavalcanti de Noronha Jr., Eva Stangler, Geusa de Freitas, Jairo de Souza, A.G.C. Signoretti, H.R. dos Santos and J.C.R. Castilho. In Brazil I also want to thank Anneliese and Michel Effenberger & family, Marcí Wagner & family, Angela Alcon & family, Christine and Ariel Dlouhy & family. Like all who work on a dissertation, I frequently had been confronted with difficulties to get access to some thesis or (un‐) published work.