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Phd Thesis, University of Würzburg Protective bacteria and attractive pheromones Symbiosis and chemical communication in beewolves (Philanthus spp., Hymenoptera, Crabronidae) Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universität Würzburg vorgelegt von Martin Kaltenpoth aus Hagen Würzburg 2006 Eingereicht am: ………………………………………………………………….…….………..… Mitglieder der Promotionskommission: Vorsitzender: Prof. Dr. Martin J. Müller Gutachter: Prof. Dr. Erhard Strohm Gutachter: Prof. Dr. Roy Gross Tag des Promotionskolloquiums: ………………………………………………….………….….. Doktorurkunde augehändigt am: …………………………………………….…………………… 2 “We are symbionts on a symbiotic planet, and if we care to, we can find symbiosis everywhere.” Lynn Margulis, “Symbiotic Planet” (1998) “There is a theory which states that if ever anyone discovers exactly what the universe is for and why it is here, it will instantly disappear and be replaced by something even more bizarre and inexplicable. There is another theory which states that this has already happened.” Douglas Adams, “The Restaurant at the End of the Universe” (1980) 3 4 CONTENTS LIST OF PUBLICATIONS..................................................................................................... 7 CHAPTER 1: GENERAL INTRODUCTION ........................................................................... 9 1.1 Symbiosis ......................................................................................................................... 9 1.1.1 Insect-bacteria symbiosis...................................................................................... 10 1.1.2 Nutritional interactions ......................................................................................... 11 1.1.3 Defensive symbioses............................................................................................. 13 1.2 Chemical communication and mate choice.................................................................... 15 1.2.1 Sexual selection and mate choice.......................................................................... 15 1.2.2 Pheromones and mate choice................................................................................ 16 1.3 Biology of Philanthus triangulum (Hymenoptera, Crabronidae)................................... 17 1.3.1 Systematic position and distribution of the genus Philanthus .............................. 17 1.3.2 Behavior of female European beewolves.............................................................. 18 1.3.3 Behavior of male European beewolves................................................................. 20 1.4 Outline of the thesis........................................................................................................ 22 1.4.1 Symbiotic microorganisms in Philanthus triangulum .......................................... 22 1.4.2 The male marking pheromone as a potential cue for female choice..................... 23 1.5 References ...................................................................................................................... 24 CHAPTER 2: SYMBIOTIC BACTERIA PROTECT WASP LARVAE FROM FUNGAL INFESTATION................................................................................................................ 35 2.1 Summary......................................................................................................................... 35 2.2 Results and Discussion................................................................................................... 36 2.3 Methods .......................................................................................................................... 42 2.3.1 PCR and sequencing ............................................................................................. 42 2.3.2 Phylogenetic analysis............................................................................................ 42 2.3.3 FISH...................................................................................................................... 42 2.3.4 Fungal infestation bioassays with beewolf cocoons ............................................. 43 2.3.5 Survival of larvae with and without white substance............................................ 43 2.4 References ...................................................................................................................... 44 CHAPTER 3: BAKTERIEN SCHÜTZEN WESPEN-NACHWUCHS VOR PILZBEFALL .......... 47 3.1 Zusammenfassung .......................................................................................................... 47 3.2 Einleitung ....................................................................................................................... 47 3.3 Ergebnisse und Diskussion............................................................................................. 48 3.4 Literaturverzeichnis........................................................................................................ 51 CHAPTER 4: MORPHOLOGY AND ULTRASTRUCTURE OF A BACTERIA CULTIVATION ORGAN:THE ANTENNAL GLANDS OF FEMALE EUROPEAN BEEWOLVES, PHILANTHUS TRIANGULUM (HYMENOPTERA,CRABRONIDAE) ...................................53 4.1 Summary......................................................................................................................... 53 4.2 Introduction..................................................................................................................... 54 4.3 Materials and Methods.................................................................................................... 55 4.3.1 Specimens.............................................................................................................. 55 4.3.2 Semithin sections and 3D-reconstruction.............................................................. 55 4.3.3 Electron microscopy.............................................................................................. 56 4.4 Results............................................................................................................................. 56 4.4.1 Overall morphology .............................................................................................. 56 4.4.2 Ultrastructure......................................................................................................... 61 4.4.3 Reservoir contents................................................................................................. 63 4.5 Discussion....................................................................................................................... 64 4.6 References....................................................................................................................... 66 CHAPTER 5: ‘CANDIDATUS STREPTOMYCES PHILANTHI’, AN ENDOSYMBIOTIC STREPTOMYCETE IN THE ANTENNAE OF PHILANTHUS DIGGER WASPS.......................69 5.1 Summary......................................................................................................................... 69 5.2 Introduction..................................................................................................................... 70 5.3 Methods .......................................................................................................................... 71 5.3.1 Specimens.............................................................................................................. 71 5.3.2 Electron microscopy.............................................................................................. 71 5.3.3 DNA extraction, PCR and sequencing.................................................................. 72 5.3.4 Fluorescence in situ hybridization (FISH) ............................................................ 73 5.3.5 Phylogenetic analysis ............................................................................................ 73 5.4 Results............................................................................................................................. 73 5.4.1 Localization of endosymbionts ............................................................................. 73 5.4.2 Distribution of symbionts among philanthine wasps ............................................ 76 5.4.3 Phylogenetic position of ‘Candidatus Streptomyces philanthi’............................ 78 5.5 Discussion....................................................................................................................... 78 5.5.1 Description of ‘Candidatus Streptomyces philanthi’............................................ 80 5.6 Appendix......................................................................................................................... 81 5.6.1 Attempts to cultivate ‘Candidatus Streptomyces philanthi’.................................. 81 5.6.2 Media Formulations .............................................................................................. 83 5.7 References....................................................................................................................... 84 2 CHAPTER 6: CHEMICAL COMPOSITION OF THE ANTENNAL GLAND SECRETION OF FEMALE EUROPEAN BEEWOLVES, PHILANTHUS TRIANGULUM (HYMENOPTERA, CRABRONIDAE)............................................................................................................ 87 6.1 Summary......................................................................................................................... 87 6.2 Introduction ...................................................................................................................
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