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The Evolution of Sexually Selected Traits

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The Evolution of Sexually Selected Traits The Evolution of Sexually Selected Traits Comparative Morphology and Chemistry of the Cephalic Glands of Beewolves (Hymenoptera, Crabronidae) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Fakultät für Biologie und Vorklinische Medizin der Universität Regensburg vorgelegt von Katharina Weiß aus Amberg im Jahr 2017 The Evolution of Sexually Selected Traits Comparative Morphology and Chemistry of the Cephalic Glands of Beewolves (Hymenoptera, Crabronidae) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Fakultät für Biologie und Vorklinische Medizin der Universität Regensburg vorgelegt von Katharina Weiß aus Amberg im Jahr 2017 Das Promotionsgesuch wurde eingereicht am: Die Arbeit wurde angeleitet von: Prof Dr. Erhard Strohm Unterschrift: I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists entertain, and which I formerly entertained - namely, that each species has been independently created - is erroneous. Charles Darwin, The Origin of Species (1859) Intelligent life on a planet comes of age when it first works out the reaso for its o eistee. […] Liig orgaiss had eisted o earth, without ever knowing why, for over three thousand million years before the truth finally dawned on one of them. His name was Charles Darwin. Richard Dawkins, The Selfish Gene (1976) CONTENTS List of Publications ............................................................................................................ 4 CHAPTER 1: General Introduction .................................................................................. 5 1.1 Sexual selection and the evolution of interspecific variation ...................................... 6 1.1.1 Dari’s uresoled prole: The asetr of seual seletio ..................... 6 1.1.2 The evolution of secondary sexual characters ..................................................... 7 1.1.3 Sexual selection and interspecific variation ......................................................... 8 1.2 Chemical communication and mate choice ................................................................. 8 1.2.1 Pheromones in mate choice ................................................................................ 8 1.2.2 The origin of pheromone diversity ...................................................................... 9 1.3 Exocrine glands in Hymenoptera ................................................................................. 10 1.3.1 The morphology of exocrine glands ..................................................................... 10 1.3.2 Exocrine glands and antimicrobial defense ......................................................... 11 1.3.3 Exocrine glands and sexual communication ........................................................ 12 1.4 The Philanthinae .......................................................................................................... 13 1.4.1 Systematics and geographic distribution ............................................................. 13 1.4.2 General biology.................................................................................................... 14 1.5 Outline of the thesis .................................................................................................... 17 1.6 References ................................................................................................................... 19 CHAPTER 2: Comparative Morphology of the Postpharyngeal Gland in the Philanthinae (Hymenoptera, Crabronidae) and the Evolution of an Antimicrobial Brood Protection Mechanism ...................................................................................... 34 2.1 Summery ..................................................................................................................... 34 2.2 Introduction ................................................................................................................. 35 2.3 Methods ...................................................................................................................... 37 2.3.1 Prey embalming in Philanthus gibbosus .............................................................. 37 2.3.2 Comparative morphology of head glands ............................................................ 38 2.4 Results ......................................................................................................................... 41 2.4.1 Prey embalming in Philanthus gibbosus .............................................................. 41 2.4.2 Comparative morphology of head glands ............................................................ 43 2.5 Discussion .................................................................................................................... 50 2.5.1 Prey embalming in Philanthus gibbosus .............................................................. 50 2.5.2 Comparative morphology of head glands ............................................................ 51 2.5.3 Evolution of prey embalming and complex PPGs ................................................ 53 2.5.4 Conclusion ........................................................................................................... 55 2.6 Supplementary material .............................................................................................. 56 2.6.1 Prey embalming in Philanthus gibbosus .............................................................. 56 2.6.2 Comparative morphology of head glands: Coding of character states ................ 59 2.6.3 Supplementary table and figure .......................................................................... 63 2.7 References ................................................................................................................... 66 CHAPTER 3: Evolution and phylogenetic Distribution of antimicrobial Brood Defense in Beewolves (Hymenoptera, Crabronidae): Evidence from Postpharyngeal Gland Chemistry ......................................................................................................................... 71 3.1 Summery ..................................................................................................................... 71 3.2 Introduction ................................................................................................................. 72 3.3 Methods ...................................................................................................................... 74 3.3.1 Specimens ............................................................................................................ 74 3.3.2 Chemical analysis ................................................................................................. 74 3.3.3 Data analysis ........................................................................................................ 77 3.4 Results ......................................................................................................................... 79 3.5 Discussion .................................................................................................................... 84 3.6 References ................................................................................................................... 90 CHAPTER 4: Does the Morphology of the Postpharyngeal Gland reflect potential functional Changes in Bee- vs. Beetle-hunting Species of the Digger Wasp Genus Cerceris (Hymenoptera, Crabronidae)? ...................................................................... 95 4.1 Summery ..................................................................................................................... 95 4.2 Introduction ................................................................................................................. 96 4.3 Methods ...................................................................................................................... 98 4.3.1 Specimens ............................................................................................................ 98 4.3.2 Histology .............................................................................................................. 98 4.3.3 3D-Reconstruction ............................................................................................... 99 4.3.4 Comparison of gland volumes ............................................................................. 99 4.4 Results ......................................................................................................................... 100 4.4.1 Overall morphology of head glands ..................................................................... 100 4.4.2 Light-microscopy ................................................................................................. 101 4.4.3 Comparison of gland volumes ............................................................................. 103 4.5 Discussion .................................................................................................................... 104 4.6 Conclusion ................................................................................................................... 107 4.7 References ................................................................................................................... 108 CHAPTER 5: Sexual Selection and the Evolution of Male Pheromone Glands in philanthine Wasps (Hymenoptera, Crabronidae) ......................................................
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