Evolution of Colony Characteristics in The Harvester Ant Genus Pogonomyrmex Dissertation zur Erlangung des naturwissenschaftlichen Doktorgrades der Bayerischen Julius-Maximilians-Universität Würzburg vorgelegt von Christoph Strehl Nürnberg Würzburg 2005 - 2 - - 3 - Eingereicht am: ......................................................................................................... Mitglieder der Prüfungskommission: Vorsitzender: ............................................................................................................. Gutachter : ................................................................................................................. Gutachter : ................................................................................................................. Tag des Promotionskolloquiums: .............................................................................. Doktorurkunde ausgehändigt am: ............................................................................. - 4 - - 5 - 1. Index 1. Index................................................................................................................. 5 2. General Introduction and Thesis Outline....................................................... 7 1.1 The characteristics of an ant colony...................................................... 8 1.2 Relatedness as a major component driving the evolution of colony characteristics.................................................................................................10 1.3 The evolution of polyandry: ultimate and proximate causes ................11 1.4 Polygyny and slavery..........................................................................13 1.5 Morphological polymorphisms between workers and queens ..............14 1.6 Introduction to the study genus Pogonomyrmex ..................................16 1.7 Outline of the thesis ............................................................................26 1.8 Publications ........................................................................................28 1.9 Literature ............................................................................................28 3. Determinants of Intracolonial Relatedness in Pogonomyrmex rugosus (Hymenoptera; Formicidae): Mating Frequency and Brood Raids .................... 41 3.1 Abstract ..............................................................................................41 3.2 Introduction ........................................................................................41 3.3 Materials and methods ........................................................................43 3.4 Results................................................................................................46 3.5 Discussion ..........................................................................................48 3.6 Acknowledgements.............................................................................53 3.7 Tables .................................................................................................53 3.8 Figures................................................................................................57 3.9 References ..........................................................................................58 4. Extremely High Mating Frequency in the Florida Harvester Ant (Pogonomyrmex badius)......................................................................................... 62 4.1 Abstract ..............................................................................................62 4.2 Introduction ........................................................................................62 4.3 Methods..............................................................................................66 4.4 Results................................................................................................70 4.5 Discussion ..........................................................................................72 4.6 Acknowledgements.............................................................................76 4.7 Appendix ............................................................................................77 4.8 References ..........................................................................................88 5. A Genetic Component in the Determination of Worker Polymorphism in the Florida Harvester ant Pogonomyrmex badius ................................................ 92 5.1 Summary ............................................................................................92 5.2 Introduction ........................................................................................92 5.3 Methods..............................................................................................94 5.4 Results................................................................................................95 5.5 Discussion ..........................................................................................95 5.6 Acknowledgments ..............................................................................97 5.7 References ..........................................................................................97 5.8 Tables .................................................................................................99 6. Cladistic Analysis of Paleo-Island Populations of the Florida Harvester Ant (Hymenoptera: Formicidae) Based Upon Divergence of Mitochondrial DNA Sequences.................................................................................................... 103 6.1 Abstract ............................................................................................103 6.2 Resumen...........................................................................................103 - 6 - 6.3 Introduction ......................................................................................104 6.4 Materials and Methods......................................................................105 6.5 Results..............................................................................................109 6.6 Discussion ........................................................................................113 6.7 Acknowledgments ............................................................................114 6.8 References cited................................................................................115 7. Colony structure and morphometrics in the queen dimorphic harvester ant, Pogonomyrmex pima..................................................................................... 118 7.1 Summary. .........................................................................................118 7.2 Introduction ......................................................................................118 7.3 Methods............................................................................................120 7.4 Results..............................................................................................122 7.5 Discussion ........................................................................................125 7.6 Acknowledgments ............................................................................129 7.7 References ........................................................................................130 7.8 Table.................................................................................................133 7.9 Figures..............................................................................................135 8. Colony Structure and Sociogenetics of the Queen Dimorphic Harvester Ant, Pogonomyrmex (Ephebomyrmex) pima ....................................................... 137 8.1 Abstract ............................................................................................137 8.2 Introduction ......................................................................................137 8.3 Materials and Methods......................................................................140 8.4 Results..............................................................................................144 8.5 Discussion ........................................................................................147 8.6 Acknowledgements...........................................................................153 8.7 Tables ...............................................................................................154 8.8 Figures..............................................................................................158 8.9 References ........................................................................................159 9. Phylogeny of the Harvester Ant Genus Pogonomyrmex (Hymenoptera: Formicidae) in North-America and its relation to South-American Pogonomyrmex species ........................................................................................ 165 9.1 Abstract ............................................................................................165 9.2 Introduction ......................................................................................165 9.3 Material and Methods .......................................................................168 9.4 Results..............................................................................................170 9.5 Discussion ........................................................................................174 9.6 Acknowledgments ............................................................................177