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Phylogenies and Pheromones

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Phylogenies and Pheromones Phylogenies and pheromones Defensive symbionts, phylogenetic affiliations and olfactory communication in beewolves (Philanthini, Hymenoptera, Crabronidae) Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) der Fakultät für Biologie und Vorklinische Medizin der Universität Regensburg vorgelegt von Kerstin Roeser-Mueller aus München im Jahr 2011 Das Promotionsgesuch wurde eingereicht am 27.9.11 Die Arbeit wurde angeleitet von: Prof. Dr. Erhard Strohm Unterschrift: If you find yourself in a hole, you must stop digging. Will Rogers Für den kleinen Moritz ______________________________________________________________TABLE OF CONTENTS TABLE OF CONTENTS LIST OF PUBLICATIONS ................................................................................................... 3 CHAPTER 1 - GENERAL INTRODUCTION............................................................................ 4 1.1 Symbioses ................................................................................................................ 4 1.2 Sexual selection, pheromones and temperature ............................................................. 6 1.3 The genus Philanthus .................................................................................................. 9 1.4 Outline of the thesis ................................................................................................. 13 CHAPTER 2 - CANDIDATUS STREPTOMYCES PHILANTHI’, AN ENDOSYMBIOTIC STREPTOMYCETE IN THE ANTENNAE OF PHILANTHUS DIGGER WASPS .......................... 15 2.1 Summary ................................................................................................................ 15 2.2 Introduction ............................................................................................................ 15 2.3 Methods.................................................................................................................. 16 2.4 Results ................................................................................................................... 19 2.5 Discussion............................................................................................................... 25 2.6 Description of ‘ Candidatus Streptomyces philanthi’....................................................... 27 2.7 Online supplementary data........................................................................................ 28 CHAPTER 3 - 65 MILLION YEARS OF DEFENSIVE ALLIANCE: MOLECULAR PHYLOGENY OF BEEWOLVES REVEALS THE AGE OF A PROTECTIVE SYMBIOSIS WITH STREPTOMYCES BACTERIA....................................................................................................................... 31 3.1 Abstract.................................................................................................................. 31 3.2 Introduction ............................................................................................................ 32 3.3 Materials and methods.............................................................................................. 33 3.4 Results ................................................................................................................... 38 3.5 Discussion............................................................................................................... 44 3.6 Acknowledgements................................................................................................... 47 3.7 Supplementary material............................................................................................ 48 CHAPTER 4 - LARVAL REARING TEMPERATURE INFLUENCES AMOUNT AND COMPOSITION OF THE MARKING PHEROMONE OF MALE EUROPEAN BEEWOLVES (PHILANTHUS TRIANGULUM )................................................................................................................ 50 4.1 Abstract.................................................................................................................. 50 4.2 Introduction ............................................................................................................ 51 4.3 Materials and methods.............................................................................................. 53 4.4 Results ................................................................................................................... 55 4.5 Discussion............................................................................................................... 60 4.6 Acknowledgements................................................................................................... 63 1 ______________________________________________________________TABLE OF CONTENTS CHAPTER 5 - GENERAL DISCUSSION .............................................................................. 64 5.1 Symbiosis between beewolves and Streptomyces bacteria............................................. 64 5.2 The male pheromone and the impact of rearing temperature......................................... 69 5.3 Final conclusions and future prospects ........................................................................ 71 CHAPTER 6 - SUMMARY .................................................................................................. 72 6.1 Symbiosis between beewolves and Streptomyces bacteria............................................. 72 6.2 Male pheromone and temperature effects during larval development.............................. 73 CHAPTER 7 - ZUSAMMENFASSUNG.................................................................................. 74 7.1 Symbiose zwischen Bienenwölfen und Streptomyceten ................................................. 74 7.2 Männchenpheromon und Temperatureinfluss während der Larvalentwicklung .................. 75 REFERENCES................................................................................................................... 76 DANKSAGUNG………………………………………………………………………………………………………97 ERKLÄRUNG…………………………………………………………………………………………………………99 2 ____________________________________________________________LIST OF PUBLICATIONS LIST OF PUBLICATIONS This thesis is based on the following manuscripts: Kaltenpoth M, Goettler W, Dale C, Stubblefield JW, Herzner G., Roeser-Mueller K, Strohm, E (2006) 'Candidatus Streptomyces philanthi', an endosymbiotic streptomycete in the antennae of Philanthus digger wasps. International Journal of Systematic and Evolutionary Microbiology 56 (6): 1403-1411 (chapter 2). Roeser-Mueller K, Datzmann T, Seger J, Stubblefield JW, Herzner G, Strohm E, Kaltenpoth M (in preparation) 65 million years of defensive alliance: Molecular phylogeny of beewolves reveals the age of a protective symbiosis with Streptomyces bacteria (chapter 3). Roeser-Mueller K, Strohm E, Kaltenpoth M (2010) Larval rearing temperature influences amount and composition of the marking pheromone of the male beewolf, Philanthus triangulum . Journal of Insect Science 10:74 available online: insectscience.org/10.74 (chapter 4). 3 ______________________________________________________________________CHAPTER 1 CHAPTER 1 GENERAL INTRODUCTION The fitness of an individual and its maximisation are of central importance for evolutionary processes. There are different ways how organisms can increase their fitness and pass on their genes to the next generation. This thesis deals with two such aspects that might contribute to the fitness of an organism: an association with symbionts and sexual selection in courtship pheromones. First, the defensive symbiosis between beewolf digger wasps (Hymenoptera, Crabronidae) and Streptomyces bacteria is addressed. Another part of this thesis deals with the sex pheromone of male beewolves and how it is influenced by the temperature an individual is exposed to during development. In addition to these two fitness relevant traits, a reconstruction of the phylogenetic relationships among beewolves and closely related species is presented, that provides a basis for understanding the evolution of both the symbiosis and the composition of the sex pheromones in beewolves. The present chapter summarizes the current knowledge on symbioses and pheromones, in particular their relevance for mate choice, and it ends with an overview of the biology of beewolves. 1.1 Symbioses Symbioses have played an important role in the evolution of life on earth, and examples can be found in all kingdoms of microorganisms, plants and animals (Margulis 1999). According to the original definition of de Bary (1879), the term “symbiosis” encompasses the living together of unlike organisms over significant parts of their live span. This definition includes the whole range of biological interactions from mutualistic relationships via commensalism to parasitism, without specifying costs and benefits for the partners involved in the association (Douglas 2010). However, according to the most common definition of “symbiosis” in the current literature (Douglas 2010), in the present thesis the term “symbiosis” is used in its stricter sense, including only mutualistic associations. The variety of symbiotic interactions is characterized by different traits and definitions. Regarding the degree of interdependence between the symbiotic partners symbioses are either classified as facultative or obligate: Associations with a low dependency on the partner are defined as facultative, symbioses with a strong interdependence, meaning that a symbiont cannot survive without its partner, are defined as obligate (Douglas 2010; Ishikawa 2003). Concerning the location of
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