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Worker Policing in the European Hornet Vespa Crabro 6.1 ABSTRACT……

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Worker Policing in the European Hornet Vespa Crabro 6.1 ABSTRACT…… The Conflict over Male Production in the Vespinae Wasps A THESIS PRESENTED TO UNIVERSITY OF SHEFFIELD DEPARTMENT OF ANIMAL AND PLANT SCIENCES FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Kevin R. Foster November 2000 SUPERVISED BY FRANCIS L.W. RATNIEKS I dedicate my thesis to my wonderful parents, for this is clearly all their fault. iii Abstract Conflict is inherent in social systems. The eusocial hymenoptera (bees, ants and wasps) are no exception with several reproductive conflicts affecting their societies. This thesis investigates the resolution one of these conflicts, that over male production, in a group of known phylogeny, the Vespinae wasps (hornets and yellowjackets). Extensive conflict is predicted as both the queen and workers can produce males. However, worker behaviour is expected to depend on queen mating frequency (formally effective paternity). With single paternity, workers are more related to each other’s sons than the queen’s sons and worker male production is expected. With paternity above two, workers are more related to the queen’s sons and mutual policing (worker policing) is predicted. To test these predictions, paternity was analysed and compared to worker reproductive behaviour in seven vespine species from the genera Vespa, Dolichovespula (five species) and Vespula. In line with prediction, Dolichovespula had mostly single paternity and worker male production in colonies with a queen. Furthermore, in D. saxonica, worker male production was correlated with paternity among colonies: evidence of facultative worker policing. Vespula vulgaris had double paternity, no worker male production and worker policing. Vespa crabro did not fit relatedness predictions with single paternity but worker policing. Finally, queen loss in reproductive colonies negatively correlated with paternity across species suggesting that matricide in response to reproductive conflict may occur. Five general conclusions on the resolution of reproductive conflicts can be made. (1) Relatedness is important. (2) Other factors, such as the cost of conflict, are also important as worker policing was found at paternity below two in V. crabro and some D. saxonica and V. vulgaris colonies. (3) Worker policing is important. (4) Conflict resolution is evolutionarily labile, with otherwise similar species having very different resolutions to conflict. (5) Conflict is associated with low colony size, with most conflict in Dolichovespula which has the smallest colonies. iv Acknowledgments So many people to thank. First of all my supervisor Francis, who was always there to help and turned my confusion into science. John Gulliver and Julie McGill for making it all possible. My labmates Adam, Steve and Thibaud and soul mates Matt, Fi, Kate and Jon who make me smile. Perttü, Cia, Niclas, Anja, Matts, Palle, Simon and Per provided light in the darkness of Uppsala and Minttu, Heikki, Katja, Kriko and Lotta helped me ‘find wasps’ in Finland. Thanks to Matt Sledge, for sharing Italy and stimulating discussions, Tom Wenseleers for numerous debates over beer and Steve Martin for teaching me the ways of the tropics. In Dorset, Alan Raybould helped me overcome oppressive administration and Pete, Zoe, Andy, Deanne and Becka made the most of two long hot summers. Emily, Dan, Miki, Nick, Tim, Ross, Rob and my sister Gillian have been there throughout. Finally, there is Madeline, who is my love. This thesis was funded by a BBSRC studentship and a CASE award from the Institute of Terrestrial Ecology, Furzebrook, Dorset. v Table of Contents CHAPTER 1. Introduction 1.1 THEORETICAL BACKGROUND ..................................................................................................1 1.11 Kin structure and reproductive conflict.....................................................................................1 1.12 Conflict over male production...................................................................................................1 Empirical data ...................................................................................................................................2 1.2 WHY STUDY THE VESPINAE WASPS?......................................................................................3 1.21 A well-supported phylogeny .....................................................................................................4 Similar basic biology.........................................................................................................................4 1.23 A few important variables: kin structure, conflict resolution and colony size ..........................6 CHAPTER 2. How the thesis evolved 2.1 THE EUROPEAN HORNET, VESPA CRABRO..............................................................................7 2.11 Paternity and worker relatedness...............................................................................................7 2.12 Zombie hornets..........................................................................................................................7 2.13 The effect of genetics on cuticular chemistry............................................................................8 2.2 SPITE…………................................................................................................................................8 2.3 DOLICHOVESPULA WASPS ........................................................................................................9 2.31 New Forest collection and observation .....................................................................................9 2.32 Microsatellite analysis.............................................................................................................10 2.4 THE COMMON WASP VESPULA VULGARIS ........................................................................10 2.41 Microsatellite analysis.............................................................................................................10 2.42 Worker policing.......................................................................................................................10 2.5 THE HUNT FOR PROVESPA........................................................................................................11 2.6 THE HUNT FOR VESPULA AUSTRIACA .................................................................................12 2.7 THE EVOLUTION OF WORKER POLICING AT LOW PATERNITY......................................12 2.8 WORKER POLICING IN THE HORNET VESPA CRABRO........................................................12 CHAPTER 3. Methodology 3.1 DNA MICROSATELLITE ANALYSIS ........................................................................................13 3.11 DNA extraction .......................................................................................................................13 3.12 PCR reaction ...........................................................................................................................13 3.13 Separation and visualisation....................................................................................................13 3.2 STATISTICAL ANALYSIS OF GENETIC DATA.......................................................................14 3.21 Worker-worker relatedness and effective paternity.................................................................14 3.22 Errors in paternity estimation ..................................................................................................16 3.23 Errors in worker male production estimation ..........................................................................16 3.3 COLLECTION AND STUDY OF LIVE WASPS .........................................................................17 3.31 Finding nests ...........................................................................................................................17 3.32 Nest collection.........................................................................................................................17 3.33 Nest relocation.........................................................................................................................17 3.34 Nest observation......................................................................................................................19 3.35 Worker policing assay.............................................................................................................19 vi CHAPTER 4. Low paternity in the hornet Vespa crabro indicates that multiple mating by queens is derived in vespine wasps 4.1 ABSTRACT……............................................................................................................................21 4.2 INTRODUCTION ..........................................................................................................................21 4.3 METHODS…….. ...........................................................................................................................22 4.31 Study organism and sample collection ....................................................................................22 4.32 Molecular methods..................................................................................................................22 4.33 Statistical methods...................................................................................................................23 4.34 Pedigree estimates ...................................................................................................................23 4.35 Non-sampling error
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