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The Role of Cuticular Hydrocarbons in Determining Male Reproductive Success

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The Role of Cuticular Hydrocarbons in Determining Male Reproductive Success View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Open Research Exeter 1 THE ROLE OF CUTICULAR HYDROCARBONS IN DETERMINING MALE REPRODUCTIVE SUCCESS Submitted by Sarah Marie Lane to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences November 2015 This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. (Signature).................................................................................................................................. 2 ABSTRACT Cuticular hydrocarbons (CHCs) are found on the outer cuticle of all terrestrial arthropods. Although their primary function is in desiccation prevention, these compounds have also been shown to play a variety of roles in insect chemical communication, from species and sex recognition to providing cues of dominance and attractiveness. However, despite growing evidence of their versatility as cues, our knowledge of how CHCs are used in mating interactions is limited to Drosophila and field crickets. In this thesis I investigate the roles CHCs play in interactions at each stage of the mating process in the broad- horned flour beetle Gnatocerus cornutus. I assess the relative importance of CHCs in influencing male reproductive success and examine the complex interplay between different episodes of selection and the mechanisms of sexual selection acting on males. I use a combination of behavioural assays, experimental manipulations and gas chromatography. First, I identify the role of CHCs as cues of sperm competition risk and intensity, demonstrating how the presence of male-derived CHCs on the cuticles of virgin females elicits males to adjust their pre- and post-copulatory investment (chapter 2), by providing information on the state of their competitive environment. I then go on to look at the stability of CHCs as cues of sperm competition over time, finding that they are highly sensitive to environmental degradation (chapter 3) and do not persist in the habitat substrate of this species. Next, I investigate how male CHCs determine fighting and mating success. By estimating and comparing the strength and form of sexual selection imposed by male-male competition and female mate choice, I show that male CHCs are subject to strong antagonistic sexual selection (chapter 4). By experimentally manipulating male CHC profile, I then attempt to verify the selection gradients estimated for female choice 3 (chapter 5). However, my experimental manipulation fails to verify the importance of male CHCs for female mate choice. Finally, I explore the role of same-sex sexual behaviour (SSB) in determining male reproductive success (chapter 6). I find evidence to suggest that SSB may in fact be a form of aggression in its own right, and demonstrate that SSB and fighting may provide equivalent means for males to overcome female choice and secure a mating advantage. My results indicate that CHCs play key roles as chemical cues throughout the mating process and significantly impact male reproductive success. My thesis reveals the intricate nature of the relationships between mechanisms of sexual selection, alongside highlighting the need to consider both the social and physical environment when investigating the importance of chemical cues. I discuss the implications of these results for the evolution of male CHCs and how my findings can be used to further our knowledge of this field. 4 ACKNOWLEDGEMENTS First and foremost I would like to thank my supervisors Clarissa and Tom for steering me through the past three years. Clarissa, I know that this thesis is a million miles away from the question we started out with and I want to thank you for giving me the freedom to shape my thesis around my interests. Thank you for helping me to build resilience to the downsides of science, the exciting ideas that don’t quite work in reality and the reviewers who call your work “simply bad science”, but also to celebrate the successes. Tom, I want to thank you for taking on the arduous role of being my second supervisor. I think the three of us have made an excellent team throughout and I couldn’t have done it without your combined support, encouragement and knowledge. I hope I’ve done you proud. I would like to thank my mum and dad for never questioning my choice of career path. For their unyielding love and support, for listening to me go on and on about how great or terrible things are depending on the day, and for gracefully accepting their inevitable education on the subject of beetles (soon to extend to the exciting world of sea anemones!). Thank you for driving for seven hours just to see me for the weekend, it means so much to me to make you both proud. Thank you to the wonderful people who together make the CEC such a vibrant place to work, I can only hope that the folks at Plymouth will be half as friendly and fiercely intelligent as you all. To all the people who made the lab a fun place to be, Corrina for party Fridays (“Corrina, is that Linkin Park?”), Alex, Chris Mitchell and Melia. To my brilliant undergrads, Jo, Andy, Alice and Dan, for not only being fabulous students but for being great company too. To my cheerful and bonkers officemates past (Mitch, Dom, Harry, Faye, Kim, Feargus) and present (Sarah P, Emily, Lucy, Rebecca, Sean, Billy, Phil, Lili), thank you all for 5 making me want to come to work every day. To PhD students past who inspired me to walk the treacherous path of academia, Fiona, Ruth and the ever lovely Jenni McDonald. To all my wonderful friends. To Nicole for always being there for me, I honestly don’t know if I would have made it through these past three years without you. In fact, I do know, I wouldn’t have. I’d have gone stark raving mad. To my glorious cohort, to Sarah P for our ridiculous giggling and inspiring chats about science, and to Dave and Sean for being our big brothers. To all those who are too numerous to mention but have made my life in Cornwall so fantastic. Last but definitely not least, to Grim. Thank you for turning my life into something wonderful and filling every day with laughter. No matter how gloomy my outlook, you always always manage to make me smile. Thank you for giving me the energy to push on when I thought I couldn’t give anymore. I love you. 6 TABLE OF CONTENTS Abstract ...................................................................................................................................... 2 Acknowledgements ................................................................................................................... 4 List of tables and figures ............................................................................................................ 8 Author’s declaration ................................................................................................................ 10 CHAPTER 1: General introduction ................................................................................... 11 1.1 Sexual selection and male reproductive success ............................................................. 11 1.2 Cuticular hydrocarbons (CHCs) ....................................................................................... 14 1.3 CHCs and male-female interactions ................................................................................ 16 1.3.1 Mate choice (female assessment) ............................................................................... 16 1.3.2 Assessing sperm competition risk and female mating status (male assessment) ....... 17 1.4 CHCs and male-male interactions .................................................................................... 20 1.4.1 Dominance and fighting success ................................................................................. 20 1.4.2 Same-sex sexual behaviour (male-male) .................................................................... 22 1.5 Study system .................................................................................................................... 23 1.6 Thesis aims and outline .................................................................................................... 25 CHAPTER 2: Rival male chemical cues evoke changes in pre- and post-copulatory investment in a flour beetle ............................................................................................ 29 2.1 Abstract ............................................................................................................................ 29 2.2 Introduction ..................................................................................................................... 30 2.3 Materials and Methods .................................................................................................... 33 2.4 Results .............................................................................................................................. 41 2.5 Discussion ........................................................................................................................ 44 CHAPTER 3: Rapid degradation of chemical cues of sperm competition ........................... 56 3.1 Abstract ...........................................................................................................................
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