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Coversheet for Thesis in Sussex Research Online A University of Sussex PhD thesis Available online via Sussex Research Online: http://sro.sussex.ac.uk/ This thesis is protected by copyright which belongs to the author. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the Author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the Author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Please visit Sussex Research Online for more information and further details The Behaviour and Chemistry of Recruitment and Alarm in Social Insects Thomas Butterfield Submitted for the Degree of Doctor of Philosophy University of Sussex School of Life Sciences March 2017 Declaration I hereby declare that this thesis has not been, and will not be, submitted in whole or in part to another University for the award of any other degree. ii Summary This thesis initially focuses on the chemical ecology of two species of ant which are common in the United Kingdom; Lasius flavus, the yellow meadow ant and Lasius niger, the common black ant. The first data chapter explores the constituent chemicals present in 3 major exocrine glands located in the gaster of L. flavus and discusses their potential functions. The work presented here also highlights the need for the comparative study of the chemical composition of glands. The second data chapter investigates how chemicals present in hindgut extractions of Lasius niger vary with the temporal caste they belong to (nurse or forager), and subsequently looks at how those chemicals may be suited to the tasks performed by that caste. The third data chapter describes the development of a highly sensitive methodology to identify low-concentration pheromones that uses a combination of analytical chemistry and behavioural bioassays. This methodology was used to identify two attractive pheromones of L. flavus, one is a trail pheromone used during foraging while the other is an alarm pheromone used to warn nest-mates of danger. The trail pheromone is the lowest concentration pheromone to be successfully identified in ants to date. This chapter also highlights the need to perform comparative behavioural bioassays to demonstrate the true function of putative pheromones. The final data chapter then investigates the source of alarm signals in Nasutitermes corniger termites and assesses the differential responses of workers and soldiers. This chapter then goes on to elucidate the colony-level effects of alarm on the regularity of repairs made to experimentally manipulated foraging galleries. iii Acknowledgements First of all, I’d like to take the opportunity to thank my supervisors, Professor Jonathan Bacon and Professor Elizabeth Hill. I’m very grateful that I was given the opportunity to undertake this PhD project, and their constant support and considerable expertise have enabled me to make a great contribution to the field. I must also thank Professor Francis Ratnieks for agreeing to house me in LASI, and for his constant and unrelenting reminders of my Yorkshire heritage (and for his supply of instant noodles, which just about staved off hunger in the cases that I’d forgotten an edible lunch). I’d also like to thank Sam Jones, his help at the very beginning of my PhD allowed me to hit the ground running which helped me to achieve much more than would otherwise have been possible. I’d also like to thank the other members of the Bacon lab. Primarily Alan Gallagher, who has accompanied me on many ant gathering trips and provided me with countless stimulating conversations regarding what questions we needed to tackle next in the world of ant research. The help of Caitlyn and Sebastian with ant maintenance and collection must also be acknowledged. Thanks also go to members of the Hill lab: Julia Horwood for helping me with countless questions on the practicalities of analytical chemistry; Andrew Chetwynd for his consistently negative outlook on his own (actually fantastic) research that always made me feel that I was making good progress; and Arthur David for his patience in helping me learn, and his expertise in, multivariate statistics. Thank you to the coffee-drinkers for getting me going in the mornings; Maggie, Roger, Victoria, Julia & Kyle (although his was a hot chocolate), and RIP to the LASI coffee iv machine, I’m sure it expired doing what it loved. I would also like to thank Maggie for her fantastically positive outlook on life, and for always being available to help me with research queries and concerns. Thank you to Roger for helping me with statistics and R, I feel like I’ve gone from a complete R novice to someone who at least looks like he knows what he’s doing, it wouldn’t have been possible without his help. I send my gratitude to the other members of LASI: Nick, Luciano and Hasan, whose long, exhausting days building and carrying beehives made me grateful that I was studying ants. I’d like to thank my parents and my sister, it feels like they’ve supported me in every decision I’ve ever made. Without their patience and encouragement I wouldn’t be where I am today. Finally, and most of all, I’d like to thank my wife Sarah. Her constant, unrelenting support and sound advice has gotten me through the trials and tribulations that come along with completing a PhD project. I wouldn’t have been able to do it without her. v Table of Contents The Behaviour and Chemistry of Recruitment and Alarm in Social Insects..............................A Thomas Butterfield ..............................................................................................................A Declaration ........................................................................................................................ ii Summary .......................................................................................................................... iii Acknowledgements.......................................................................................................... iv Table of Contents ............................................................................................................. vi 1 Introduction.................................................................................................................... 1 1.1 Eusocial Insects .............................................................................................................. 1 1.1.1 Eusociality.................................................................................................................................................. 1 1.1.2 Division of labour ..................................................................................................................................... 1 1.2 Chemical Signalling in Social Insects ............................................................................... 4 1.2.1 Trail pheromones ..................................................................................................................................... 5 1.2.2 Alarm pheromones .................................................................................................................................. 9 1.3 Analytical Techniques in Chemical Ecology.................................................................... 11 1.4 Aims ............................................................................................................................ 17 2 Chemical characterization of 3 major exocrine glands in the gaster of Lasius flavus ......................................................................................................................................... 19 2.1 Abstract....................................................................................................................... 19 2.2 Introduction ................................................................................................................. 20 2.3 Materials and Methods ................................................................................................ 24 2.3.1 Chemicals................................................................................................................................................. 24 vi 2.3.2 Ants ........................................................................................................................................................... 24 2.3.3 Gland dissection & extraction.............................................................................................................. 25 2.3.4 Chemical analysis ................................................................................................................................... 26 2.4 Results......................................................................................................................... 28 2.4.1 Compounds identified in the Dufour gland extractions .................................................................. 30 2.4.2 Compounds identified in the poison gland extractions .................................................................. 35 2.5 Discussion .................................................................................................................... 39 3 The right tools for the job: Temporal Biochemical Polymorphism in Lasius niger .... 44 3.1 Abstract....................................................................................................................... 44
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