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Coversheet for Thesis in Sussex Research Online A University of Sussex DPhil 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 UNIVERSITY OF SUSSEX Submitted for the Degree of Doctor of Philosophy CHRISTOPHER TRANTER Defending the Fortress comparative studies of disease resistance in ant societies SUMMARY Parasites represent a considerable and ubiquitous threat to organisms, and studies of host-parasite interactions can demonstrate important insights into key biological processes. Identification and quantification of host defences and their role in parasite resistance is an important part of understanding these effects. Additionally, life-history traits can have significant effects on host-parasite interactions. For example, living in groups has many benefits, but also may have associated costs in terms of increased parasite transmission. Thus group-living animals may be predicted to invest heavily in disease resistance strategies, though which may depend on each species’ parasite pressure. Social insects, and ants in particular, are an ideal model with which to test these evolutionary and ecological hypotheses, as they possess an array of mechanisms to defend themselves against disease and have highly diverse life-histories. However, previous studies into disease resistance tend to have been performed on single species, often looking at just single measures of investment of defence. In this thesis I explore the comparative importance of disease resistance in different ant species. I show that ants possess a variety of defence mechanisms to protect themselves against the threat of parasites and demonstrate how investment into these important defences can vary between individuals and species, and may depend on context, type of parasite, and life- history of the host. Work such as this, demonstrating the costs of individual components of disease resistance in multiple species, is important in developing our understanding of how changes in parasite pressures can influence host biology and how organisms can survive in a world abundant with parasites. Defending the Fortress comparative studies of disease resistance in ant societies Christopher Tranter Submitted for the Degree of Doctor of Philosophy University of Sussex Faculty of Life Science September 2015 i Declaration I confirm that the work submitted is my own, except where work which has formed part of jointly-authored publications is included. The contributions of other authors are indicated for published chapters below. This thesis has not been and will not be submitted to another institution for the award of any other degree. List of published chapters and contributions: Chapter 2. Tranter, C., LeFevre, L., Evison, S.E.F. & Hughes, W.O.H. 2015. Threat detection: contextual recognition and response to parasites by ants. Behavioural Ecology. 26: 396–405. CT designed the experiments carried out the experimental work, analysed the data, and wrote the manuscript. LF conducted preliminary experimental trials. SEFE supervised the work of LF. WOHH supervised the work, assisted with the design of the experiments, the data analysis, and proofing the manuscript. Chapter 3. Tranter, C., Graystock, P., Shaw, C., Lopes, J.F.S. & Hughes, W.O.H. 2014. Sanitizing the fortress: protection of ant brood and nest material by worker antibiotics. Behavioural Ecology and Sociobiology 68: 499–507. CT designed the experiments carried out the experimental work, analysed the data, and wrote the manuscript. PG aided with the experimental work in blocking the glands of ants. CS conducted preliminary experimental trials. JFSL supplied the ant colonies. WOHH supervised the work, assisted with the design of the experiments, the data analysis, and proofing the manuscript. Chapter 4. Tranter, C. & Hughes, W.O.H. 2015. Acid, silk and grooming: alternative strategies in social immunity in ants? Behavioural Ecology and Sociobiology. 69: 1687-1699 CT designed the experiments carried out the experimental work, analysed the data, and wrote the manuscript. WOHH aided in the manuscript proofing. ii Chapter 5. Tranter, C., Fernandez-Marin, H., & Hughes, W.O.H. 2015. Quality and quantity: transitions in antimicrobial gland use for parasite defence. Ecology and Evolution (in press) CT and WOHH designed the experiments, CT carried out the fieldwork and experimental work, analysed the data, and wrote the manuscript. WOHH aided in the manuscript writing and HFM aided with fieldwork. Chapter 6. Tranter, C. & Hughes, W.O.H. 2015. Simple societies and sanitary syndromes: species and individual differences in ant social immunity. Biology Letters (in review) CT and WOHH designed the experiments, CT and TM carried out the experimental work, CT analysed the data, and wrote the manuscript. WOHH aided in the manuscript writing. Chapter 7. Tranter, C. & Hughes, W.O.H. 2015. Sanitary slaves and slovenly slavemakers: disease resistance of social parasites and their hosts. Proceedings of the Royal Society of London: B (in review) CT and WOHH designed the experiments, CT carried out the experimental work, analysed the data, and wrote the manuscript. WOHH aided in the manuscript writing. Appendix Tranter, C. & Hughes, W.O.H. 2015. A preliminary study of nest structure and composition of the weaver ant Polyrhachis (Cyrtomyrma) delecta (Hymenoptera: Formicidae) Journal of Natural History. In press CT designed the experiments carried out the experimental work, analysed the data, and wrote the manuscript. WOHH aided in the manuscript proofing. Signed ___________________________________ Date ______________________ iii Acknowledgments Thanks to Bill Hughes for the opportunity to undertake this PhD, for being ever supportive, no matter where in the world either of us might have been, and for giving me the chance to prove myself as a scientist through work that has been both exciting and rewarding. To the members of the evolving Hughes Lab: thank you to those who welcomed me to Leeds and to PhD life, when I was first starting out, and provided me with the technical knowledge and friendship that has carried through my project and to the current members of the Hughes Lab who have helped feed ants, or with feedback on manuscripts, and have enabled me to produce the best work I can. I always manage to find someone to organise me (a skill in itself), and Jasmine Parkinson, this was you. We’ve shared 4 years together and I am immeasurably grateful for your help, patience and compassion. Thank you to Victoria Norman for being a tolerant fieldworker, an enthusiastic biker, and a great friend. I’d also like to mention all the other students, scientists and organisations, I’ve met and collaborated with, both at home and abroad, who helped with ant collection and identification, or lent equipment, or a hand, and who without which this PhD would have been considerably more challenging. Thanks to my family who have always been supportive of whatever I’ve done and have helped get me where I am today. To my mother and father, thank you for the years of selfless commitment, the hours of your evenings spent driving me to and from activities, the places you took me, and chances you gave. I have certainly not said it enough, but that these things seemed routine, that you made these opportunities normal, demonstrates how truly lucky I have been. Finally thanks to Jen for unending support; I’ve taught you about ants, and you’ve taught me about myself. Thank you for your patience at the countless times we’ve been out and I’ve chased around after an ant, for the 5am starts in the dark for the perfect photo, for the mountains we’ve climbed, for the memories, for our future. iv Abstract Parasites represent a considerable and ubiquitous threat to organisms, and studies of host-parasite interactions can demonstrate important insights into key biological processes. Identification and quantification of host defences and their role in parasite resistance is an important part of understanding these effects. Additionally, life-history traits can have significant effects on host-parasite interactions. For example, living in groups has many benefits, but also may have associated costs in terms of increased parasite transmission. Thus group-living animals may be predicted to invest heavily in disease resistance strategies, though which may depend on each species’ parasite pressure. Social insects, and ants in particular, are an ideal model with which to test these evolutionary and ecological hypotheses, as they possess an array of mechanisms to defend themselves against disease and have highly diverse life-histories. However, previous studies into disease resistance tend to have been performed on single species, often looking at just single measures of investment of defence. In this thesis I explore the comparative importance of disease resistance in different ant species. I show that ants possess a variety of defence mechanisms to protect themselves against the threat of parasites and demonstrate how investment into these important defences can vary between
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