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The Pennsylvania State University The Graduate School Intercollege Graduate Degree Program in Ecology SOCIAL INSECTS AS SOLITARY VEHICLES A Dissertation in Ecology by Emilia Solá Gracia 2017 Emilia Solá Gracia Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy August 2017 ii The dissertation of Emilia Solá Gracia was reviewed and approved* by the following: David P. Hughes Assistant Professor of Entomology and Biology Dissertation Advisor Chair of Committee Nina Jenkins Senior Research Associate Victoria Braithwaite Professor of Fisheries and Biology Ephraim Hanks Assistant Professor Jason Kaye Professor of Soil Biogeochemistry Professor in Charge *Signatures are on file in the Graduate School iii ABSTRACT Parasite-host interactions affect more than the two main characters. Organisms living in tight-knit communities depend on each other in order to survive. Individuals harboring a parasite can affect the delicate balance within these communities. While on the other hand the community could also have a strong effect on a parasite’s life cycle. I explore the effects infected individuals have on the within-nest dynamics of ant colonies. Additionally, I investigate the effects of infectious and non-infectious cadaver exposure have on worker behavior, as well as determine the effects workers have on the fungus protruding from the infectious cadavers. Using both a coevolved fungus, Ophiocordyceps unilateralis sensus lato, and a generalist fungus, Beauveria bassiana, I found infected workers do not strongly affect the within nest dynamics of Camponotus castaneus, the natural host of O. unilateralis s. l. Furthermore C. castaneus and C. pennsylvanicus manage infectious and non-infectious cadavers differently. Workers from C. castaneus colonies are adept in quickly removing cadavers with internal and external fungal development. While C. pennsylvanicus workers must come in contact with fungal tissue in order to recognize the disease threat. However, I found C. pennsylvanicus workers to be highly effective in reducing the infectivity of hazardous fungal conidia (i.e. asexual spores). Such findings could be caused by the wood nesting ecology this species has, as well as the higher aggression towards infectious cadavers. My research lays the foundation for future studies which focus on using semi-natural settings to study parasite- host and community dynamics. iv TABLE OF CONTENTS LIST OF FIGURES ................................................................................................................. vi LIST OF TABLES ................................................................................................................... ix ACKNOWLEDGEMENTS ..................................................................................................... x DEDICATION ......................................................................................................................... xi Chapter 1: Introduction ....................................................................................................... 1 Background ...................................................................................................................... 1 Behavioral mechanisms for parasite defense ................................................................... 2 Objectives......................................................................................................................... 6 Chapter 2: Within the fortress: the tolerance of a specialized parasite in a social insect society ......................................................................................................... 8 Introduction ...................................................................................................................... 9 Materials and Methods ..................................................................................................... 12 Results .............................................................................................................................. 19 Discussion ........................................................................................................................ 22 Conclusion ....................................................................................................................... 28 Chapter 3: Observations on a generalist entomopathogen within a social insect fortress............................................................................................................................. 36 Introduction ...................................................................................................................... 37 Materials and Methods ..................................................................................................... 40 Results .............................................................................................................................. 45 Discussion ........................................................................................................................ 47 Conclusion ....................................................................................................................... 51 Chapter 4: Dynamics surrounding the dead: Do ant colonies recognize disease threats? ................................................................................................................. 60 Introduction ...................................................................................................................... 61 Materials and Methods ..................................................................................................... 65 Results .............................................................................................................................. 76 Discussion ........................................................................................................................ 79 Conclusion ....................................................................................................................... 84 Chapter 5: The effectiveness of social behaviors in reducing fungal pathogenicity ........ 103 Introduction ...................................................................................................................... 104 v Materials and Methods ..................................................................................................... 107 Results .............................................................................................................................. 114 Discussion ........................................................................................................................ 116 Conclusion ....................................................................................................................... 118 Chapter 6 Conclusion anf future directions ....................................................................... 128 Appendix A: Supplementary material for Chapter 2 ......................................................... 132 Appendix B: Supplementary material for Chapter 3 ......................................................... 137 Appendix C: Supplementary material for Chapter 4 ......................................................... 140 Appendix D: Supplementary material for Chapter 5 ......................................................... 143 References ............................................................................................................................... 146 vi LIST OF FIGURES Figure 2.1- Proportion of time spent in trophallaxis 6 days post-injection. ............................ 29 Figure 2.2- Extended trophallaxis observations ..................................................................... 30 Figure 2.3- Mean distance between focal individuals and nest entrance ............................... 32 Figure A .1- Mortality cuves ................................................................................................... 132 Figure A.2- Time spent within the nest .................................................................................. 133 Figure 3.1- Proportion of time workers spent in trophallaxis while inside the nest ............... 53 Figure 3.2- Number of trophallaxis events occuring within the nest ..................................... 55 Figure 3.3- Percent of the time individuals spent within the nest during our observational bouts .......................................................................................................... 57 Figure 3.4- Proportion time individuals performed vigilance behaviors while inside the nest ................................................................................................................................... 58 Figure B.1- Nest vigilance categorization .............................................................................. 137 Figure B.2- Healthy Beauveria bassiana growth from cadaver ............................................. 138 Figure B.3- Survival rate per treatment.................................................................................. 139 Figure 4.1- Time spent within th nest, percent of contacts, mortality, and infection for Camponotus castaneus nests which are open for the first 12 hours of exposure ............. 86 vii Figure 4.2- Time spent within th nest, percent of contacts, mortality, and infection for Camponotus pennsylvanicus nests which are open for the first 12 hours of exposure .... 88 Figure 4.3- Time spent within