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Patterns and Mechanisms of the Exploitation of Mutualisms

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Patterns and Mechanisms of the Exploitation of Mutualisms PATTERNS AND MECHANISMS OF THE EXPLOITATION OF MUTUALISMS A Thesis Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Master of Science in The Department of Biological Sciences by Ellen McGrail Reid B.A. Boston University, 2005 August 2010 ACKNOWLEDGEMENTS I would like to thank the members of my advisory committee: Kyle Harms, Jim Cronin, and Richard Stevens for their guidance and advice. I would especially like to thank Kyle Harms, my advisor, for his encouragement, patience, and assistance over the somewhat difficult course of my studies. I would like to thank my lab mates: Adriana Bravo and Jonathan Myers, two wonderful scientists and great people who were always ready to listen and help. I am indebted to the Lowy Foundation at Louisiana State University for funding my participation in a course through the Organization for Tropical Studies, Tropical Biology: An Ecological Approach in 2008. My work during that course provided inspiration for this thesis. I would also like to thank the Organization for Tropical Studies for funding my proposal for the grant that allowed me to carry out much of this work in Palo Verde National Park in Costa Rica. I would not have been able to complete my thesis without the help and support of several friends and colleagues: Joe Sapp, Susan Whitehead, Andre Kessler, Katja Poveda, Amanda Accamando, Natalia Aristizabal, Jessie Deichmann, Meche Gavilanez, Noah Reid, Maria Sagot, and many others. I owe many thanks to Prissy Milligan for helping me navigate the logistics of finishing my thesis. I thank Dr. C. Kelly Swing, the inspirational teacher, biologist, and conservationist who first introduced me to fieldwork in the tropics. Thanks to my parents, Robert Reid and Kathleen McGrail, who had the confidence that I could chart my path and supported me along my way. Finally, I would like to thank David Johnson for his encouragement and support. ii TABLE OF CONTENTS ACKNOWLEDGEMENTS .......................................................................................................................... ii ABSTRACT................................................................................................................................................. iv CHAPTER 1: INTRODUCTION ................................................................................................................. 1 CHAPTER 2: EXPLOITATION OF AN ANT-ACACIA MUTUALISM BY MOZENA SP. (HEMIPTERA: COREIDAE)....................................................................................................................... 3 INTRODUCTION .................................................................................................................................... 3 METHODS............................................................................................................................................... 5 Study site and collection of study organisms ....................................................................................... 5 Specificity to ant colony of origin........................................................................................................ 5 Species-specificity to Vachellia host.................................................................................................... 5 Species-specificity to ant host species.................................................................................................. 6 Ant response to glandular compounds ................................................................................................. 6 Ant response to lanolin-dissolved cuticular and glandular compounds............................................... 6 Removal of coreid cuticular hydrocarbons .......................................................................................... 7 Ant response to methyl salicylate......................................................................................................... 7 Chemical analyses ................................................................................................................................ 7 RESULTS................................................................................................................................................. 8 Specificity to ant colony of origin........................................................................................................ 8 Species-specificity to Vachellia host.................................................................................................... 8 Species-specificity to ant host species.................................................................................................. 8 Ant response to glandular compounds ................................................................................................. 8 Ant response to lanolin-dissolved cuticular and glandular compounds............................................. 11 Removal of coreid cuticular hydrocarbons ........................................................................................ 12 Ant response to methyl salicylate....................................................................................................... 12 Chemical analyses .............................................................................................................................. 12 DISCUSSION......................................................................................................................................... 13 Species-specificity to ant- and plant-hosts ......................................................................................... 13 The role of coreid glandular and cuticular compounds...................................................................... 14 The role of methyl salicylate in ant-coreid interactions..................................................................... 15 Chemical analyses .............................................................................................................................. 15 Conclusions ........................................................................................................................................ 15 CHAPTER 3: THE COSTS OF EXPLOITATION OF MUTUALISMS .................................................. 17 INTRODUCTION .................................................................................................................................. 17 METHODS............................................................................................................................................. 18 RESULTS............................................................................................................................................... 19 DISCUSSION......................................................................................................................................... 22 CHAPTER 4: CONCLUSIONS ................................................................................................................. 24 LITERATURE CITED ............................................................................................................................... 25 VITA ........................................................................................................................................................... 31 iii ABSTRACT Mutualisms are reciprocally exploitative interactions providing net benefits to both partners. These interactions can be exploited, in turn, by individuals that take advantage of benefits offered by one or both partners in a mutualism, while offering no benefits in return. For many mutualist-exploiter interactions the mechanisms allowing exploitation, and the maintenance of mutualisms in the face of exploitation, are still poorly understood. Here I describe manipulative field and laboratory experiments to investigate the mechanisms used by an exploiter to invade an ant-plant mutualism. I tested two non-mutually exclusive hypotheses for how a coreid (Mozena sp., Hemiptera: Coreidae) feeds on mymecophytic acacia trees (Vachellia spp.) while avoiding attack by resident ants: chemical defense and chemical mimicry. I found that chemical compounds produced by Mozena sp. in both the metathoracic gland and the cuticle reduced the number of ant attacks and cuticular compounds appeared to be essential in escaping recognition on ant-occupied Vachellia spp. trees. The compounds were effective on multiple colonies and for multiple ant species, thus they are not strictly host- or species-specific. In addition, gas chromatography and mass spectrometry analyses of cuticular compounds revealed a close match between chemical profiles of Mozena sp. and Pseudomyrmex spinicola ants, suggesting chemical mimicry is the primary mechanism by which Mozena sp. exploits the ant- acacia mutualism. To examine the prevalence of a cost of exploitation for plant partners in exploited mutualisms, I conducted a meta-analysis of studies from the published literature. I found that exploitation has a weak, negative, but insignificant impact on the reproductive success of mutualistic plants. Collectively, these analyses illuminated methods by which exploiters may succeed in infiltrating mutualisms and suggested that the relatively low costs of exploitation may account for the lack of destabilization and degradation by exploiters of some mutualistic interactions. iv CHAPTER 1 INTRODUCTION Mutualisms are reciprocally
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