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Eco-Physiological Causes and Consequences of Sexually Selected Color Variation in Dragonflies

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Eco-Physiological Causes and Consequences of Sexually Selected Color Variation in Dragonflies ECO-PHYSIOLOGICAL CAUSES AND CONSEQUENCES OF SEXUALLY SELECTED COLOR VARIATION IN DRAGONFLIES by MICHAEL P. MOORE Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Advisor: Ryan A. Martin Department of Biology CASE WESTERN RESERVE UNIVERSITY August, 2019 CASE WESTERN RESERVE UNIVERSITY SCHOOL OF GRADUATE STUDIES We hereby approve the dissertation of Michael P. Moore Candidate for the degree of Doctor of Philosophy* Committee Chair Dr. Ryan A. Martin Committee Member Dr. Sarah E. Diamond Committee Member Dr. Michael F. Benard Committee Member Dr. Patrick D. Lorch Date of Defense May 13, 2019 * We also certify that written approval has been obtained for any proprietary material contained therein 1 TABLE OF CONTENTS LIST OF TABLES .......................................................................................................8 LIST OF FIGURES ...................................................................................................10 ACKNOWLEDGEMENTS ......................................................................................12 ABSTRACT ................................................................................................................14 CHAPTER 1: AN INTRODUCTION TO THE ECO-PHYSIOLOGICAL CAUSES AND CONSEQUENCES OF SEXUALLY SELECTED COLOR VARIATION IN DRAGONFLIES ..........................................................................16 BACKGROUND ........................................................................................................16 Ecological causes of variation in sexually selected characters ....................................18 Consequences of sexual selection for variation in ecological adaptation ....................21 ECO-PHYSIOLOGICAL CAUSES AND CONSEQUENCES OF DRAGONFLY WING COLOR VARIATION .......................................................23 A primer on wing coloration in dragonflies and damselflies .......................................24 Pachydiplax longipennis: a future model system? .......................................................25 Research Overview ......................................................................................................27 Sexual selection on wing coloration ............................................................................27 Potential eco-physiological causes of sexually selected color variation .....................28 Potential eco-physiological consequences of sexual selection on wing coloration .....30 2 CHAPTER 2: INTRASEXUAL SELECTION FAVORS AN IMMUNE-CORRELATED ORNAMENT IN A DRAGONFLY..........................33 ABSTRACT ................................................................................................................33 INTRODUCTION .....................................................................................................34 METHODS .................................................................................................................37 Study System ...............................................................................................................37 Study Site, Population Monitoring, and Phenotypic Measurements ...........................38 Do territory holding success and longevity vary with wing melanization? .................39 Do males with greater wing melanization receive less aggression from territorial rivals? ...........................................................................................................41 Is wing melanization associated with fighting ability? ................................................43 Does wing melanization correlate with melanin-based immune responses? ...............44 Statistical Analyses ......................................................................................................45 RESULTS ...................................................................................................................46 Phenotypic variation ....................................................................................................46 Do territory holding success and longevity vary with wing melanization? .................46 Do males with greater wing melanization receive less aggression from territorial rivals? ...........................................................................................................47 Is wing melanization associated with fighting ability? ................................................47 3 Does wing melanization correlate with melanin-based immune responses? ...............48 DISCUSSION .............................................................................................................48 CHAPTER 3: TEMPERATURE SHAPES THE COSTS, BENEFITS, AND GEOGRAPHIC DIVERSIFICATION OF SEXUAL COLORATION IN A DRAGONFLY ............................................................................................................59 ABSTRACT ................................................................................................................59 INTRODUCTION .....................................................................................................60 METHODS .................................................................................................................63 Wing coloration in Pachydiplax longipennis ...............................................................63 Prediction 1: Wing coloration increases male body temperatures ...............................63 Prediction 2: Thermal consequences of wing coloration translate into performance variation ..................................................................................................66 Prediction 3: Wing coloration provides its greatest territorial advantages on the coldest and/or most thermally variable days ..........................................................67 Prediction 4: Reduced wing coloration in hottest parts of range .................................69 RESULTS ...................................................................................................................72 Prediction 1: Wing coloration increases male body temperatures ...............................72 Prediction 2: Thermal consequences of wing coloration translate into performance variation ..................................................................................................72 Prediction 3: Wing coloration provides its greatest territorial advantages 4 on the coldest and/or most thermally variable days ....................................................73 Prediction 4: Reduced wing coloration in hottest parts of range .................................73 DISCUSSION .............................................................................................................74 CHAPTER 4: TRADE-OFFS BETWEEN LARVAL SURVIVAL AND ADULT ORNAMENT DEVELOPMENT DEPEND ON PREDATOR REGIME IN A TERRITORIAL DRAGONFLY .....................................................................84 ABSTRACT ................................................................................................................84 INTRODUCTION .....................................................................................................85 METHODS .................................................................................................................88 Focal species and study overview ................................................................................88 Aim 1: ornament development and viability selection ................................................89 Ornament development ................................................................................................91 Larval Survival .............................................................................................................92 Aim 2: performance, morphological, and physiological correlates of body condition .............................................................................................................93 RESULTS ...................................................................................................................95 Aim 1: ornament development and viability selection ................................................95 Ornament development ................................................................................................95 Aim 2: Performance and morphological correlates of body condition ........................96 5 DISCUSSION .............................................................................................................97 CHAPTER 5: IMMUNE DEPLOYMENT INCREASES LARVAL VULNERABILITY TO PREDATORS AND INHIBITS ADULT LIFE-HISTORY TRAITS IN A DRAGONFLY ...................................106 ABSTRACT ..............................................................................................................106 INTRODUCTION ...................................................................................................107 METHODS ...............................................................................................................110 Study Species and Study Overview ...........................................................................110 Collection and Manipulating Immune Deployment .............................................112 Melanin deposition and immune challenge duration .................................................113 Laboratory Experiment: Does larval immune deployment affect predator avoidance traits? ..........................................................................................114
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