Do Butterflies Use “Hearing Aids”? Investigating the Structure and Function of Inflated Wing Veins in Nymphalidae

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Do Butterflies Use “Hearing Aids”? Investigating the Structure and Function of Inflated Wing Veins in Nymphalidae Do butterflies use “hearing aids”? Investigating the structure and function of inflated wing veins in Nymphalidae by Penghui (Carrie) Sun A thesis submitted to the Faculty of Graduate and Postdoctoral Affairs in partial fulfillment of the requirements for the degree of Master of Biology in Biology Carleton University Ottawa, Ontario © 2018 Penghui (Carrie) Sun Abstract Many butterfly species within the subfamily Satyrinae (Nymphalidae) have been informally reported to possess a conspicuous “inflated” or “swollen” subcostal vein on each forewing. However, the function and taxonomic diversity of these structures is unknown. This thesis comprises both experimental and comparative approaches to test hypotheses on the function and evolution of these inflated veins. A laser vibrometry study showed that ears in the common wood nymph, Cercyonis pegala, are tuned to sounds between 1-5 kHz and the inflated subcostal vein enhances sensitivity to these sounds. A comparative study showed that all species with inflated veins possess ears, but not all species with ears possess inflated veins. Further, inflated veins were better developed in smaller butterflies. This thesis provides the first evidence for the function of inflated wing veins in butterflies and supports the hypothesis that they function as aids to low frequency hearing. ii Acknowledgements I thank my supervisor Dr. Jayne Yack for the continued guidance and support, throughout my academic program and in beginning my career, as well as an inspired and newfound appreciation I never knew I could have for insects. I thank my committee members Dr. Jeff Dawson and Dr. Charles-Antoine Darveau for their guidance, advice, and support. I thank Dr. Natasha Mhatre for being an amazing researcher and teacher; I am grateful for her patience and skills when answering all my questions and helping me understand. I thank Dr. Andrew Mason for collaboration and mentoring the brilliant Natasha that I got to work with, and Dr. Akito Kawahara for collaboration and sending many cool exotic specimens I got to sample. I thank Rick Cavasin for his beautiful photos that are used with permission in my thesis and publications. I thank Jeff Skevington, Michael Olsen, Peter Hall, Chris Schmidt, Ross Layberry, and members of the Ottawa annual butterfly count for their help and advice in specimen collection. I thank Laura Hall, Jessica King, Chelsea Stunden, and Caitlin Preston for their specimens and unpublished data. Lastly, I thank lab members Andras Dobai, Melanie Scallion, Conrado Denadai, Chanchal Yadav, and Mairelys Naranjo for their support and encouragement. I thank Alexander Pochopsky for his continuous support and encouragement throughout this higher education journey, and for his enthusiasm and encouragement to take the journey in the first place. I am forever thankful for, and could not have succeeded, without his one-liners about butterflies, his home-cooked meals, his beefy computer, and his role as a practice audience member, life partner, and best friend. Lastly, I thank anyone who reads this in the future, for any reason. iii Table of Contents Do butterflies use “hearing aids”? Investigating the structure and function of inflated wing veins in Nymphalidae ................................................................................................................................. i Abstract ............................................................................................................................................ ii Acknowledgements ......................................................................................................................... iii List of Tables ................................................................................................................................. vii List of Figures ............................................................................................................................... viii List of Appendices ........................................................................................................................... x Chapter 1. General Introduction .................................................................................................... 10 1.1. Acoustic communication in insects ................................................................................ 10 1.2. Butterfly ears .................................................................................................................. 11 1.2.1. Acoustic communication in butterflies .................................................................. 11 1.2.2. Hearing in butterflies ............................................................................................. 12 1.3. “Inflated” wing veins ..................................................................................................... 13 1.3.1. What are they? ............................................................................................................. 13 1.3.2. Who has inflated wing veins? ...................................................................................... 14 1.3.3. What is/are the function(s) of inflated veins? .............................................................. 14 Chapter 2. Butterflies use Inflated Wing Veins as “Hearing Aids” ............................................... 19 2.1. Abstract ............................................................................................................................... 20 2.2. Introduction ......................................................................................................................... 20 2.2. Methods .............................................................................................................................. 21 2.2.1. Animals ........................................................................................................................ 21 2.2.2. Tympanal and wing vein morphology ......................................................................... 22 2.2.3. Laser vibrometry and acoustic stimuli ......................................................................... 22 2.2.4. Data analysis ................................................................................................................ 23 2.3. Results ................................................................................................................................. 24 2.3.1. Morphology.................................................................................................................. 24 2.3.2. Mechanical response of tympanum .............................................................................. 26 iv 2.4. Discussion ........................................................................................................................... 29 Chapter 3. Wing Veins as “Hearing Aids” in Butterflies: A Comparative Study .......................... 31 3.1. Abstract ............................................................................................................................... 32 3.2. Introduction ......................................................................................................................... 33 3.2.1. Hearing in butterflies ................................................................................................... 34 3.2.2. Inflated wing veins ....................................................................................................... 37 3.2.3. Inflated wing veins relative to ears, taxa, sex, and body size ...................................... 39 3.2.4. Do small butterflies tend to have larger inflations? ..................................................... 40 3.3. Methods .............................................................................................................................. 42 3.3.1. Specimens .................................................................................................................... 42 3.3.2. Morphology of body size, wing vein inflations, and ears ............................................ 43 3.3.3. Taxonomic distribution of subcostal vein and ears ...................................................... 50 3.3.4. Analyses of vein and body size .................................................................................... 50 3.4. Results ................................................................................................................................. 52 3.4.1. Vein and Ear Morphology............................................................................................ 52 3.4.2. Ears .............................................................................................................................. 61 3.4.3. Relationship between veins and ears ........................................................................... 62 3.4.4. Sexual Dimorphism ..................................................................................................... 65 3.4.5. Taxonomic distribution ................................................................................................ 67 3.4.6. Relationship of subcostal vein size to body size .......................................................... 69 3.5. Discussion ........................................................................................................................... 79 3.5.1. What is a Wing Vein Inflation? ................................................................................... 79 3.5.2. Are Inflated
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