Repair of the Proboscis of Brush-Footed Butterflies (Lepidoptera: Nymphalidae)" (2014)
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Clemson University TigerPrints All Theses Theses 8-2014 REPAIR OF THE PROBOSCIS OF BRUSH- FOOTED BUTTERFLIES (LEPIDOPTERA: NYMPHALIDAE) Suellen Pometto Clemson University, [email protected] Follow this and additional works at: https://tigerprints.clemson.edu/all_theses Part of the Entomology Commons Recommended Citation Pometto, Suellen, "REPAIR OF THE PROBOSCIS OF BRUSH-FOOTED BUTTERFLIES (LEPIDOPTERA: NYMPHALIDAE)" (2014). All Theses. 1881. https://tigerprints.clemson.edu/all_theses/1881 This Thesis is brought to you for free and open access by the Theses at TigerPrints. It has been accepted for inclusion in All Theses by an authorized administrator of TigerPrints. For more information, please contact [email protected]. REPAIR OF THE PROBOSCIS OF BRUSH-FOOTED BUTTERFLIES (LEPIDOPTERA: NYMPHALIDAE) ______________________________________ A Thesis Presented to the Graduate School of Clemson University ______________________________________ In Partial Fulfillment of the Requirements for the Degree Master of Science Entomology ______________________________________ by Suellen Floyd Pometto August 2014 ______________________________________ Accepted by: Dr. John C. Morse, Committee Co-Chair Dr. Peter H. Adler, Committee Co-Chair Dr. John Hains ABSTRACT A key feature of the order Lepidoptera is the coilable proboscis, present in over 99% of lepidopteran species. The proboscis is used to obtain liquid nutrition, usually floral nectar. The proboscis is assembled from two elongate galeae immediately after emergence of the adult from the pupa. What happens if the galeae become separated? I studied the process of repair of the proboscis, behaviorally and functionally, at the organismal level. My research questions were as follows: 1) is the proboscis capable of repair, 2) is saliva necessary to proboscis repair, and 3) is the repaired proboscis able to acquire fluids? Test organisms were Danaus plexippus (Linnaeus) (Lepidoptera: Nymphalidae: Danainae) and Vanessa cardui (Linnaeus) (Lepidoptera: Nymphalidae: Nymphalinae). To assess repair capability, at 24 hours after eclosion the galeae were separated totally and compared to control specimens without galeal separation. Both species were able to accomplish repair of the proboscis, with 10–100% achieving reunion of the galeae and recovery of the coiled resting state. The role of saliva in repair of the proboscis was tested for D. plexippus, with the hypothesis that removal of saliva would impede repair. Three groups were compared: 1) the proboscis was manipulated with a capillary tube to remove saliva for five minutes, 2) the proboscis was similarly manipulated for five minutes but no saliva was removed, and 3) the proboscis was not manipulated and no saliva was removed. Both manipulated groups were similarly impeded, indicating that manipulation impeded repair, rather than removal of saliva. To test functionality of the repaired proboscises, at 24–48 hours after separation of the galeae all butterflies were offered dyed sucrose-water, with the hypothesis that the repaired ii proboscis can acquire fluids. After feeding, butterflies were placed in separate cages lined with filter paper. Successful feeding was confirmed by the presence of dye in the gut exudate on the filter paper or in the gut by dissection for 64–100% of all groups. Physical and functional self-repair of the butterfly proboscis could provide a model for artificially engineered microfluidic devices. iii DEDICATION This work is dedicated to my father, Robert Forsythe Floyd†, who nurtured my love of Nature from the time I was a small child, and to my husband, Anthony Louis Pometto, III, who made many sacrifices to allow me to pursue graduate studies. I thank my children for their encouragement and support. Three of my five children have attended college concurrently with me. I give thanks to God for allowing me to use His gifts to study His creatures and for granting me healing from ovarian cancer and restoration to good health to continue this work. iv ACKNOWLEDGMENTS Funding for this project was from National Science Foundation grants EFRI 0937985 and PoLS 1305338 for the project “Biomechanics of Self-Assembly of the Lepidopteran Feeding Devices” with co-principal investigators Dr. Peter H. Adler and Dr. Konstantin G. Kornev, and NSF grant IOS-1354956, “Mechanisms of Fluid Feeding in Insects, from Nanoscale to Organism” to Dr. Peter H. Adler and Dr. Konstantin G. Kornev. I thank the Greater Charleston Pest Control Association for the James D. “Denny” Ford award which provided support for my graduate research. I thank my committee members, Dr. John C. Morse, Dr. Peter H. Adler, and Dr. John Hains, for their mentoring during my research and study. Dr. Morse accepted me as his student and assisted me with project ideas and funding. Dr. Morse inspired me with his enthusiasm and helped me with his patient personal assistance. Dr. Adler provided my final project and funding through a graduate research assistantship. Dr. Adler has given me ceaseless encouragement from the time I took my first entomology course, Taxonomy of Insects, in 2009. I thank Dr. Charles “Eddie” Beard for his calm and wise presence and for his practical help and guidance with this project. I thank Dr. James Rieck for assistance with statistics. I thank Dr. Alfred P. “Hap” Wheeler for giving me a graduate teaching assistantship in Biological Sciences, without which I could not have pursued this course of study. I thank my daughter, Mary Ann, for formatting assistance and for using her graphics arts skills on Figure 1.1. v TABLE OF CONTENTS Page ABSTRACT ..................................................................................................................................... ii DEDICATION ................................................................................................................................ iv ACKNOWLEDGMENTS ................................................................................................................ v LIST OF TABLES .......................................................................................................................... ix LIST OF FIGURES .......................................................................................................................... x CHAPTER 1: INTRODUCTION .................................................................................................... 1 CHAPTER 2: LITERATURE REVIEW ......................................................................................... 5 Phylogenic Context for Study of the Lepidopteran Proboscis ................................................ 5 Evolutionary History of Lepidoptera .............................................................................. 5 Order Level Phylogenetic Considerations....................................................................... 7 Synapomorphies of Amphiesmenoptera ......................................................................... 8 Synapomorphies of the Order Lepidoptera ..................................................................... 9 Phylogeny within Lepidoptera ........................................................................................ 9 Basal Suborders of Lepidoptera .................................................................................... 11 The Suborder Glossata .................................................................................................. 11 Molecular Phylogenetic Evidence ................................................................................. 15 Feeding Habits of Adult Lepidopterans ................................................................................ 16 Non-Feeding Adults ...................................................................................................... 18 Water Drinking ............................................................................................................. 18 Nectar Feeding .............................................................................................................. 19 Feeding on Sap, Honeydew, and Honey ....................................................................... 20 Feeding on Over-Ripe or Rotting Fruit ......................................................................... 21 Fruit Piercing ................................................................................................................. 22 Feeding on Animal Fluids ............................................................................................. 22 Pollen feeding ............................................................................................................... 24 Model Species for Study of the Lepidopteran Proboscis ...................................................... 25 Pieris brassicae (Linnaeus) ........................................................................................... 25 Manduca sexta (Linnaeus) ............................................................................................ 32 vi (Table of Contents Continued) Structure of Adult Mouthparts of the Lepidoptera: ............................................................... 34 The Larval Mouthparts .................................................................................................. 34 Adults of Primitive Suborders ....................................................................................... 34 Adults of the Suborder Glossata: General Structure of the Proboscis .......................... 35 Proboscis Specializations