THE BIOACOUSTICS OF TABANIDAE (DIPTERA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Bastiaan Meijer Drees. B.A., M.S. * * * * * The Ohio State University 1980 Reading Committee: Approved By Dr. Donald E. Johnston Dr. Glen R. Needham Dr. Donald J. Borror ACKNOWLEDGEMENTS This investigation could not have been undertaken without the help and support of many individuals. I am deeply grateful to my adviser, Professor Donald E. Johnston, for encouragement and guidance throughout my studies, and to my father, Mr. Jan Meijer Drees, for providing technical assistance for the aerodynamic aspects of insect flight. I am also indebted to Professor Donald J. Borror for the time and effort he gave in teaching me the fundamentals and techniques of bioacoustics, and to Dr. l. L. Pechuman of Cornell University for his continued assistance with the difficult dipterous family, Tabanidae. Special thanks must also be given to Professor linda Butler of West Virginia University for giYing me a broad base of entomological knowledge and the enthusiasm to pursue my interests. I have enjoyed working with my examination committee members, Dr. Glen R. Needham and Professor Willard C. Myser and I thank them for undertaking a project which was slightly out of the ordinary. Other members of The Ohio State University faculty who have had a profound influence on my years as a doctorate candidate include, Or. D. J. Horn, Dr . G. Ekis, Dr. D. L. Denlinger, Dr. C. A. Triplehorn, Dr. G. W. Wharton, Dr. W. A. Foster and Dr. W. C. Rothenbuhler. Day to day ii existence in Columbus, Ohio was possible and even enjoyable only because of the close friendships and support of my fellow graduate students including Dr. Dan Potter, Dr . Jay Bradfield, Beth Lenoble, Maury Walsh, Greg Walker, Lucille Antony, Connie Rogers, Larry Ross, Ann Gnagey, and Rick Helmich. Finally, I wish to thank the two women in my life: my mother, Jacoba Meijer Drees, and my wife-to-be, Carol Frost, to whom this work is dedicated . iii VITA June 28, 1952 Born, Amsterdam, Netherlands 1974 B.A., Biology with minors in Chemistry and Art, West Virginia University, Morgantown, wv. 1974-1976 • • . • • • • • • laboratory Technician, curator of the Rhopalocera and Tabanidae sections of the Insect Museum, and application of green house pest control in the Horticulture Greenhouse for the Department of Entomology, West Virginia University, Morgantown, WV. 1975-1976 . • • • • • • . • Insect Specialist, identifications and control recommendations, assisting Or. J. F. 8aniecld, West Virginia Cooperative Extension Service, West Virginia University, Morgantown, WV. 1976 Andrew Delmar Hopkins Scholarship in Entomology, West Virginia University, Morgantown, WV. 1976 M.Sc., Entomology, West Virginia University, Morgantown, WV. 1977 Teaching Associate, Economic Entomology, The Ohio State University, Columbus, OH. 1977-1979 Ohio's Survey Entomologist, in cooperation with the United States Department of Agriculture, the Ohio Department of Agriculture, the Ohio Agricultural Research and Development Center, and the Ohio Cooperative Extension Service, The Ohio State University, Columbus, OH. 1979 • . • • . • . • • • Teaching Associate, General Biology, Department of Zoology, The Ohio State University, Columbus, OH. iv 1979 Administrative Assistant and insect specialist for Dr. R. l. Miller, Ohio Cooperative Extension Service, The Ohio State University, Cc~~bus, OH. 1979-1980 ••.•.••.• Teaching Associate, Insect Morphology, Economic Entomology, and General Entomology, Department of Entomology, The Ohio State University, Columbus, OH. 1980 • • • • • . • • • • Instructor, Biology- The World of Insects, Urban Extension and Community Programs, The Ohio State University, Columbus, OH. 1980 ••.•...•••• Ph.D., Entomology, The Ohio State University, Columbus, OH. PUBLICATIONS Baniecki, J. F., and B. M. Drees. 1979. General leaf Feeding Caterpillars, Pest Information Series 87. Cooperative Extension Service, West V1rginia University, Morgantwon, WV. 1976. Ants and their control. Pest Information Series 95. Cooperative Extension Service, West V1rginia Unlversi~-­ Morgantown, WV. 1976. Fleas and their control. Pest Information Series 96. Cooperative Extension Service, West VTrgln1a University--.----­ Morgantown, WV. Drees, B. M. 1977 . in The Cooperative Plant Pest Report 2(15-36). APHIS-PPQ, Unitea-states Department-or-Agriculture, Hyattsville, MD. Citations from the 1977 Ohio Cooperative Economic Insect Report (1-26) . -- --- 1978. in The Cooperative Plant Pest Report 3(16-36). APHIS­ PPQ, United~tates Department Of'~riculture, Hyattsville, MD. Citations from the 1978 Ohio Cooperative Economic Insect Report {1-26). - -- Drees, B. M. and l. Butler. 1978. Rhopalocera of West Virginia. J. lepidopterist's Society 32(3):1g2-206. Clement, S. L., G. Szatmari-Goodman, and B. M. Drees. 1979. The status and control of the Ohio corn rootworms. Ohio Report 64(1): 8-10. v Drees, B. H.• L. Butler, and L. L. Pechuman. In lress. The horse flies and deer flies of West Virginia; An-rl ustrated key (Diptera, Tabanidae). Biological Bulletin, West Virginia Experiment Station, West Virginia University, Morgantown. WV. FIELDS OF STUDY Major field: Entomology Areas of Specialization: Biology of Rhopalocera and Tabanidae, insects of economic importance. and insect behavior. vi TABLE OF CONTENTS Page ACKNOWLEDGEMENTS ii VITA ••••• iv LIST OF TABLES ix LIST OF FIGURES X INTRODUCTION • • literature Review of Tabanid Sexual Behavior and Rationale for Investigating the Sounds of Tabanid Flight 2 METHODS AND MATERIALS • • 11 Collection Techniques 11 Recording Techniques. • • • • • • 12 Wing Modifications or Mutilations 15 iemperature Modifications 15 Sonagrams • • . • . • . 16 loudness. • . • 17 Anatomical Measurements 17 Wet Weight • 17 Wing Slides. 18 Wing Length. 18 Chord. • • 18 Wing Area. • 19 Wing Loading . • • . • . • 19 Effective Aerodynamic Center 19 RESULTS AND DISCUSSION . • . • • 23 Elements of Tabanid Flight Sound. 23 Temperature and Sound • . • . • • . • . • • . • 25 The Effect of Wing Dimensions, Wet Weight and Wing Loading on Flight Sound . • . • 28 Wing length. • . 31 Chord. • . • 32 Wing Area . • . 32 Effective Aerodynamic Center . 33 vii Page Wet Weight •••••••••• 35 Wing Loading ••••••••• 37 Loudness or Sound Intensity ..•. 41 Sexual Differences of Tabanid Sound • 42 Relation of Wing Beat Rate to Habitat 43 Concluding Remarks. 45 SUI-t!ARY. 84 APPENDIX A. Collection Data for Specmens Recorded 86 B. Flight Mode of Diptera • • • • • • • 92 C. Sample Calculation of the Effective Aerodynamic Center. 93 BIBLIOGRAPHY . • . • • • • • • • • • • • • • • • • • • . • • 94 viii LIST OF TABLES Table Page 1 Observations of Tabanid "Courtship" Flights ... 6 2 Means, Standard Deviations, and Sample Sizes for the Wing Lengths, Wing Areas, Wet Weights, Wing Loads and Wing Beat Frequencies Between 26 and 2B°C for Thirty-nine Species of Tabanidae .•. 29 3 Factors Influencing the Sounds of Tabanidae (Di ptera) . • . • • . • . • • . • • . • . • • 47 ix LIST OF FIGURES Figure Page Container used for rearing field collected late instartabanid larvae • . • . • • • . • • • 48 2 Two dimensional modified Malaise trap baited with dry ice and a shiny black sphere for field collecting adult Tabanidae •.•.••••••.•.•.•.• 50 3 Recording chamber - top view with lid removed and leaning against one side .•.••••••••.• 52 4 Specimen holder, constructed of 1/4 in. plexiglass and 1/8 in. diam. plexiglass rods. 54 5 The outline of a tabanid wing •.•••••.••. 56 6 Chrysops cincticornis attached to the tethering rod with rubber cement, resting on the trap door •••. 58 7 Chrysoys cincticornis in typical attitude of flight after osing ground contact with the trap door ••• 58 8 Sound production • • . • • 60 9 Flight sounds of Tabanidae (Diptera) 62 10 The relationship of the wing beat frequency to hannoni cs. • • • • • • • • • • • . • • • • • • 64 11 Average wing beat rates (cps) and standard deviations calculated for 17 species of Pangoniinae • • . • • • 66 12 Average wing beat rates (cps) and standard deviations calculated for 22 species of Tabaninae • • • . • . 68 13 Sonogram of Chrysops vittatus (female) showing the fundamental tone or wing beat frequency (Hz or cps) and 5 harmonic overtones • . • • . • • . • . • • . 70 X Figure Page 14 The effect of temperature changes on the wing beat rates of individual tabanid specimens. • . • 72 15 The effect of temperature range and sample size on the presentation of results. • • . • . • • • • 74 16 The relationship between mean wing beat rates (Y axis) and mean wing lengths (X axis) of 12 species of Pangoniinae and 20 species of Tabaninae. • • . 76 17 Wing beat rate and wing length relationships of Tabanidae and Culicidae, and Greenewalt's (1962) boundary line. • . • . • . • • • • . 78 18 The effect of increased wing length (X axis) on wing shape (Y axis) . • • • • • • . • • • 80 19 The relationship of wing length (X axis) to body weight (Y axis). • . • . • • . • . • 82 xi INTRODUCTION Since Roth (1948) demonstrated conclusively that male Aedes aegypti were attracted to the sound produced by the flight of the female, bioacoustically and ethologically oriented entomologists have been searching for biological significance of the noises produced by the wing movements of other Diptera. Much work has been done with Culicidae (Costello, 1974); Wishart and Riordan, 1959), Drosophila (Williams and Gambos, 1950; Sharp et al., 1975; Bennet-Clark