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Xerox University Microfilms INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is custGmary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 74-10,990 KOLSON, Richard Arthur, 1927- BEHAVIOR OF THE BULLFROG (RANA CATESBEIANA) WITH SPECIAL EMPHASIS ON VOCALIZATION. The Ohio State University, Ph.D., 1973 Zoology University Microfilms, A XEROX Company, Ann Arbor, Michigan 0 1974 RICHARD AUTHUR KOLSON ALL RIGHTS RESERVED THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. BEHAVIOR OF THE BULLFROG (RANA CATESBEIANA) WITH SPECIAL EMPHASIS ON VOCALIZATION DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Richard Arthur Kolson, B.Sc., M.Sc. * * * * * The Ohio State University 1973 Dr. Donald J. Borror Dr. Theodore A. Bookhout Approved by Dr. Loren S. Putnam Dr. Walter C. Rothenbuhler Adviser // Department of Zoology TABLE OF CONTENTS Page ACKNOWLEDGEMENTS . .............................. ii VITA ................................................. iii LIST OF T A B L E S ' . iv LIST OF ILLUSTRATIONS............................ V INTRODUCTION ....................................... 1 MATERIALS AND METHODS ..... ................ 7 Construction of the Experimental Pond . 7 The Pond The Observation Building Fencing the Area Maintenance of the Area Bullfrogs Used During the S t u d y ........... 18 Capture of Specimens Marking of Specimens Maintenance of Specimens Instrumentation of Experimental Area .... 31 Barometric Pressure Rainfall Wind Maximum-Minimum Temperature Weather Description Temperature Humidity Index Sound R e c o r d i n g ......................... 40 Recordings from Natural Ponds Recordings from the Experimental Pond Sonagram Terminology Use of a. Ceramic Model for Behavior Studies 51 RESULTS, ANALYSES, AND DISCUSSION .............. 60 Self Maintaining Behavior .................. 60 Hibernation and Emergence Daily Activity Hunting and Food Seeking Behavior Influence of Sex and Age RESULTS, ANALYSES, AMD DISCUSSION (Continued) Vocal Behavior................................ 100 Types of Vocalizations Produced The Effects of Temperature on Vocalization The Call Energy Index (CEI) Territorial Behavior .................... 145 Maintenance of Territory Playback and Model Presentation Mating B e h a v i o r ......................... 154 CONCLUSIONS ....................................... 157 SUMMARY ............................................ 169 LITERATURE CITED .................................. 176 ACKNOWLEDGEMENTS Although there are many people who have helped me, I am especially indebted to Dr. Donald Borror, whose knowledge and advice were indispensable. Even though he is not here to read this I wish to express my deep gratitude to the late Arthur M. Vorys. He was more a friend than a landlord and never hesitated to allow me to construct my experimental area on his property. I also wish to extend special thanks to Dr. Walter Rothenbuhler and Dr. Kraig Adler for their encouragement and contagious enthusiasm. My sincere thanks to Dr. C. Benjamin Meleca for insuring that I had the time to complete the final writing. And finally, my wife Nancy, who was and is my severest and most trusted critic. VITA December 5, 1927 . Born— Cleveland, Ohio 1946 ................ Willoughby Union High School, Willoughby, Ohio 1947-1950 ...... Employed as chemist at Barium and Chemicals, Inc., of Willoughby, Ohio 1951-1953.............. Army service. Medical Basic Training; transferred to Army •Chemical Corps, Edgewood Arsenal, Maryland 1957 ......... B.Sc., The Ohio State University, Columbus, Ohio. Major: Pathogenic Bacteriology 1961-1962 Research Fellow, Wildlife Research Unit, The Ohio State University, Columbus, Ohio 1962 ......... M.Sc., The Onio State University, C o l u m b u s O h i o . Thesis title: Pathogenesis of Leptospirosis in some Small Mammals Following Experimental Infection 1962-1970 Graduate Teaching Assistant and Associate, Department of Zoology, The Ohio State University, Columbus, Ohio 1970-present Research Specialist, Introductory Biology Program, The Ohio State University, Columbus, Ohio LIST OF TABLES Table Page 1. Vocalizations produced by the Bullfrog .............................. 101-102 2. Parameter Estimates for the "Mating C a l l " ................................ 124 3. Parameter Estimates for the "Territorial C a l l " ................................ 125 LIST OF ILLUSTRATIONS Figure Page 1. Completed pond excavation ................ 10 2 . Elevation of the observation building and enclosure ......................... 17 3. Top view of the experimental area .... 17 4. The experimental pond in 1969 . 20 5. Collection data and identification form . 24 6. Young male bullfrog with bead marker . 27 7. Thermocouple and sunshade . 35 8 . Positions of thermocouple on pole and f l o a t ................................... 39 9. Float in w a t e r ............................ 39 10. Experimenral area with parabolae reflectors set-up for sound recording ............ 43 11. Vocalisation terminology ......... 49 12. Ceramic model of bullfrog used during playback and model presentation exoeriments ............. 53 13. Wooden slide block ....................... 55 14. Arrangement of suspension wires, control strings and p u l l e y s ........... 55 15. The ceramic model floating in the "low posture"................................ 59 16. The ceramic model floating in the "high posture" and being approached by a female bullfrog ....................... 59 v 17. Relationship of air and mud temperature of experimental pond, August 1958 to August 1970 ......................... 63 18. Various temperatures of the experimental area for a typical late spring day (May 21, 1 9 5 9 ) ......................... 72 19.. Comparison of the relative humidity of the experimental area with that of the adjacent f i e l d ..................... 80 20. Various temperatures of the experimental area for a typical fall day (October 8, 1 9 5 8 ) ..................... 95 21. Juvenile cluck followed by adult pop . 105 22. Adult pop followed by two non-pulsatile territorial call notes ........... 105 23. Pulsatile territorial call note followed by adult p o p ............................ 105 24. Adult male growl followed by male pop . 109 25. Adult male g r o w l .......................... 109 25. Adult male g r o w l .......................... 109 27. Adult alarm call or s q u a w k .............. Ill 28. Two non-pulsatile call notes followed by four burps ending in a p o p ............ Ill 29. Two juvenile c l u c k s ....................... Ill 30. Female release calls ..................... 115 31. Female release calls ..................... 115 32. Male clasping calls ............ 115 33. Male orowl followed by 3 female release c a l l s .................. 117 34. Four female release calls followed by male growl ....................... 117 vi 35. Male growl followed by 2 female release calls ......................... 117 35. Mating call, June 10, 1959 . 120 37. Mating call, June 18, 1953 . 120 33. Mating call, June 21, 195 9 . 120 39. Territorial call, August 11, 1969 120 40. Territorial call, August 14, 1959 120 41. Territorial call, August 19, 1969 120 42 . Mating call, July 5 , 1970 . 122 43. Mating call, July 21, 196 7 .. 122 44. Mating „all., July 28, 195 5 . 122 45. Territorial call, August 26, 1969 122 AS Territorial call, August 28, 1959 122 47. Territorial call, September 1, 19? 122 48. Non-pulsatile terri tor.i al call 128 49. Mon-nulsatile territorial call .128 50. Mon—uulsati1e territorial call • 128 si; Female territorial call ......... 131 52. Female territorial call ......... 131 53. Female territorial
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