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Studies on the Acoustical Behavior and Taxonomy Op STUDIES ON THE ACOUSTICAL BEHAVIOR AND TAXONOMY OP THE THEE CRICKETS (ORTHOPTEEA: OECANTHMftE) OF THE EASTERN UNITED STATES DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State U n iv e rs ity By THCMAS JEFFERSON WALKER, J R ., B. A ., M. S. ****** The Ohio State University 1957 Approved hy Deparmnem; o r Ziooj-ogy ana Entomology ACKNOVnjaXSMENTS Many persons contributed to the research reported in this dissertation. I wish to acknowledge their share in any merit the work may hare. Special acknowledgment is due my adviser, Donald J. Borror, who has given support, advice, and constructive criticism in all stages of the work. The knowledge and enthusiasm of Richard D. Alexander started me on this work, and discussion and field work with him have been a continuing source of stimulation. George Potor, Division of Biophysics, gave generously of his time and skill in constructing the electronic apparatus which produced the imitation tree cricket calls used in some of the bdiavioral studies. Thanks are due Basil Pames, Department of Physics, for the design of the apparatus used in study­ ing the periodicity of singing. I am indebted to Henry Wave Shaffer, Department of Physics, for advice concerning the mechanism of sound production in tree crickets. Many taxonomists were obliging in allowing me to examine their collections and in giving me the benefit of their experience in namenclatorial problems. Particularly helpful were Ashley B. Gurney, U. S. National Miseum, who helped in locating and obtaining type specimens; J. N. Khull, Ohio State University Museum, who brought specimens from southern Texas; and Edward S. Thomas, Ohio State Miseum, who informed me of an unusual form of tree cricket he had found on willows in several areas in Ohio. B. B. Pulton, T. H. ii iii Hubbell, and J. A. G. Rehn gave patient advice concerning nomencla- torial questions. Alvah Peterson and F. W. Fisk were generous in the loan of equipment used in the study of the effect of physical factors upon singing behavior. Finally I wish to thank the Graduate School, who made the last year of this work free from alien worries by supplying an ample research grant. TABLE OF CONTENTS INTRODUCTION................................................................................................................ 1 SOUND PRODUCTION AMONG TREE CKCCKETS........................................................... 3 Occurrence of Sound Production ............................................................... 3 Methods of Recording and A n aly sis ............................« ...................... 6 Mechanics of Sound Production ............................................................... 8 Structures Associated with Sound Production ......................... 8 Nature of Tegminal Movement ............................................. 10 Determination of P itc h ................................. .................................16 Variations in the Calling Song ....................... ......... 26 Major C ategories ...................................................................................... 26 Physical C haracteristics ................................................. 28 Effects of Environmental Factors ............................................ 34 Temperature ................................... ............................................ 34 H u m id ity ............................................................................ 63 Air Currents ................................... .............................................68 L i g h t ....................................................................................................71 S o u n d .................................................. ... ........................................72 Effects of Intrinsic Factors ......................................... 72 Individual V ariation ............................ ..................................72 Population V ariation ...............................................................75 Relation of Sound Production to the Diurnal Cycle ..... 78 Field O bservations ..................................................................................79 Laboratory Observations .................................................................... 79 TAXONOMY OF THE OEGANTHINAE OF EASTERN UNITED ST A T E S .............................. 89 Introduction ................................................................................ 89 Plan of Presentation ...........................................................................................91 Key to the Oecanthinae of Eastern United S ta te s .............................. 94 iv V Genus N eoxabea .................................................................... , ...................... 99 Neoxabea bipunctata (De Gear) ......................................... 99 Genus Oecanth.ua ............................................ 107 The rileyl Group ......................................,................................ 108 Oecanth.ua exclamation!s D avis .........................................109 O ecanthus n iv eu s (De G e e r ) ...................................................115 Oecanthus rileyl Baker ....................................................... 12? The latipennls Group ......................................................................... 158 Oecanthus latipennls R ile y ............................... 139 O ecanthus v a r ic o r n is F. W a l k e r .................................... 145 The nigricornis Group ..... .............................................. 152 Geographical Distribution .... 154 Ecological Distribution .. 160 Seasonal Distribution ........................................................... 165 Calling Songs ............................................................................. 168 M o r p h o lo g y ........................................................................ 175 Hybridization and Crossing Experiments ..... 189 Summary of Distinguishing C haracters ............................191 Oecanthus nigrleornls F. W alker ..........................................193 Oecanthus celerinictus n. sp .................................... 195 Oecanthus argentinus Saussure .......................... 197 Oecanthus quadrlpunctatus Beutenmuller ..... 200 Oecanthus pini Beutenmuller ............................... 202 vi BEHAVIORAL EFFECTS OF THE CALLING SONG.................................................... , 204 Introduction * 204 Synchronous Singing in the Snowy Tree Cricket ...................................204 M e t h o d s ........................................................................................................ 206 R e s u l t s ................................... 208 Significance of Synchronous Singing ..................................... 213 Specificity in the Response of Females to Calling Songs of the M ales ................................................................................ 213 M e t h o d s ........................................................................................................ 214 Results . .................................................................... ..... 220 Basis of Specificity Among Species Producing Continuous T rills .................................................................... 223 Specificity in Species Producing Broken T rills . 228 Specificity in 0. rileyl ..... ................................. 228 Discussion ...........................................................................................230 Summary ....................................................................... 230 APPENDIX: DISTRIBUTION RECORDS OF THE OECANTHINAE OF THE EASTERN UNITED STATES AND CANADA ..................................................................................232 REFERENCES CITED ..... ................................................................................. 269 AUTOBIOGRAHIY................................................................................................................. 280 LIST OF TABLES T able I. Effect of mutilation of the tegraina upon the calling song of Oecanthus nigricom is II. Effect of mutilation of the tegmina upon the calling song of Oecanthus latipennis III. Laboratory recordings made in the study of effects of temperature upon calling songs IV. Effect of high and low humidities upon the calling song of an individual of £. rileyi V. Effect of air currents upon the calling songs of three tree crickets 71. Fulton’s (1926b) classification of the eastern Oecan- thinae and changes indicated by the present study VII. T rill characters of Neoxabea bipunetata; field re c o rd in g s VIII. T rill characters of Oecanthus exclamationls; field re c o rd in g s IX. T rill characters of Oecanthus niveus; field recordings X. Frequency of occurrence of chirps with various numbers of pulses in the song of 0, r i l e y i XI. Measurements of 0. varicom is from Hidalgo and Cameron Counties, Texas XII. Antennal markings of the nigricom is group, pini excluded XIII. Measurements of members of the nigricom is group XEV. File characters of the nigricom is group 37. Usefulness of various characters in identifying the members of the nigricom is group XVI. Response scores and chi square values relating to specificity of response in females of species found in weedy fields viii Table X 7II. Response scores and chi square values relating to specificity of response in females of tree- dwelling
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