INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type ofcomputer printer. The quality ofthis reproduction is dependent upon the quality ofthe copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back ofthe book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A BeU & Howell Information Company 300 North Zeeb Road, Ann Arbor MI 48106·1346 USA 3131761-4700 800/521-0600 HAWAIIAN ENDEMIC COPIPHORINI: SYSTEMATICS AND ACOUSTICS (ORTHOPTERA: TETTIGONIIDAE: CONOCEPHALINAE) A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI" IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENTOMOLOGY MAY 1996 By John Sanford Strazanac M. Lee Goff, Chairperson Scott E. Miller Whitlow W. L. Au Sheila Conant Bruce E. Tabashnik UMI Number: 9629857 UMI Microform 9629857 Copyright 1996, by UMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 ACKNOVVLEDGMENTS The history of this dissertation can be divided into the research done while in Hawaii and the written portion, which was done on the mainland. I would like to express my appreciation to the members of my committee for sticking with me through a dissertation that required more time than one should. I would especially like to acknowledge Dr. Scott Miller's support and input throughout the entire process; Dr. Whitlow Au's discussions and guidance in acoustics; Dr. Sheila Conant's guidance in Hawaiian Natural History; and Dr. Tabashnik's support. Dr. Lee Goff came onto my committee as chair when I was moving from one institution to a more supportive environment at West Virginia University with Dr. Linda Butler. This combination seemed to have a synergistic effect as the finishing up went very quickly and smoothly. It is questionable that I would have been able to finish if it were not for the following individuals: Dr. Jon Gelhaus, Dr. Ed Theriot, Keith Russell, and Margaret Fisher of the Academy of Natural Sciences; Dr. Sue McCombs of the University of Hawaii; and Dr. Neil Reimer of Hawaii State Department of Agriculture. Also I would like to thank for assistance and friendship, Hideyuki Chiba, Keith Arakaki, Steve Montgomery, and Jong-Yong Kim. Invaluable for keeping the process moving along Millie Uegawachi and Lynn Hata. I would like to thank my daughter for her patience as I worked on my ll IIputer and sharing her projects with me. I need to deeply thank my wife Cynthia Fritzler for her faith through some tough times that prolonged the completion of this degree. iii ABSTRACT The endemic Hawaiian Copiphorini fauna consists of two lineages: Conocephaloides and Banza. The genus Conocephaloides is resurrected, but the type species appears to be extinct. A species found on Nihoa Island described as a Banza is transfered to Conocephaloides. Banza is represented on all of the main islands at mid to upper elevations in wet forests. The first acoustic analysis of the extant Hawaiian Copiphorini provides a guide to compare morphological and coloration characters. With new characters for Banza the status of Kauai Island species is changed, a sibling species not previously recognized is described from Maui Island and the variation of the Hawaii Island Banza species now appears to form a cline. The song of the Nihoa Island Conocephaloides species is unusual for Copiphorini katydids. iv ------_.._---- - _.. TABLE OF CONTENTS Acknowlegements . iii Abstract . iv List of Tables ' . vi List of Figures . vii Introduction . 1 Materials and Methods . 11 Taxonomy . 19 Acoustics . 94 Phylogenetics ............................................. .. 125 Appendix: Leg Spine Counts. ................................ .. 131 References. .............................................. .. 147 v LIST OF TABLES Table Page 1. Characteristics of wing closure rates of Conocephaloides nihoa (Hebard) and Banza species .....•........................ 153 2. Tooth click number and tegmen closure duration of Banza kauaiensis, B. parvula, B. unica, B. molokaiensis, B. brunnea, B. mauiensis, B. pilimauiensis, and B. deplanata 154 3. Tooth click number and tegmen closure durationof Banza nitida populations 155 vi LIST OF FIGURES Figure Page 1. Subgenital plate of male Conocephaloides nihoa (Hebard) . .. 156 2. Stridulatory file of Conocephaloides nihoa (Hebard) . .. 157 3. Individual teeth of s!ridulatory file of Conocephaloides nihoa (Hebard) ........................................... .. 158 4. Terminalia of male Conocephaloides nihoa (Hebard) . .. 159 5. Sculpturing of Conocephaloides nihoa (Hebard) egg surface anterior region ...................................... .. 160 6. Sculpturing of Conocephaloides nihoa (Hebard) egg surface posterior region. .................................... •. 161 7. Sculpturing of Conocephaloides nihoa (Hebard) egg surface with an indistinct transition zone between anterior and mid regions . .. 162 8. Stridulatory file of Banza kauaiensis affinis (Perkins) .......... .. 163 9. Individual teeth of stridulatory file of Banza kauaiensis affinis (Perkins) ........................................... .. 164 10. Terminalia of male Banza parvula (Walker) ................. .. 165 11. Sculpturing of Banza parvula (Walker) egg surface anterior region. ............................................ .. 166 12. Sculpturing of Banza parvula (Walker) egg surface posterior region. ............................................ .. 167 vii 13. Sculpturing of Banza parvula (Walker) egg surface mid region .... 168 14. Sculpturing of Banza parvula (Walker) egg surface with an indistinct transition zone between anterior and mid regions ............. 169 15. Terminalia of male Banza kauaiensis (Perkins) .•......•...•... 170 16. Sculpturing of Banza kauaiensis affinis (Perkins) egg surface ant'enor region. ........................................ 171 17. Sculpturing of Banza kauaiensis affinis (Perkins) egg surface posterior region ...........................•........... 172 18. Sculpturing of Banza kauaiensis affinis (Perkins) egg surface mid region ............................................... 173 19. Terminalia of male Banza unica (Perkins) ..................... 174 20. Sculpturing of Banza unica (Perkins) egg surface anterior region .. 175 21. Sculpturing of Banza unica (Perkins) egg surface mid region ...... 176 22. Sculpturing of Banza unica (Perkins) egg surface posterior region.. 177 23. Terminalia of male Banza molokaiensis (Perkins) .............. 178 24. Sculpturing of Banza molokaiensis (Perkins) egg surface anterior region ............................................... 179 25. Sculpturing of Banza molokaiensis (Perkins) egg surface posterior region ............................................... 180 viii -------------------------_ ..__._----------- 26. Sculpturing of Banza molokaiensis (Perkins) egg surface mid region. ............................................ .. 181 27. Terminalia of male Banza brunnea (Perkins) •..... .. .. ... .. .. 182 28. Sculpturing of Banza brunnea (Perkins) egg surface anterior region. ................•........................... .. 183 29. Sculpturing of Banza brunnea (Perkins) egg surface posterior region ............................................... 184 30. Sculpturing of Banza brunnea (Perkins) egg surface mid region ... 185 31. Terminalia of male Banza mauiensis (Perkins) ................. 186 32. Sculpturing of Banza mauiensis (Perkins) egg surface anterior region ............................................... 187 33. Sculpturing of Banza mauiensis (Perkins) egg surface posterior region ............................................... 188 34. Sculpturing of Banza mauiensis (Perkins) egg surface mid region .. 189 35. Terminalia of male Banza pilimauiensis new species ............ 190 36. Sculpturing of Banza pilimauiensis new species egg surface anterior region ........................................ 191 37. Sculpturing of Banza pilimauien~is new species egg surface posterior region. .................................... .. 192 38. Sculpturing of Banza pilimauiensis new species egg surface mid region. ............................................. .. 193 ix ----------------------- - -- -------- 39. Terminalia of male Banza deplanata (Brunner v. W.) ............ 194 40. Sculpturing of Banza deplanata (Brunner v. W.) egg surface anterior region ........................................ 195 41. Sculpturing of Banza deplanata (Brunner