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University Microfilms International A Bell & Howell Information Company 300 North Zeeb Road, Ann Arbor, Ml 48106-1346 USA 313/761-4700 800/521-0600 Order Number 0001002 A systematic study of the leafhopper tribe Hecalini (Homoptera: Cicadellidae: Deltocephalinae) of North America Ma, Ning, Ph.D. The Ohio State University, 1989 UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 A SYSTEMATIC STUDY OF THE LEAFHOPPER TRIBE HECALINI (HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE) OF NORTH AMERICA DISSERTATION Presented in the Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University by Ning Ma, M. S. * * * * * •* The Ohio State University 1989 Reading Committee: Approved by: Charles A. Triplehorn Norman F. Johnson David J. Horn Advisor Department of Entomology To my wife, Xin ii ACKNOWLEDGMENTS I would like to express my appreciation to Dr.. Charles A. Triplehorn, my principle advisor, who was always encouraging, supporting and understanding during my stay at The Ohio State University. I also cherish the memory of the late Dr. D. M. DeLong, my original advisor, who played a key role in choosing this project for me. OSU faculty, Drs. Norman F. Johnson and David J. Horn kindly criticized this manuscript. Drs. Paul H. Freytag, University of Kentucky, and William P. Morrison, Texas Agricultural Extension Service, have been very helpful with some questions referring to this study. Many graduate students, faculty and research scientists have given tremendous assistance and help on the systematic theory in general and microscope work enabling me to complete this project. These include: Drs. John Wilcox, Paul Cwikla, Shawn Clark, and Richard Miller. I also thank Paul W. Oman, Oregon State University, iii K. G. A. Hamilton, Biosystematics Research Institute, Canada, Robert W. Brooks, University of Kansas, Paul H. Freytag, University of Kentucky, Raymond J. Gagne, and James Kramer, U.S. National Museum (Natural History), W. J. Knight, British Museum, Frank W. Mead, Florida Department of Agriculture, Norman D. Penny, California Academy of Sciences, and H. Derrick Blocker, Kansas State University, who kindly loaned the specimens in their care for this study. A special thanks to my wife, Xin and my family, who are always supportive and understanding about my frequent absence. VITA May 9, 1955 ----------- Born - Shenyang City, P.R. China 1978 ------------------ Graduated, Shenyang Agricultural University, Shenyang City, P.R. China 1981 ------------------ M.S., N.W. University of Agriculture, Shaanxi Province, P.R. China 1981-1983 ------------- Teaching Associate, Shenyang Agricultural University, Shenyang City, P.R. China 1984-1989 ------------- Research/Teaching Associate, Department of Entomology, Ohio State University PUBLICATIONS Chou, I. and N. Ma, 1981. On some new species and new records of Typhlocybinae (Homoptera: Cicadellidae) from China. Entomotaxonomia, 1981, 111(3):191-210, Ma, N. , 1983. A new species of Tautoneura and a new record from China (Homoptera: Cicadellidae: Typhlocybinae). Entomotaxonomia, 1983, V(2):149-150. v Ma, N., 1984. A new record of Typhlocybinae from China (Homoptera: Cicadellidae). Entomotaxonomia, 1984, VI(1):157. Cwikla, P. and N. Ma, 1986. A new species of Cariancha Oman Homoptera: Cicadellidae: Deltocephalinae) with a discussion on the tribal placement of the genus. J. New York Entomol. Soc. 94(1) 115-117. Freytag, P. and N. Ma, 1987. An unusual new genus and species of Hecalini leafhoppers from Venezuela (Homoptera: Cicadellidae). Ent. News 99(3):153-156. Ma, N., 1988. A review of the genus Spangbergiella (Homoptera: Cicadellidae: Deltocephalinae). J. Kansas Entomol. Soc. 61(4):000-000. Ma, N., A revision of the genus Dicyphonia Homoptera: Cicadellidae: Deltocephalinae). In press (JKES). Ma, N., A revision of the genus Memnonia (Homoptera: Cicadellidae: Deltocephalinae) with a phylogenetic analysis of species. In press (JKES). Ma, N., Systematic and phylogenetic study of genus Hecalus (Homoptera: Cicadellidae: Deltocephalinae) of North America. In press (JKES). Ma, N., The cladistic study of of Cicadellidae subfamilies (Homoptera). In manuscript. FIELDS OF STUDY Major Field: Systematic Entomology Studies in Leafhopper Systematics vi TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................ iii VITA ........................................ v LIST OF TABLES .............................. x LIST OF FIGURES ............................. xi INTRODUCTION ................................ 1 Taxonomic History of Hecalini ......... 4 Natural History of Hecalini ........ 6 Morphology and Terminology ............ 7 Material and Methods ................... 10 CHAPTER I. SYSTEMATIC STUDY OF HECALINI ....... 12 Check list of Hecalini ........... 12 Key to genera ..................... 19 Phylogeny of Hecalini genera ..... 20 II. SYSTEMATIC STUDY OF GENUS MEMNONIA 26 key to species .................... 31 Phylogeny of Memnonia species .... 52 III. SYSTEMATIC STUDY OF GENUS DICYPHONIA ................................... 56 Key to species .................... 59 Phylogeny of Dicyphonia species ... 77 IV. SYSTEMATIC STUDY OF GENUS HECULLUS 82 Key to species .................... 83 V. SYSTEMATIC STUDY OF GENUS NEOHECALUS ................................... 87 Key to species .................... 89 VI. SYSTEMATIC STUDY OF GENUS vii SPANGBERGIELLA .................... 92 Key to species .................... 94 Phylogeny of Soangbergiella species ........................... 105 VII. SYSTEMATIC STUDY OF GENUS HECALUS ........................... 108 Key to species .................... Ill Phylogeny of Hecalus species ..... 146 LIST OF REFERENCES ........................... 151 APPENDIX A: ILLUSTRATIONS .................. 156 viii LIST OF TABLES TABLES PAGE 1. Characters and their polarized conditions of tribe Hecalini .......... 23 2. Data matrix of characters or character states of tribe Hecalini, the numbers referring to Table 1 ...... 25 3. Characters and their polarization for Memnonia species .................. 51 4. Data Matrix of Memnonia species, numbers refer to characters used in table III. O=plesiomorphic, l=apomorphic .......................... 52 5. Characters of Dicyphonia used in the cladistic analysis and their polarized conditions by outgroup comparison. Codes (6* etc.) represent the same characters but different states ...................... 79 6. Data matrix of polarized characters or character states of Dicyphonia for cladistic analysis. 1 = apomorphic state, 0 = plesiomorphic state ..... 81 7. Characters used in the phylogenetic analysis of the species of Spangbergiella and their polarized condition ...... 106 8. Data matrix of the species of Spangbergiella. l=apomorphic states, plesiomorphic states ........ 107 9. Characters used in phylogenetic study of Hecalus and their polarized conditions ................. 148 10. Data matrix of the polarized characters of Hecalus. 1 = apomorphic condition, 0 = plesiomorphic conditions ...... 150 ix LIST OF FIGURES FIGURES PAGE 1-7. Hecalus flavidus. 1) male, dorsal aspect; 2) face, ventral aspect; 3) right forewing, dorsal aspect; 4) pygofer, lateral aspect; 5) first abdominal segment and apodemes, ventral aspect; 6) connective, dorsal aspect; 7) aedeagus, lateral aspect ........................... 157 8 . Hypothesized phylogenetic relationship among species of Memnonia Ball, the numbers equal to those of characters in Table 1, 2 and x = reverse condition .................................. 159 9-15. Memnonia ornata. 9) male head, pronotum and scutellum, dorsal aspect; 10) male right forewing; 11) connective; 12) pygofer, ventral aspect; 13) apex of aedeagus; 14) aedeagus, lateral aspect; 15) male first abdominal segment .......... 161 16-22. Memnonia albolinea. 16) male right forewing; 17) male head, pronotum and scutellum, dorsal aspect; 18) pygofer, lateral aspect; 19) male first abdominal segment; 20) aedeagus, posterior aspect; 21) aedeagus, lateral aspect; 22) connective.