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Z eeb Road, Ann Arbor, Micfiigan 48106 8618856 Stribling, James Bentley WORLD GENERIC REViSION OF PTILODACTYLIDAE (COLEOPTERA;DRYOPOIDEA) The Ohio State University Ph.D. 1986 University Microfilms I nternsti 0 nâl SOO N. zeeb Road, Ann Arbor, Ml 48106 WORLD GENERIC REVISION OF PTILODACTYLIDAE (COLEOPTERA: DRYOPOIDEA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By James B. Stribling, B.S., M.S. ***** The Ohio State University 1986 Dissertation Committee: Approved by C. A. Triplehorn N. F. Johnson Û- Adviser D. E. Johnston Department of Entomology B. D. Valentine Copyright by James Bentley Stribling 1986 To My Parents ii ACKNOWLEDGEMENTS There are many people to whom I am very grateful for academic, moral, physical, and financial support during my graduate career at The Ohio State University: my adviser. Dr. Charles A. Triplehorn, for sound advice for the duration and understanding when I needed it most; Dr. Norman F. Johnson for numerous discussions, suggestions, and welcome advice on systematic theory and taxonomic procedure; my officemates, colleagues, and fellow graduate students, especially, Paul S. Cwikla, Michael A. Ivie, Richard S. Miller, and John A. Shuey, for seemingly unending discussions on systematics, for companionship on collecting trips, and for tolerance during these last few months; John F. Lawrence for much advice and comment on this as well as smaller simultaneous projects; W. Calvin Welbourn for many hours spent in setting up the AT&T 3B2 and QTREE and then tutoring me on their use; John F. Lawrence, Charles W. O'Brien, and Arthur J. Gilbert for gifts of specimens; Roland L= Seymour (Department of Botany, The Ohio State University) for fungal identifica­ iii tions; Ho-Yih Liu (also Department of Botany, 0. S. 0.) for translating some of Nakane's works from Japanese; all of the curators in the "Materials" section for giving me the opportunity to study their specimens (even past expiration dates on loan forms); Niki Lowell and Adam Rubinstein for the habitus illustrations (Figures 3 (N. L«) and 1 and 2 (A. R.); and Helen W. Monk for typing part of the manuscript. Partial funding of museum trips was provided by Sigma Xi (Grant-in-Aid of Research), The Ohio State University Graduate School (Graduate Student Alumni Research Award), and the Knull Fund, Department of Entomology. I am very grateful to them. I owe immense thanks to my wonderful parents. Dr. and Mrs. Loutrelle, for their financial support, endless moral support, encouragement, and for all of those times I needed "a good talkin' to" and got it. Finally, I thank my wife, Sally, for the hours of assistance she has given me in my research, taking time out of her own to process specimens, to type, or to take over household duties when I might be nearing a deadline. My greatest thanks to her are for her unwavering love, encouragement, and companionship. IV VITA October 13, 1958.............. Born - Jackson, Mississippi 1980.................. .. .B. S., The University of Mississippi, Oxford, Mississippi 1982 # ••••«•••••• # #M# S #, The Ohio State University, Columbus, Ohio 1980-present..................Graduate Research/Teaching Associate, Department of Entomology, The Ohio State University PRESENTATIONS 1985. Phylogenetic Implications of Metathoracic Wing Venation in Dryopoidea (Coleoptera). Annual Meeting of Entomological Society of America, Hollywood, Florida. The Biology and Systematics of Anchvtarsus (Coleoptera: Ptilodactylidae). Annual Meeting of the North Central Branch, Entomological Society of America, Lexington, Kentucky. The Biology, Phylogeny, and Zoogeography of the Wing-Toe Beetles (Ptilodactylidae). Departmental seminar. Department of Entomology, The Ohio State University, Columbus. 1984. The Faces of Systematic Biology. Invited seminar. Department of Biology, The University of Mississippi, Oxford. 1981. The New World Genera of the Ptilodactylidae (Coleoptera). Annual Meeting of the Entomological Society of America, San Diego, California. PUBLICATIONS 1986. Revision of Anchvtarsus (Coleoptera: Dryopoidea) and a key to the New World genera of Ptilodactylidae. Ann. Entomol. Soc. Amer. 79: 219-234. 1984. Taxonomic and nomenclatorial notes on Caribbean Tropicus Pacheco (Coleoptera: Heteroceridae). Proc. Ent. Soc. Wash. 86 (4): 946-950. (with M. A. Ivie) Report on the LeConte symposium on the classification and phylogeny of the Coleoptera, Detroit and Columbus, 1983. Coleop. Bull. 38 (1): 98-102. (with M. A. Ivie) FIELDS OF STUDY Major Field: Systematic Entomology with specialization in Coleoptera biosystematics. VI TABLE OF CONTENTS ACKNOWLEDGEMENTS.................................... iii VITA........................................ V LIST OF TABLES......................... ix LIST OF FIGURES............................ x CHAPTER PAGE I. INTRODUCTION...................................1 II. METHODS AND MATERIALS .........................4 Methods Phylogenetic...................... 4 General........................... 8 Material............................... 11 III. NATURAL HISTORY..............................16 IV. CHARACTERS, POLARIZATION, PHYLOGENETIC ANALYSIS, AND CLASSIFICATION........... 19 Monophyly of the Ptilodactylidae. 19 Character discussion................... 19 H e a d ................... 20 Thorax........................... 23 Abdomen........................... 28 Character polarization................. 29 Discussion of phylogenetic analysis and resulting classification ..... 32 V. TAXONOMY......................................38 Key to World Genera of Ptilodactylidae (Adults)......................... 38 The World Genera of Ptilodactylidae Anchvtarsus Guerin-Meneville . 42 vii Cladotoma Westwood .............. 44 Paraiichas White ................ 47 Hovactvla Fairmaire. ....... 50 Pseudocladotoma Pic. ....... 52 Octoqlossa Guérin-Méneville. 54 Bradvtoma Guérin-Méneville .... 57 Aploqlossa Guérin-Méneville. 60 Anchycteis Horn................... 63 Epilichas White................... 66 Bvrrocrvptus Broun .............. 69 Araeopidius Cockerell............. 72 Stenafricus NEW GENUS............. 74 Daemon de Laporte........ .. 77 Falsotherius P i c ............... 8 1 Pseudoepilichas Armstrong and Nakane ................................. 83 Ectvphodactvla NEW GENUS ..... 85 Ptiloderes NEW G E N U S ............. 89 Pherocladus Fairmaire. ...... 91 Lachnodactvla Champion .......... 94 Ptilodactvla Illiger . ......... 97 Stirophora Champion.............. 118 Chelonariomorphus Pic............ 121 Epiptyqma NEW GENUS. ........ 123 Therius Guerin-Mènev i l i e ........ 126 Incertae sedis....................... 129 Exclusions. .................. 129 LITERATURE CITED.................................. 130 APPEmiX A. Figures ................................. 144 Vlll LIST OF TABLES TABLE PAGE 1. Fungal spores and hyphae identified from gut content of some species of Ptilodactlidae. 18 2. Hypothesized polarities of characters used in phylogenetic analysis of the genera of Ptilodactylidae............ 35 3. Phylogenetic hypothesis resulting from QTREE analysis of characters in Table 2. Reversals are indicated by dots at the upper right hand side of the character numbers; characters with multiple apomorphic appearances are 2, 3, 4, 5, 7, 9, 11, 12, 13, 14, 15, 16, 18, 20, 23, 25, 27, 29, 31, 35, and 3 8 ................................... 36 4» Proposed supraspecific classification of the Ptilodactylidae based on phyletic sequencing • 37 ix LIST OF FIGURES FIGURES PAGE 1. Dorsal habitus. Anchvcteis velutina, male............... 146 2. Dorsal habitus, Ptilodactvla militaris, male. ............................... 148 3. Dorsal habitus. Ptiloderes sp., female. » . 150 4. Serrate antenna; Bradvtoma lineata........... 152 5. Ramous antenna, nonarticulated rami; Paralichas q u e r i n i .......... 152 6. Ramous