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Information to Users 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 ^ewriter face, while others may be from aiy type of computer printer. The quality of this reproduction is dependent upon the quali^ of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photogr^hs, print bleedthrough, substandard margins, and improper alignment can adversely afreet 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 of the book. Photogr^hs 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 aiQr photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell Information C om pany 300 North Z eeb Road. Ann Arbor. Ml 48106-1346 USA 313.'761-4700 800/521-0600 Order Number 9517035 Phylogeny, chaetotaxy, and ecology of the Onthophilinae and Tribalinae (Coleoptera: Histerdae) Kovarik, Peter William, Ph.D. The Ohio State University, 1994 UMI 300 N. Zeeb Rd. Ann Arbor, MI 48106 PHYLOGENY, CHAETOTAXY, AND ECOLOGY OF THE ONTHOPHLLINAE AND TRIBALINAE (COLEOPTERA; HISTERDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Peter William Kovarik, B.A., M.S. ***** The Ohio State University 1994 Dissertation Committee: Approv W. A. Foster D. J. Horn N. F. Johnson Co-Adviser C. A. Triplehom Department of Entomology To Horace R. Burke and Rupert L. Wenzel “The study o f entomology is one o f the mostfascinating ofpursuits. It takes its votaries into the treasure-houses of Nature, and explains some o f the wonderful series o f links whichform the great chain o f creation. It Ic^s open before us another world, o f which we have been hitherto unconscious, and shows us that the tiniest insect, so small perhaps that the unaided eye can scarcely see it, has its work to do in the world, and does it. ” Rev. J.G. W ood u Acknowledgments My initial thanks go to my advisors Dr. Norman F. Johnson and Dr. Charles A. Triplehom for their guidance, support, and camaraderie, I wish to also thank close friends and committee members Dr. Woodbridge A. Foster and Dr. David J. Horn for sharing their ideas on science and philosophy. I am grateful for the tutelage of Rupert L. Wenzel and Alfred F. Newton, who were functionary, though unoflScial, committee members. The collective editorial comments of the aforementioned people substantially improved this document. I am particularly indebted to George Keeney for key assistance in beetle rearing. Special thanks are due P. Skelley, K. Stephan, R. W. Jones, A. F. Newton, J. W. Summerlin, M. Archangelsky, W. Godwin, B. Carr, D. Verity, and R. Westcott for providing me with preserved immatures and /or culture stock. Thanks are also due K. Roach, P. Kaufl&nan, P. Lima, and D. Knott (USDA/APHIS/PPQ) for providing me with importation permits. I owe much to the following individuals for financial assistance for the period beyond my allotted departmental support: N. F. Johnson, J. Kovarik, B. Armitage, K. Summers, D. McIntyre and L. Summers. I wish to acknowledge the artistic and technical contributions of R. Hancock, S. Hakola, J. Mitchell, and J. Diaz. I also wish to thank J. Wenzel and C. Marshall for assistance with HENNIG86. Ill I am extremely grateful to Dr. James W. Kimbrough for fungal spore identifications. I wish to thank the following individuals and lending institutions for the specimens I borrowed: L. H. Herman, American Museum of Natural History, New York; E. DuBoise, British Museum of Natural History, London; E. R. Hoebeke, Cornell University, Ithaca; H. R. Burke and E. G. Riley. Texas A&M University, College Station; H. M. André, Musee Royal De L’Afiique Centrale, Tervuren; J. F. Lawrence, Australian National Insect Collection, Canberra; E. Matthews, South Australian Museum, Adelaide; R. L. Wenzel and A. F. Newton, Field Museum of Natural History, Chicago; N. Berti, Paris Museum, M. Uhlig, Humboldt University Museum, Berlin. Thanks are due Karl Stephan, C. Klopp, and J. Wenzel for translation assistance and Kim Summers for formatting and proofing this document. Finally I wish to thank Miguel Archangelsky, Brian Burrell, Flavia Ejchel, Bob Jones, Bob Hancock, Jolanda Kovarik, John Mitchell, Steve Passoa, Keith Philips, Brady Porter, Paul Skelley, Jim Stoeckel, Kim Summers, Nancy Tomei, and Bruce Weaver for moral support. IV VTTAE May 25,1957 Bom - Norwalk, CT 1975-1979 ...................................................................................................B.A. Biology Quinnipiac College Quinnipiac, CT 1979-1983 ............................................................................................. M.S. Entomology Texas A&M College Station, TX MAJOR AREA OF STUDY Entomology Table of Contents Dedication ........................................................................................................................ii Acknowledgments ...........................................................................................................iii Vita................................................................................................................................. iv List of Tables .................................................................................................................vii List of Figures...............................................................................................................viii CHAPTER I Chaetotaxy of Larval Histeridae (Coleoptera: Hydrophiloidea) based on a Description of Onthophilus nodatus LeConte ..................................................1 Introduction .......................................................................................................... 1 Materials and Methods ......................................................................................... 4 Terminology and Homology ................................................................................. 3 Discussion .......................................................................................................... 16 CHAPTER n Pupal Chaetotaxy of Histeridae (Coleoptera: Hydrophiloidea) with a Description of the Pupa of Onthophilus kimi Ross .............................................. 39 Introduction .............................................................................................................39 Materials and Methods .............................................................................................40 Results.....................................................................................................................40 Discussion ................................................................................................................45 CHAPTER m Development of Epierus divisus Marseul (Coleoptera: Histeridae) Introduction .............................................................................................................50 Materials and Methods .............................................................................................51 Results.....................................................................................................................53 Discussion ................................................................................................................56 VI CHAPTER IV A Revision of the Genera of Onthophilinae and Tribalinae (Coleoptera: Histeridae) .................................................................................................61 Introduction ............................................................................................................. 61 Materials and Methods .............................................................................................63 Terminology and Morphology ...................................................................................64 Results..................................................................................................................... 75 Phylogentic Analysis ............................................................................................... 158 CHAPTER V Chaetotaxy of the Immature Stages of the New World Idoliini (Coleoptera: Histeridae) Based on Idolia gibba Lewis and Eutribalus americamis (LeConte) ..............................257 Introduction ...........................................................................................................257 Results................................................................................................................... 258 Discussion ..............................................................................................................269 APPENDICES Appendix A ............................................................................................................ 307 Appendix B ...........................................................................................................
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