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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 of computer printer. The quality of this reproduction is dependent upon the quality of the 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 of the 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. 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 9130523 A revision of the Leptolycini (Coleoptera:Lycidae) with a discussion of paedomorphosis Miller, Richard Stuart, Ph.D. The Ohio State University, 1991 Copyright ©1991 by Miller, Richard Stuart. All rights reserved. UMI 300 N. Zccb Rd. Ann Arbor, MI 48106 A REVISION OF THE LEPTOLYCINI (COLEOPTERA: LYCIDAE) WITH A DISCUSSION OF PAEDOMORPHOSIS DISSERTATION Presented in Partial Fulfillment of the Requirements f the Degree Doctor of Philosophy in the Graduate School of The Ohio State University by Richard S. Miller, B.A., M.S. ***** The Ohio State University 1991 Dissertation Committee: Approved By: Dr. David J. Horn tA Dr. Donald E. Johnston Dr. Charles A. Triplehorn Advisor Dr. Barry D. Valentine Department of Entomology Copyright by Richard S. Miller 1991 I To Dr. Roy A. Crowson for his study and his provocative queries i i ACKNOWLEDGEMENTS How dull it is to pause, to make an end, To rust unburnished, not to shine in use! As though to breathe were life itself! -A. Tennyson, 1833. The immensity of biology can either elate or overwhelm the student. There are, essentially, two responses to this conundrum. One is the quixotic charge here and there at the fog of unexplained phenomena. The other is to succumb to a Faustian despair of ever encompassing the morass of little suspected knowledge. While fully appreciating the absurdity, I chose the former when most of my cohort were increasing their biological fitness (sensu Colinveaux, 1982:394). This current milestone in my studies has given me pause to reflect on those who have contributed to my formal and informal education. It is impossible to differentiate adequately between them and, thereby, classify them - for they have made varying contributions of infinitely subtle distinction. A "lumper" might thank Western Civilization as probably more than 997. of my thoughts are a result of inculcation from this source. A "splitter" would begin a list of individuals. Aristotle, a member of Western Civilization, has urged moderation and, so, I will mention a few stellar contributions. Of course Niki comes to mind first. Besides her readily apparent artistic abilities demonstrated in this dissertation, her support and encouragement have significantly contributed to its conclusion and my i i i happiness. Again, my family has been a source of support, even though they kindly find my interests "different." I thank them for everything. My friends and former office- mates of B & 2 309 have been inspirational in our discussions. These peers include Michael A. Ivie, William F. Abeles, James B. Stribling, Paul S. Cwikla (really only 309-A), and recently Peter W. Kovarik. I thank Nick Calderone, an honorary member of our clique, who has done much to enrich my appreciation of philosophy and other arcane subjects. The various members of the faculties of CSUS, UCD, and OSU who have contributed to my education are appreciated. Especially important among these are Harold Wiedmann and Rollo Darby; Martin C. Birch; Charles A. Triplehorn, Barry D. Valentine, David J. Horn, Donald E. Johnston, Norman F. Johnson, and J. Bruce Griffing respectively. The staff of the OSU biology library has been immensely helpful. I would like to thank them all, and especially Susan Ward, for their patience, understanding, and assistance. Several of ray colleagues around the world have assisted in various aspects of my work including the loan of specimens. The latter are listed in the materials and methods section, although I would like to acknowledge all here. This paragraph is reserved for my reading committee should they decide to pass me after the laborious task of reading this manuscript. Additionally, I would like to thank Dr. David Horn for his question during my general exam on testability by use of hypothesized phylogenies. It was seminal to some of the ideas developed here. iv Finally, I thank all those natural historians and scientists, whether trained or untrained, who for whatever motives have left their thoughts in print. If I have been remiss in failing to cite their work, I here apologize and can only plead the finite limitations of a human mind. History <ie. pattern) is not intrinsically useful unless its lessons reveal process. It cannot explain process. Hence, the validity of a phylogenetic hypothesis is not as important as the testable questions it compels. This dissertation gently asks how ontogeny relates to beetle evolution and the significance of paedomorphosis in cantharoid evolution. I hope this relationship of ontogeny and phyiogeny can be examined more cogently by providing this arena for further study. If I have failed, the enclosed cladograms must be viewed merely as art to be stored in some attic. Art is the culmination of human imagination, but it was not my purpose. v VITAE 15 February 1945 .......... Nativity, Los Angeles, California 1970 — 1976 . ............ Bachelor of Arts in Biology with a concentration in Zoology, California State University, Sacramento 1975 - 1976.............. University of California, Davis 1976 - 1981.............. Economic Entomologist I- III, California Department of Food and Agriculture, Sacramento 1981 - 1985 .............. Master of Science in Entomology, The Ohio State University, Columbus 1988 -1989 ............... Insect Identifier, United States Department of Agriculture 1985 - present ........... Ph.D. in Entomology, The Ohio State University, Columbus PUBLICATIONS 1984. The Buprestidae of the Virgin Islands. Florida Entomologist 67(2 ):288“300. (co-authored with M. A. Ivie) 1988. The behavior of Calooteron reticulatum (F. ) larvae (Coleoptera: Lycidae). Ohio Journal of Science 88(3 ):115— 116- FIELDS OF STUDY Major Field: Entomology vi TABLE OF CONTENTS ACKNOWLEDGEMENTS ............................................................ iii VITA ........................................................................... vi TABLE OF CONTENTS ............................................................ ix LIST OF TABLES ............................................................... xi LIST OF FIGURES ............................................................. xi i INTRODUCTION ................................................................... 1 MATERIALS AND METHODS ........................................................ 3 Methods .................................................................... 3 Systematic analysis ............................................... 3 Taxonomic conventions ............................................. 6 Paedomorphosis ...................................................... 7 Material studied .......................... .............................. 9 ONTOGENY AND PHYLOGENY ...................................................... 12 Introduction .................................................. 14 Definition of the problem ....... 18 An example ................ 19 A model . ................................................................ 30 Synergism of morphology and systematics ............................. 34 Phylogenetic implications ............................... 37 Possible taxa for further study ...................................... 40 PHYLOGENETIC RELATIONSHIPS WITHIN THE CANTHAROIDEA .................... 45 vi i Relations within the Elateriformia ................... 47 Monophyly of the Cantharoidea............ 49 Current hypotheses of Cantharoid relationships ............ 50 Family relations within the Cantharoidea ........................... 51 Omalysidae ......... .................................................. 51 Plastoceridae ................................... 53 Cantharidae .......... 54 Omethidae ........................................... ................... 55 Phengodidae ................................. 55 Telegeusidae ........................................................... 57 Dri 1 idae .............................................. 57 Lampyridae ........................................................ 58 Lycidae ...................................................... 59 THE TRIBE LEPTOLYCINI .....................................................