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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 (fid not send UMI a complete manuscript and there are missing pages, these wilt 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. Photographs included in the original manuscript have been reproduced xerographicaity 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. ProQuest Information and Learning 300 North Zeeb Road. Ann Arbor, Ml 48106-1346 USA 800-521-0600 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. THE EVOLUTION OF LAMPYRIDAE, WITH SPECIAL EMPHASIS ON THE ORIGIN OF PHOTIC BEHAVIOR AND SIGNAL SYSTEM EVOLUTION (COLEOPTERA: LAMPYRIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for The Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Marc A. Branham The Ohio State University 2002 Dissertation Committee: Approved by Dr. John W. Wenzel, Advisor Dr. Norman F. Johnson w. Co. Sl Dr. Woodbndge A. Foster Advisor ^ Dr. Douglas A. Nelson Department o f Entomology Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. UMI Number 3048999 Copyright 2002 by Branham. Marc Alexander All rights reserved. UMI* UMI Microform 3048999 Copyright 2002 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17. United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor. Ml 48106-1346 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Copyright by Marc A. Branham 2002 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT The distribution, life history, and behavior of both larval and adult Lampyridae are summarized, as well as larval and adult morphological characters that define this family of beetles. The last larval and pupal stages of the North American firefly, Lucidota atra (G~A~ Olivier 1790), are described and illustrated. The larva of L atra was misidentified in the literature as a species in the genus P hotinus. A discussion of the homology of abdominal sclerites in larval, pupal, and adult fireflies is provided. This work represents the first cladistic analysis of genera in the family Lampyridae and other closely related beetles. A monophyledc concept of Lampyridae is established. The phylogenetic positions of the luminous cantharoid families [Omalisidae, Rhagophthalmidae and Phengodidae] in relation to Lampyridae are discussed, as well as the implications of the evolution of bioluminescence and photic s ign alin g in this group of beetles. The Rhagophthalmidae appears to include D ioptom a and Diplocladon (formerly located in Phengodidae) and the Phengodidae apparently includes Stenocladius (formerly of Lampyridae). Harmatelia, Drilaster and P terotus are transferred to Elateroidea m certae seeds and not included in Lampyridae where they were sometimes placed. Through a phylogenetic analysis using adult morphological characters, it is shown that ii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. the origin ofbioluminescence in cantharoid beetles appears to predate the origin o f the family Lampyridae. The ability to produce and emit photic signals was first gained by larvae and appears to function as an aposematic warning display, while subsequently being gained in adults and used as sexual signals. This analysis also suggests that while pheromonal sexual signals are used basally in the fam ily, these are used in conjunction with, and then subsequently replaced by, photic signals in some Iampyrid lineages. Both photic signals and the photic organs used to produce them have become greatly elaborated in the fireflies that no longer employ pheromonal sexual signals. In addition, the ability to produce a flashed sexual signal appears to have arisen at least three times in the family Lampyridae. Convergent evolution also is evident in adult male photic organ morphology. Further, it is recommended that individual signal-system components be compared, rather than overall signal system complexity. The use of this strategy may allow one to recognize and better interpret adaptive correlations despite convergence or loss. ' Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. To Frank A. McDermott, a pioneer in the study ofLampyridae and a man I would have liked to have met “Bat there is no ram tonight no clouds, in fact to blot out the showering of starlight The moon has not risen from its sluggish daybed, and the only competition offered the stars in this clean night air is from lightning bogs over the drtchwater. They are leisurely about sharing their miracle, never overdoing the effect - one here, then another off in the distance, a fluid interval and then three airborne grace notes of yellow rr Blue Rise By Rebecca Hill hr Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS I would like to acknowledge the members of my doctoral committee. First and foremost, I am indebted to my advisor Dr. John Wenzel for continually challenging me to look a little deeper and think cladistically. He is a wonderful mentor and Mend. I am still amazed that he was brave enough to take on a student who studied not wasps, but beetles. And to John’s credit, he has learned many difficult Iampyrid and cantharoid names. He now knows a lot about beetles, for a hymenopterist! Dr. Norman Johnson’s impressive knowledge of nomenclature, languages, geography and entomology has been invaluable to me. One day 1 was in his office some four times... I appreciate his patience and willingness to be of help. Dr. Woodbridge Foster is one of my favorite instructors. His attention to detail is superhuman and his knowledge so broad that conversations with him always leave me thinking. Dr. Douglas Nelson always encouraged me to think about communication systems in the broadest terms, rather than only thinking about insects. It is, of course, always important to place one’s work within a larger context. Doug’s quiet and careful ways provide an excellent example of professionalism in science. Dr. James Lloyd is both a Mend and mentor. I treasure the time I spend looking out over a field of flashing fireflies with Jim pointing out this and that, or just talking V Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. over beers about firefly staff nr fishing-1 often feel like Jim is the only one I can talk to about the nitty gritty details of fireflies. Jim and his wife Dorothy have generously put me up in a spare bedroom in their home during my visits to Gainesville, FL. They have always treated me like family and I enjoy their company very much. I would like to thank Dr, Hans Klompen and Dr. Peter Kovarik for our discussions concerning morphology and the morphological similarities (though subtle) between mites, histerid beetles and fireflies. The following friends and colleagues have supplied particularly helpful ideas, criticism, and friendship: Keith Philips, Todd Blackledge, Hojun Song, Chi Feng Lee, and Ming Luen Jeng. I’d like to give special thanks to Miguel Archangelsky for teaching me about scientific illustration and how to keep beetle larvae alive and happy in the lab. Kurt Pickett should also receive special mention, as we arrived in the Wenzel Lab together and were partners-in-crime for my entire stay at OSU. Kurt has always been willing to help, listen, argue, and proofread whenever the need arose. I would also like to gratefully acknowledge Mr. Bruce Leech, Ms. Susan Ward and Ms. Karen Ronga, librarians at the Biological and Pharmacy Library, Ohio State University, for their willingness to help me track down obscure references that were vi Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. oftentimes incorrectly cited. I often wondered if I could come up with a Iibrary-research related problem that they could not solve; I never could. They are experts and I appreciate them skills. I would like to thank my wife Lori Sandhoidt and my brother Aric Branham for their patient tolerence of my entomological endeavors. I would also like to thank my mother, who never minded my filling up her freezer with bug jars when I was a boy. I would