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IM w sity Microfilms International V lalj* 1.0 Itt _ to. y£ 22 £ b£ 2.0 1.1 UL 11.25 111 1.4 1.6 MICROCOPY RESOLUTION TEST CHART NATIONAL BUREAU OF STANDARDS STANDARD REFERENCE MATERIAL 1010a (ANSI and ISO TEST CHART No. 2) University Microfilms Inc. 300 N. Zeeb Road, Ann Arbor, MI 48106 INFORMATION TO USERS This reproduction was made from a copy of a manuscript sent to us for publication and microfilming. While the most advanced technology has been used to pho­ tograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. Pages in any manuscript may have indistinct print. In all cases the best available copy has been filmed. The following explanation of techniques is provided to help clarify notations which may appear on this reproduction. 1. Manuscripts may not always be complete. When it is not possible to obtain missing pages, a note appears to indicate this. 2. When copyrighted materials are removed from the manuscript, a note ap­ pears to indicate this. 3. Oversize materials (maps, drawings, and charts) are photographed by sec­ tioning the original, beginning at the upper left hand comer and continu­ ing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or in black and white paper format.* 4. Most photographs reproduce acceptably on positive microfilm or micro­ fiche but lack clarity on xerographic copies made from the microfilm. For an additional charge, all photographs are available in black and white standard 35mm slide format.* *For more information about black and white slides or enlarged paper reproductions, please contact the Dissertations Customer Services Department. Dissertation U-M-I Information Service University Microfilms International A Bell & Howell Information Company 300 N. Zeeb Road, Ann Arbor, Michigan 48106 8618850 Shuey, John Allen THE ECOLOGY AND EVOLUTION OF WETLAND BUTTERFLIES WITH EMPHASIS ON THE GENUS EUPHYES (LEPIDOPTERA: HESPERIIDAE) The Ohio State University Ph.D. 1986 University Microfilms International 300 N. Zeeb Road, Ann Arbor, Ml 48106 PLEASE NOTE: In all cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark V . 1. Glossy photographs or pages_____ 2. Colored illustrations, paper or print ______ 3. Photographs with dark background_____ 4. Illustrations are poor copy______ 5. Pages with black marks, not original copy______ 6. Print shows through as there is text on both sides of page_______ 7. Indistinct, broken or small print on several pages. 8. Print exceeds margin requirements______ 9. Tightly bound copy with print lost in spine_______ 10. Computer printout pages with indistinct print______ 11. Page(s)___________ lacking when material received, and not available from school or author. 12. Page(s) ___________ seem to be missing in numbering only as text follows. 13. Two pages numbered . Text follows. 14. Curling and wrinkled pages______ 15. Dissertation contains pages with print at a slant, filmed as received_________ 16. Other________________________________________________________________________ University Microfilms International THE ECOLOGY AND EVOLUTION OF WETLAND BUTTERFLIES WITH EMPHASIS ON THE GENUS EUPHYES (LEPIDOPTERA: HESPERIIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Doctor of Philosophy in the Graduate School of The Ohio State University 1986 By John Allen Shuey, B.S., M.S. * * * The Ohio State University Reading Committee: Approved By: Dr. Gordon Stairs A r Dr. David Horn Adviser Dr. Norman Johnson Department ofEntomology Dr. Donald Johnston ACKNOWLEDGMENTS I express sincere appreciation to; Dr.s G. Stairs, D. Horn, N. Johnson, and D Johnston for their guidance and suggestions during the course of this research; Mr.s m. Neilsen, J. Calhoun, and L. Martin for contributing data from wetland sites; Dr. D. Iftner, for helping collect some of the data herein presented and for accompanying me, along with Dr. J. Peacock, on collecting expeditions to South Florida; Dr. J. Briggs for the use of his computer; and Dr. R. Hall for statistical advice. The following curators and institutions lent material for the systematic portion of this research and I wish to thank them for their assistance; Dr.s F. Rindge and K. Johnson, The American Museum of Natural History; Dr. J. Burns, The National Museum of Natural History; Dr. G. Ekis, The Carnegie Museum of Natural History; Dr. D Bowers, Museum of Comparative Zoology, Harvard University; Dr. C. Triplehorn, The Ohio State University; and, Dr. J. Franclemont, Cornell University. The following organizations provided access to and information about wetlands under their care; The Ohio Department of ii Natural Resources, Division of Natural Areas and Preserves; The Nature Conservancy, Ohio Chapter; The Toledo Metropark System; and the Indiana Department of Natural Resources. The Following graduate students of the Department of Entomology made my stay here pleasant and influenced my research greatly; James (Sam) Stribling, Sally Rockey, Erik Neff, Mike Hamerski, Mike Ivie, Rich Miller, Paul Cwikla, Ed Larue, Dave Bezanson, and Norm Riekenbach. Finally, I wish to express my deepest and most sincere appreciation to Judy for her support, understanding, guidance and love during the three plus years that expired during this adventure. iii VITA March 29, 1958 ............... Born - Zanesville, Ohio 1980 B.S., Ohio University, Athens, Ohio 1982 M.S., Ohio University, Athens, Ohio PUBLICATIONS Shuey, J. A. 1983. An annotated checklist of the butterflies of Athens County, Ohio. Ohio J. Sci. 83: 262-269. Shuey, J. A. 1984. A bilateral sexual mosiac of Mitoura grvneus (Lepidoptera: Lycaenidae). J. Lepid Soc. 38: 144-146. Shuey, J. A. 1985. Habitat associations of wetland butterflies near the glacial maxima in Ohio, Indiana, and Michigan. J. Research Lepid. 24: 176-186. Shuey, J. A. and J. W. Peacock, 1985. A bilateral gynandromorph Celastrina ebenina (Lycaenidae) J. Research Lepid. 24: 192-193. FIELDS OF STUDY Insect Ecology ..................... Dr.G. Stairs Dr. D. Horn Biosystematics Dr. N. Johnson Dr. D. Johnston TABLE OF CONTENTS ACKNOWLEDGMENTS........................... ii VITA ........................................ iv LIST OF T A B L E S ............................. vii LIST OF FIGURES ...............................viii CHAPTER PAGE I. GENERAL INTRODUCTION ................. 1 Introduction ..................... 1 The Wetland Community ............. 4 Objectives ......... • • • 12 II. HABITAT ASSOCIATIONS OF WETLAND BUTTERFLIES NEAR THE GLACIAL MAXIMA IN OHIO, INDIANA, AND MICH IGAN ............. 14 Introduction ..................... 14 Material and Methods ............... 17 R e s u l t s .............................. 23 D i s c u s s i o n ..........................28 III. PATTERNS OF RESOURCE UTILIZATION IN A GUILD OF WETLAND HESPERIIDAE (LEPIDOPTERA) . ..................... 43 Introduction .................. 43 Materials and Methods . ■............. 47 R e s u l t s .............................. 51 D i s c u s s i o n ..........................57 v IV THE PHYLOGENETIC POSITION OF AROTIS 67 Introduction ..................... 67 Materials and Methods ............. 68 R e s u l t s .......... 70 Character Analysis ............... 76 D i s c u s s i o n ......................... 79 V. THE PHYLOGENY AND BIOGEOGRAPHY OF EUPHYES . ........................... 85 Introduction ..................... 85 Materials and Methods ............. 85 Character Analysis ............... 86 Phylogenetic Analysis ............. 104 Biogeographic Considerations . 108 VI. THE ECOLOGY OF EUPHYES AS IT RELATES TO PHYLOGENY .......................... 113 Introduction ...................... 113 Ecology and its Relationship to Phylogeny...........................114 LIST OF R E F E R E N C E S .................. 128 APPENDIX I ......................... 135 vi LIST OF TABLES TABLE PAGE 1. Descriptions and locations of wetland sites discussed ............... 18 2. Wetland butterfly species recorded from wetland si t e s .................... 27 3. Wetland butterflies and their habitat associations as recorded in this study ............................... 30 4. Estimated total numbers of flowers (= resources) present at Mud Lake in 1983 and 1984, relative percentages of resources composed of. flowers from each resource species, total flower minutes for each resource species, and relative percent flower minutes for each resource species ........... 54 5. Non-random resource utilization by hesperiids at Mud L a k e ............. 56 6. Electivity measures (D^j) of hesperiid utilization for each resource species demonstrating the differential utilization patterns between hesperiid s p e c i e s ............................. 58 7. 2 X N contingency tables between the patterns of resource utilization by the hesperiid s p e c i e s ............... 61 8. Character matrix for Euphves Species . 88 vii LIST OF FIGURES FIGURE PAGE 1. Approximate locations of wetland complexes discussed ................... 21 2. The distribution of Poanes spp. at Mud Lake relative to the Carex and Carex-Typha mats ..................... 25 3. The hypothesized phylogeny of Satvrodes. 34 4. A tentative hypothesis of phylogeny in the species group of Poanes that contains P^ massasoit and P^. viator. 38 5. A tentative phylogenetic hypothesis of Epidemia....................
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