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Viewed My Progress Throughout This Study INFORMATION TO USERS This reproduction was made from a copy o f a document sent to us for microfilming. While the most advanced technology has been used to photograph and reproduce this document, the quality of the reproduction is heavily dependent upon the quality of the material submitted. The following explanation of techniques is provided to help clarify markings or notations which may appear on this reproduction. 1. The sign or “ target” for pages apparently lacking from the document photographed is “ Missing Pagets)” . If it was possible to obtain the missing page(s) or section, they are spliced in to the film along w ith adjacent pages. This may have necessitated cutting through an image and duplicating adjacent pages to assure complete co ntin u ity. 2. When an image on the film is obliterated with a round black mark, it is an indication of either blurred copy because of movement during exposure, duplicate copy, or copyrighted materials that should not have been filmed. For blurred pages, a good image of the page can be found in the adjacent frame. If copyrighted materials were deleted, a target note will appear listing the pages in the adjacent frame. 3. When a map, drawing or chart, etc., is part of the material being photographed, a definite method of “ sectioning” the material has been followed. It is customary to begin film in g at the upper left hand corner o f a large sheet and to continue from left to right in equal sections with small overlaps. If necessary, sectioning is continued again -beginning below the first row and continuing on until complete. 4. For illustrations that cannot be satisfactorily reproduced by xerographic means, photographic prints can be purchased at additional cost and inserted into your xerographic copy. These prints are available upon request from the Dissertations Customer Services Department. 5. Some pages in any document may have indistinct print. In all cases the best available copy has been film ed. University Micrdfilms International 300 N. Zeeb Road Ann Arbor, Ml 48106 8510544 Akers, Rodney Cliff REPRODUCTIVE BIOLOGY OF THE BRONZE BIRCH BORER, AGRILUS ANXIUS GORY, IN OHIO (COLEOPTERA: BUPRESTIDAE) The Ohio State University Ph.D. 1985 University Microfilms International300 N. Zeeb Road, Ann Arbor, Ml 48106 REPRODUCTIVE BIOLOGY OF THE BRONZE BIRCH BORER, AGRILUS ANXIUS GORY, IN OHIO (COLEOPTERA: BUPRESTIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Rodney Cliff Akers, B.A., M.S. ***** The Ohio State University 1985 Reading Committee: Approved By Dr. David G. Nielsen Dr. Harry D. Niemczyk Dr. Benjamin R. Stinner Department of Entomology ACKNOWLEDGMENTS I would like to express my gratitude to Professors D. G. Nielsen, H. D. Niemczyk, B. R. Stinner, C. A. Triplehorn, G. R. Stairs, and R. D. Mitchell. Drs. Triplehorn, Stairs, and Mitchell offered helpful suggestions at the initiation of this study. Drs. Niemczyk and Stinner critically re­ viewed my progress throughout this study. Their interest and concern are appreciated. Dr. Nielsen served as my committee chairman and helped in the design and implementation of this study. His persis­ tence in helping me strive for excellence, professionally and personally, was invaluable to my growth and success during this study. I wish to thank others who contributed to this work. D. Cox, C. Birk, L. Birk, L. Blockus, and M. Dunlap provided much assistance in data collection. Special thanks to Michael Dunlap for his friendship. Finally, I would like to thank my wife, Anne Trice, for editing and typing this dissertation, but above all, for her encouragement, confidence, patience, and understanding throughout the last four years. VITA Peer uary 21, 1d 53 norn - aecford, Indiana LjI o U.S., biology, Lynci'.burr Cc lie g e , i-.ynciibr.rg, Virginia i :< 7 -iy7c Gracuato Kesearch civ Teaching /issirtant, ueyarl- raont of Lnto.noiogy, V i r g i n ­ ia Polytechnic Institute and State university, uiacksourg, Virginia i i) ~hj .S . , entomology, Vir ini a Polytechnic Institute ant.; State University, alacks- burg, Virginia 1 D 7 o Extension agent, Virginia C o o p e r a t i v e l. >: t e r: a ion c. r - vice, uUck ing nan:, Virginia, I I O J or an ui; te h e s e a re., . u a e l — ate, Department of Lnto;.cl­ og y , on io .Ig r icuitu :a i desearcr. an... Development Center, 1 ne ukio state University, coo; ter, unio Ph.D., n n toir.c- log y, Tne on io State University, Colu...bus, on io i i i TABLE OF CONTENTS Page ACKNOWLEDGMENTS i i VITA.................................................... iii LIST OF TABLES.......... .............................. vi LIST OF FIGURES........................................ ix INTRODUCTION........................................... I LITERATURE REVIEW..................................... 5 EXPERIMENTATION........................................ 20 I. Influence of Post-Felling Treatment of Birch Logs on Adult Emergence............ 20 Materials and Methods......................... 20 Results and Discussion........................ 21 II. Spatial Emergence Patterns from European White Birch................... 29 Materials and Methods........................ 29 Results and Discussion....................... 30 III. Predicting Adult Emergence by Heat Unit Accumulation............................ 40 Materials and Methods........................ 40 Results and Discussion....................... 41 IV. Host and Host Quality Influences on Reproductive Biology...................... 57 Materials and Methods............. 57 Results and Discussion....................... 62 iv Page V. Mating Behavior............................. 82 Materials and Methods........................ 82 Results and Discussion....................... 85 DISCUSSION........................................... 93 SIGNIFICANT FINDINGS................................ 101 APPENDICES............. .............................. 10 2 A. Leaf Collection and Extraction................ 102 B. Determination of Total Phenols by Folin-Denis Method.......................... 103 C. Determination of Hydrolysable Tannins by KIO4 Method.................................. 10 4 D. Determination of Condensed Tannins by HC1 in n-BuOH Method................................ 10 5 E. Determination of Tanning Coefficient by Protein Precipitate Method................. 106 F. Determination of Total Proteins by Bradford Method........................................... 107 LIST OF REFERENCES................................... 108 v LIST OF TABLES TABLE PAGE 1 Analysis of variance of factors and interactions influencing A. anxius emergence success under laboratory conditions, 1981 to 1983........................ 22 2 Influence of felling date on A. anxius emergence under laboratory conditions.......... 24 3 Influence of bolt end treatment on A. anxius emergence under laboratory conditions.......... 25 4 Influence of weeks of cold storage on A. anxius emergence under laboratory conditions.. 28 5 Comparison of A. anxius, parasite, wood­ pecker predation, and total holes from B. pendula vs height near Wooster, OH............. 32 6 Comparison of A. anxius. parasite, wood­ pecker predation, and total holes from B. pendula vs circumference near Wooster, OH 33 7 Comparison of A. anxius, parasite, wood­ pecker predation, and total holes from B. pendula vs bark thickness near Wooster, OH.... 34 8 Comparison of A. anxius, parasite, wood­ pecker predation, and total holes from B. pendula vs compass direction near Wooster, OH. 36 9 Comparison of coefficient of variation for 10% A. anxius emergence in Ohio. Eight base temperatures and 4 starting dates were used for calculations........................... 47 10 Julian dates of actual 10% A. anxius emergence compared to predicted emergence based on °D and linear regression........... 53 v i TABLE PAGE 11 Julian dates of actual and predicted 10% emergence of A. anxius adults in Ohio localities and years not incorporated into predictive models........................... 54 12 Julian dates of actual 10% A., anxius emergence compared to recommended spray date.. 56 13 Longevity and length of maturation feeding period of A. anxius on selected trees near Wooster, OH, during 1980 through 1983............................................... 63 14 Host influnces on A. anxius fecundity and egg incubation period near Wooster, OH, during 1980 through 1983........................ 65 15 Foliage consumption per A. anxius adult on selected hosts near Wooster, OH, in 1983........ 68 16 B . pendula leaf treatment effects on A. anxius adult longevity in 1983 near Wooster, OH........ 71 17 B . pendula leaf treatment effects on A. anxius adult reproductive biology in 1984 near Wooster, OH....................................... 72 18 B . pendula leaf traits assayed on 3 sampling dates during 1984 . Values are X ± S.E.......... 74 19 Influence of cage movement on A. anx ius reproductive biology in 1984.................... 80 20 Percent of A. anxus beetles copulating at different ages under laboratory conditions near Wooster, OH................................. 86 21 Duration of copulation and time prior to first copulation of naive A. anxius adults under laboratory conditions near Wooster, OH......... 88 22 Comparison of copulation duration and time prior to copulation between naive and 3-min- interrupted A. anxius adults under laboratory conditions near Wooster, OH..................... 90 v i i TABLE PAGE 23 Influence of mating frequency on longevity, fecundity,
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