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University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left handcorner of a large sheetand to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St. John's Road, Tyler's Green High Wycombe, Bucks, England HP10 0HR 7 9 0 2 2 3 3 STQFFER» RICHARD LAWRENCE A COMPARATIVE BEHAVIORAL STUDY OF THE DIPTERA FAMILY CHIRONOMIDAE. • THE OHIO STATE UNIVERSITY• PH»D*« 1978 University M icrofilm s international 300 n zeeb r o a d, a n n a r b o r, mi 4bio6 © Copyright by Richard Lawrence Stoffer 1978 A COMPARATIVE BEHAVIORAL STUDY OF THE DIPTERA FAMILY CHIRONOMIDAE DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University BY Richard Lawrence Stoffer, B.S., M.S. The Ohio State University 1978 Reading Committees Approved By Dr. Barry D . Valentine Dr. N. Wilson Britt Dr. David H. Stanshery '.M \. v iisor Department of Zoology ACKNOWLEDGMENTS I am indebted to a number of people who have influ­ enced this study. To my advisor Dr. Barry D. Valentine I owe much for his guidance and confidence in advisees to encourage original research. I thank Dr. Donald E. Johnston for his assistance with the NTSYS computer program; Dr. James E. Sublette of Eastern New Mexico University for his time spent checking the identification of the Tanytarsini; and Dr. Ole A. Saether, formerly of the Freshwater Institute in Canada, for his verification of several Orthocladiinae. To Dr. Saether, Dr. Andrew L. Hamilton, also of the Freshwater Institute, and Dr. D. R. Oliver of the Biosystematics Research Institute in Canada, I am indebted for the use of their unpublished keys to Chironomidae. I also thank Water Resources for the use of one of their Ekman dredges. I thank Ohio State University and the Zoology Department for their financial assistance, mainly in the form of a University Fellowship and Teaching and Research Associateships. I wish to thank the reading committee for their time and comments concerning the preparation of this work. To my fellow graduate students, especially those who shared room 373 BZ , I am indebted for their friendship, intellectual discussion, and many cherished remembrances. Most of all I wish to acknowledge my wife Cathy for her assistance in the collection of specimens and data, her editorial and typing time on this manuscript, and her patience during the course of this study. VITA December 13, 19^8.... Born - Cleveland, Ohio' 1970.................. B .S ., Ashland College, Ashland, Ohio 1972-1976............ University Fellowship, The Ohio State University, Columbus, Ohio 1975............... M.S., The Ohio State University, Columbus, Ohio 1978.................. Research Assistant, Zoology Department, The Ohio State University, Columbus, Ohio TABLE OF CONTENTS Page ACKNOWLEDGEMENTS ii VITA iv LIST OF TABLES vii LIST OF FIGURES ix LIST OF ABBREVIATIONS xi INTRODUCTION 1 Survey of Behavioral Systematic Literature 2 General Information About Chironomidae 12 Larval Lifestyles 13 Larval Feeding Behavior 19 Larval Irrigation 24 Larval Grooming 25 Behavior Associated with the Pupae 25 Fore Leg Positioning in Adults 27 Adult Grooming Behavior 27 STATEMENT OF PURPOSE 30 METHODS AND MATERIALS 31 Collection and Sorting of Specimens 31 Rearing of Specimens 35 Observation of Specimens 39 v Page Mounting Procedures 44 Analysis of Data 48 TAXONOMIC TREATMENT 50 RESULTS 68 Larval Habitation 68 Larval Feeding Behavior 83 Larval Irrigation 85 Larval Grooming 88 Behavior Associated with the Pupae 89 Fore Leg Positioning in Adults 91 Adult Grooming Behavior 106 ANALYSIS OF DATA AND CONCLUSIONS 125 Selection of Characters 125 Character Analysis 136 Comparison of Taxa 144 Discussion of Behavioral Relationships of Taxa 160 Significance of this Study 167 REFERENCES 169 APPENDIX A: ADDITIONAL TABLES 193 APPENDIX B: COLLECTION LOCALITIES AND NOTES 221 vi LIST OF TABLES Table Page 1 . Taxa Examined 51 2. Adult Grooming Frequencies 107 3. Number of Observed Grooming Combinations 122 4. Behavior of the Larvae and Pupae 126 5- Fore Leg Positioning in Adults 129 6. Product-moment Correlation Coefficients for Characters 194 7. Principal Component Analysis of Characters 196 8. Product-moment Correlation Coefficients for Chironominae 198 9. Principal Component Analysis of Chironominae Characters 200 10. Product-moment Correlation Coefficients for OTU's 202. 11. Distance Matrix of OTU's 205 12. Principal Component Analysis of OTU's 208 13- Five Dimensional MDS of OTU's 210 14. Distance Matrix of OTU's Based on Characters from Immatures 211 15* Distance Matrix of OTU's Based on Adult Fore Leg Positioning Characters 213 vii Table Page 16. Distance Matrix of OTU's Based on Characters of Adult Self-grooming 217 17- Three Dimensional MDS of OTU's Based on Characters of Adult Self-grooming 220 LIST OF FIGURES Figure 1. Frame for Collecting Tent 2. Section from a Test Tube Rearing Apparatus 3- A. Longitudinal Cross-section of a Typical Tanypodinae Burrow B. Tube and Feeding Net of Rheotanytarsus, redrawn from Walshe (1950) U. Tubes of Polypedilum digitifer 5. Leaf-miner Nets 6. Substrate Leg Position of Psectrotanypus dyari 7- Fore Leg Positioning in Chironomus attenuatus 8. Positioning of FFA in Chironomus attenuatus 9. Fore Leg Positioning in Dicrotendipes nervosus 10. Fore Leg Positioning in Stenochironomus hilaris 11. Plot of the first three PCA factors using all OTU's and characters 12. Plot of the first three MDS dimensions using all OTU's and characters Figure Page 13. Phenogram showing distances in the behavior of immatures 15^ 14. Phenogram showing distances in adult fore leg positioning behavior 155 15- Plot of the first two MDS dimensions using all OTU's andgrooming characters 157 x LIST OF ABBREVIATIONS Ab - grooming movement in which the abdomen is rubbed by one or both hind legs. ABLMAL - OTU abbreviation for Ablabesmyia mallochi ,j ACRSEN - OTU abbreviation for Acricotopus senex. An - grooming movement in which one or both antennae are rubbed by one or both fore legs. BL - a line bisecting the body longitudinally as seenjfrom above . CHIATT - OTU abbreviation for Chironomus attenuatus. CHIPLU - OTU abbreviation for Chironomus plumosus. CHIRIP - OTU abbreviation for Chironomus riparius. CHIRON - OTU abbreviation for Chironomus summary. CLADOT - OTU abbreviation for Cladotanytarsus summary. CLIPIN - OTU abbreviation for Clinotanypus pinguis. COELOT - OTU abbreviation for Coelotanypus summary. CONCHA - OTU abbreviation for Conchapelopia summary. CORYNO - OTU abbreviation for Corynoneura summary. CRIBIC - OTU abbreviation for Cricotopus bicinctus. CRICOT - OTU abbreviation for Cricotopus summary. CRISP3 - OTU abbreviation for Cricotopus sp. 3* CRISP4 - OTU abbreviation for Cricotopus sp. 4. CRISP5 - OTU abbreviation for Cricotopus sp. 5* xi CRITRI - OTU abbreviation for Cricotopus trifasciatus. CRYGAL - OTU abbreviation for Cryptocladopelma galeator. CRYPTO - OTU abbreviation for Cryptochironomus summary. DEMBRA - OTU abbreviation for Demei.jerea brachial is . DICFUM - OTU abbreviation for Dicrotendipes fumidus. DICMOD - OTU abbreviation for Dicrotendipes modestus. DICNER - OTU abbreviation for Dicrotendipes nervosus. DICROT - OTU abbreviation for Dicrotendipes summary. dvW - grooming movement in which the ventral wing surfaces are rubbed by the contralateral hind leg in conjunc­ tion with dWH. dWH - grooming movement in which the dorsal wing surfaces are rubbed by the ipsilateral hind legs. dWM - grooming movement in which the dorsal wing surfaces are rubbed by the ipsilateral mid legs. EINCHE - OTU abbreviation for Einfeldia chelonia. ENDNIG - OTU abbreviation for Endochironomus nigricans. FF - grooming movement reported by Heinz (19^9) in which
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