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. 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. 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 BIOLOGY OF THE HETEROZERCONIDAE DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Beverly Swaim Gerdeman, M.S. The Ohio State University 2002 Dissertation Committee: Approved by Assistant Professor Hans Klompen, Adviser Associate Professor Dana Wrensch Professor David Horn // /^Adviser Professor Emeritus Rodger Mitchell itomofogy Grad- Reproduced with permission of the copyright owner. Further reproduction prohibited without permission UMI Number 3039471 ___ © UMI UMI Microform 3039471 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. ABSTRACT Narceoheterozercon ohioensis n. gen., n. sp. (Acari: Heterozerconidae) the first North American representative of Heterozerconidae, is described. The description includes the first immatures for the infraorder Heterozerconina. A new location for the secondary genital opening in the Acari is discovered in the inner, anterior comer of the suckers in females. The phenology of a tropical heterozerconid provides a comparison for that in the temperate climate. Both field studies and laboratory observations establish the biology and ecology of Narceoheterozercon ohioensis. The phenology between the mites and their millipede host, Narceus annularis (Diplopoda: Spirobolida) shows coinciding periods of oviposition and mating. The three immature mite instars develop through the summer beginning in June and continuing through August. Teneral adults appear in late August and early September and males may briefly outnumber females. The mating millipedes may provide a cross-species mating stimulus for their heterozerconid commensals. Evidence suggests post-mating dispersal occurs in the millipedes. Mites appear to overwinter on their hosts. A hypothesis for the origin of the spermatodactyi on the fixed digit of the males is proposed. The spermatodactyi appears to have evolved from structures associated with the fixed digit, including the pilus dentilis. This indicates the spermatodactyi in the Heterozerconina arose independently from that in the Dermanyssina. ii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. A world survey of spermatodactyls reveals an enormous diversity. The diversity may be due to lengthening and fusing of structures associated with the fixed digit. Differences in the appearance of the spermatodactyls allows them to be grouped into 4 major zoogeographic regions: African, Oriental, Neotropical and Nearctic. Characteristics in the spermatodactyls of each region are discussed. The diversity in the spermatodactyls suggests a variety of reproductive methods occur in the Heterozerconidae. Podospermy appears present in all four zoogeographic regions. Evidence from the Oriental region suggests a transition stage between tocospermy and podospermy may occur in the Heterozerconidae. Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Dedicated to the kingdoms of the hollow iv Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGMENTS I wish to thank my adviser, Hans Klompen, for sharing his enthusiasm in my project and for his patience, intellectual support and willingness to listen to my ideas. I am grateful to Rodger Mitchell for introducing me to the classical aspects of scientific writing and illustration and for never running out of time. I also thank Dana Wrensch for understanding and supporting the addition of a creative artistic approach to the logical processes of scientific problems. My sincere appreciation to my husband Robert, who never grew weary of the words “millipede” and “heterozerconid", uttered at all times of the day and night and for allowing me to pursue my goals. To my four daughters: Ana, Erikah, Mauria and Diedre for their unwavering support and surviving on promises. The tropical research was made possible through the support of the Fulbright Association, The Institute of International Education and the Philippine American Education Foundation. Thanks to Ms. Tessie Camero-Hawes, Leonila Raros and Mercedes Delfinado-Baker for their support and enthusiasm during my stay in the Philippines. My sincere gratitude to Phin Garcia for sharing his tropical collecting expertise and for having the courage to guide me through the islands. v Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. VITA July 14, 1952 ................................. Born - Hot Springs, Arkansas 1996 ............................................... M.S. Entomology, The Ohio State University 1977 ............................................... B.S. Education, Henderson State University 1974 ............................................... B.S. Wildlife Management, Arkansas Tech College 1993-present ................................ Graduate Teaching and Research Associate, The Ohio State University PUBLICATIONS Research Publication 1. Gerdeman, B.S., Klompen, J.S.H. and J.A. Yoder. 1999. Description of the larva of Gromphadorholaelaps schaeferi (Acari: Laelapidae) a parasite of the giant Madagascar hissing-cockroach. Int. J. Acarol., 24: 301-305. 2. Gerdeman, B.S., Klompen, J.H.S., and L. K. Tanigoshi. 2000. Insights into the biology of a mite/millipede association. In: Wytwer J. and S. Golovatch (eds.), Progress in Studies on Myriapoda and Onychophora. Fragm. faun., Warszawa 43:223-227. FIELDS OF STUDY Major Field: Entomology Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS PACE Abstract ............................................................................................................................ ii Dedication .................................................................................................................. iv Acknowledgments ...................................................................................................... v Vita ............................................................................................................................... vi Table of Contents ................................................................................................... vii List of Tables ........................................................................................................... x List of Figures ......................................................................................................... xi Chapters: 1. Introduction ...................................................................................................... 1 2. A new North American heterozerconid, Narceoheterozercon ohioensis, n. gen., n. sp., with first description of immatures of Heterozerconidae (Acari: Mesostigmata) ................................................................................................. 6 2.1 Introduction .............................................................................. 6 2.2 Materials and Methods ............................................................ 8 2.2.1 Study Site ................................................................................ 8 2.2.2 Collecting and Rearing Techniques .......................................... 8 2.2.3 Material Examined.................................................................... 8 vii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2.2.4 Specimen Preparation and Measurements................................. 10 2.2.5 Specimen Depositories ..............................................................10