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Information to Users INFORMATION TO USERS While the most advanced technology has been used to photograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. For example: • Manuscript pages may have indistinct print. In such cases, the best available copy has been filmed. • Manuscripts may not always he complete. In such cases, a note will indicate that it is not possible to obtain missing pages. • Copyrighted material may have been removed from the manuscript. In such cases, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, and charts) are photographed by sectioning the original, beginning at the upper left-hand corner and continuing 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 as a 17”x 23” black and white photographic print. 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For an additional charge, 35mm slides of 6”x 9” black and white photographic prints are available for any photographs or illustrations that cannot he reproduced satisfactorily by xerography. 8703510 Antony, Lucille Marilyn May Kriger d’Amorim A PHYLOGENETIC ANALYSIS OF THE RHODACAROIDEA (ACARI: MESOSTIGMATA) ' The Ohio State University Ph.D. 1986 University Microfilms I nternetionel300 N. zeeb Road, Ann Arbor, Ml 48106 Copyright 1986 by Antony, Lucille Marilyn May Kriger d’Amorim All Rights Reserved A PHYLOGENETIC ANALYSIS OF THE RHODACAROIDEA (ACARI; MESOSTIGMATA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Phylosophy in the Graduate School of the Ohio State University By Lucille Marilyn May Kriger d*Amorim Antony, B.Sc., M.Sc. ****** The Ohio State University 1986 Dissertation Committee: Approved by N.F. Johnson S.W. Fisher D.L. Wrensch ^ idviser Department of Entomology Copyright by Lucille Marilyn May Kriger d'Amorim Antony 1986 To My Mother and to the Memory of My Father ii ACKNOWLEDGMENTS I want to thank my major Professor Dr. Donald E. Johnston, for his advice, and especially for the very stimulating discussions through the course of my studies at Ohio State. Thanks to him (and others), my memories of OSU will remain indelible. I am very grateful to the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico and to The Organization of American States for granting two-year scholarships during my graduate studies. I want to thank the acarologists and curators of collections who generously provided me with the material studied. Among them, special thanks to Mr. Keith Hyatt, Miss Anne Baker, Mr. Kieran Martyn of the British Museum of Natural History, Dr. Fausta Pegazzano of the Istituto Sperimentale per la Zoologia Agraria, and Dr. Jo Ann Tenorio of the Bernice Pauahi Bishop Museum, for their hospitality and for providing access to the collections. I thank the Instructional and Research Computer Center of The Ohio State University, for granting invaluable computer time. I extend my appreciation to Dr. Norman F. Johnson, for spending a great deal of his time helping me to get acquainted with the "mini-link" and other bugs of the computer world. iii My warmest thanks to the good friends who, directly or indirectly contributed to the accomplishment of this study. Finally, and most specially, I thank for all the love and constant support with which my very special family have gifted me all through the years. To them, my deepest love and appreciation. iv VITA October 15, 1950 . Born - Manaus, Amazonas, Brasil 1972-1975........... Trainee student. Laboratory of Pedobiology, Instituto Nacional de Pesquisas da Amazonia (INPA), Manaus, Brasil. 1973 ............... B.Sc. - Pharmacy, Universidade Federal do Amazonas, Manaus, College of Health Sciences. 1975-1977........... Research Assistant, Laboratory of Pedobiology, INPA, Manaus. 1978-1979........... Graduate studies, Department of Biology, Georgia Southern College, Statesboro. Sponsored by The Organization of American States. 1980 ............... M.Sc. - Entomology, The Ohio State University, Columbus, Ohio, sponsored by The Organization Of American States. 1981-1983........... Graduate Teaching Associate, Department of Entomology, The Ohio State University. 1983-1985........... Doctoral Student, Department of Entomology, The Ohio State University. Sponsored by the Conselho Nacional de Desenvolvimento Cientifico e Tecnologico, Brasil. 1985-1986........... Graduate Teaching Associate, Department of Entomology, The Ohio State University. Publications Antony, L.M.M.K.d'A. and Johnston, D.E., 1984. A Cladistic Analysis of the Genus Ologamasus Berlese (Acari; Mesostigmata). In; Griffiths, D.A. and Bowman, C.E.(eds.). Acarologv VI. v.l. Ellis Norwood Publ., Chichester, England, p.216-221. Presentations Antony, L. M, and Johnston, D, E. Phylogenetic studies of primitive Dermanyssina: The Rhodacaroidea (Acari). Entomological Society of America, Annual Meeting, Hollywood, Florida, 8-12 December, 1985. FIELDS OF STUDY Major Field; Entomology Studies in Taxonomy and General Biology of Oribatid Mites, Laboratory of Pedobiology, Instituto Nacional de Pesquisas da Amazonia, Manaus, Brasil. Dr. Herbert 0. R. Schubart. Studies in Reproductive Biology of Dermanvssus gallinae De Geer (Acari: Dermanyssidae). Department of Biology, Georgia Southern College, Statesboro, Georgia. Dr. James H. Oliver Jr., Callaway Professor. Studies in Taxonomy of Free-living Gamasina (Acari: Mesostigmata). Laboratory of Acarology, The Ohio State University, Columbus. Dr. Donald E. Johnston, Professor. Vi TABLE OF CONTENTS Page ACKNOWLEDGEMENTS............................................ i ü VITA....................................................... V LIST OF TABLES.............................................. ix LIST OF FIGURES ............................................ x INTRODUCTION................................................ 1 MATERIALS AND METHODS...................................... 26 CHARACTER ANALYSIS.......................................... 34 RESULTS AND DISCUSSION...................................... 57 Suprafamilial relationships............................. 57 Interfamilial relationships............................. 71 Intrafamilial relationships............................. 78 SYSTEMATIC ACCOUNTS ........................................ 130 Superfamily Rhodacaroidea Ouderaans ..................... 130 Family Euryparasitidae n. fam........................... 138 Family Rhodacaridae Ouderaans ........................... 144 Family Gamasellidae n. f a m ............................. 155 Family Ologaraasidae Ryke ....................... 162 Subfamily Ologamasinae Ryke............................. 169 Subfamily Epiphidinae Kethley........................... 171 Subfamily Gamasiphinae Lee ............................. 172 vii Taxa removed from the Rhodacaroidea Oudemans................173 Key to the families of Rhodacaroidea....................... 178 Superfamily Laelaptonyssoidea Womersley .......... .... 182 Family Panteniphidae n. f a m ............................... 188 SUMMARY........................................................ 205 LIST OF REFERENCES............................................ 207 APPENDICES.................................................... 213 A. MATERIAL EXAMINED (LISTED ALPHABETICALLY BY REPOSITORY) . ...................................................... 213 B. DATA MATRIX...........................................256 viii LIST OF TABLES TABLE Page 1. Family Rhodacaridae Oudemans sensu Ryke, 1962. 9 2. Superfamily Eugamasoidea Karg, 1965................ 13 3. Family Rhodacaridae Oudemans sensu Lee, 1970, 1973 .... 16 4. Family Rhodacaridae Oudemans sensu Bregetova, 1977 .... 22 5. Family Rhodacaridae Oudemans sensu Shcherbak, 1980 .... 24 6. Character Polarization ................................. 49 7. Proposed classification of the Rhodacaroidea and Laelaptonyssoidea. The arrangement of families is phyletically sequenced ................................. 126 IX LIST OF FIGURES FIGURE Page 1. Suprafamilial and interfamilial relationships of the Rhodacaroidea. Numbers on figure refer to stems. Characters are listed following sequence of stems. Character states are listed between parentheses........ 192 2. Intrafamilial relationships of the Euryparasitidae n. fam. Refer to Figure 1 legend for explanation............. 194 3. Intrafamilial relationships of the Rhodacaridae. Refer to Figure 1 legend for explanation.................. 196 4. Intrafamilial relationships of the Gamasellidae. Refer to Figure 1 legend for explanation.................. 198 5. Intrafamilial relationships of the Ologamasinae. Refer to Figure 1 legend for explanation.................. 201 6. Intrafamilial relationships of the Epiphidinae and Gamasiphinae. Refer to Figure 1 legend for explanation. 203 X INTRODUCTION The Rhodacaroidea is a large group of ecologically and morphologically diverse free-living mites of the suborder Mesostigmata. Together with the Dermanyssoidea, Eviphidoidea, Ascoidea and Veigaioidea they comprise the cohort Dermanyssina (Johnston, 1982). Females of this cohort are characterized by the presence of an accessory genital system - the spermatheca - which consists of paired ducts that open in the region of legs III and IV. The males possess a sperm-transfer device - the spermadactyl - located on the movable digit of the chelicera. Their genital
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