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 corner and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. 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. UMI A Bell & Howell Information Company 300 North Zeeb Road, Ann Arbor MI 48106-1346 USA 313/761-4700 800/521-0600 STUDIES ON THE BIOLOGY, ECOLOGY AND SYSTEMATICS OF THE PREIMAGINAL STAGES OF NEW WORLD HYDROPHILOIDEA, WITH CONSIDERATIONS ON THEIR PHYLOGENY (COLEOPTERA: STAPHYLINIFORMIA) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Miguel Archangelsky The Ohio State University 1996 Dissertation Committee: Approved by J. W. Wenzel Co-Advisl D. L. Denlinger C. Welty C. A. Triplehom Co-Adviser Department of Entomology UMI Number: 9630845 Copyright 1995 by Archangelsky, Miguel All rights reserved. UMI Microform 9630845 Copyright 1996, by UMI Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 Copyright by Miguel Archangelsky 1995 ABSTRACT This is a taxonomic and bionomical study of the immature stages of the New World hydrophiloids. Most of the information presented here comes from specimens collected in the field and reared under laboratory conditions; this allows for unequivocal associations between immature stages and adults. Description of collecting and rearing techniques for beetles coming from different habitats is discussed; also a general account of the biology of hydrophiloids is summarized. A generic key to larvae of all known New World hydrophiloids is provided, as well as diagnostic descriptions for each genus; these descriptions include the egg cases, larvae, and pupae. Further information on number of species, distribution, species with known immature stages, and additional biological characteristics is provided for each genus. Detailed illustrations for each genus supplements the key and the diagnostic descriptions. The immature stages of five genera are described here for the first time: Notionotus, Oocyclus, Pelosoma, Cryptopleurum, and Pemelus. A phylogenetic analysis using characters from larvae, pupae, egg cases, and adults was conducted; the outgroups included Silphidae, Hydraenidae, and Histeridae. Characters of the immature stages were obtained from the material studied, those from the adults were obtained from M. Hansen’s generic revision for the superfamily. The results support the position of Hydraenidae within the Staphylinoidea, and not as part of Hydrophiloidea; Histeroidea is proposed as the sistergroup of Hydrophiloidea. At a family level two clades are distinguished, the first one formed by Georissidae, Epimetopidae, and Helophoridae; the second one formed by Hydrophilidae, Spercheidae, and Hydrochidae. Larval characters are most informative at the base of the tree, especially the characters associated to the development of the spiracular atrium in the larvae; adult characters were more informative at the apex of the tree. To my parents and Don Johnston “In the final years of the twelfth century, from twilight of dawn to twilight of dusk, a leopard looked upon some wooden planks, some vertical iron bars, men and women who were always different, a thick wall and, perhaps, a stone trough filled with dry leaves. The leopard did not know, could not know, that what he craved was love and cruelty and the hot pleasure of rending and the odor of a deer in the wind; and yet something within the animal choked him and something rebelled, and God spoke to him in a dream: You live and will die in this prison, so that a man I know may look at you a certain number of times and not forget you and put your figure and your symbol in a poem which has its precise place in the scheme of the universe. You suffer captivity, but you will have furnished a word to the poem. In the dream, God enlightened the rough beast, so that the leopard understood God’s reasons and accepted his destiny; and yet, when he awoke, he felt merely an obscure resignation, a gallant ignorance, for the machinery of the world is overly complex for the simplicity of a wild beast. Years later, Dante lay dying in Ravenna, as little justified and as much alone as any other man. In a dream, God revealed to him the secret purpose of his life and labor; in wonderment, Dante knew at last who he was and what he was and he blessed his bitter days. Tradition holds that on awakening he felt he had received and then lost something infinite, something he could not recuperate, or even glimpse, for the machinery of the world is overly complex for the simplicity of men.” Inferno 1,32 JORGE LUIS BORGES Translated by Anthony Kerrigan ACKNOWLEDGMENTS I would like to express my most sincere appreciation to my advisors Dr. John W. Wenzel, Dr. Charles A. Triplehom, and Dr. Don E. Johnston. Their support, guidance, criticism, and insight have been invaluable during my research. I would also like to thank the other members of the advisory committee, Drs. D. L. Denlinger C. Welty, Dana Wrensch and Dr. Dan Crawford. Many people have devoted their time and assistance during these past years, especially during my fieldwork, without them a lot of the information presented in this dissertation would have been difficult to gather; they are P. W. Kovarik, T. K. Philips, K. Stephan, R. Androw, and George Keeney. I am also indebted to A. F. Newton, D. L. Gustafson, C. Costa, L. Fem&ndez, M. Toledo, and P. Skelley for providing me with preserved specimens for this study. I have to thank several institutions: the INTA in Argentina, the people form UCV (Maracay), and Rancho Grande Biological Station in Venezuela, and the Archbold Biological Station in Florida. All of them provided me with support and permits in order to collect some of the specimens used in this study. The Ohio State University through the Graduate Alumni Awards, and the Tinker Foundation made possible my collecting trip to Venezuela. A special thanks goes for George Keeney for his assistance in beetle rearing, the same goes for M. E. Durand. The illustrations that accompany this dissertation have been possible because T. L. Temple spend time teaching me different drawing techniques, and tactfully criticizing my artwork. My former colleagues from Argentina, especially Dr. A. O. Bachmann from the Museo Argentino de Ciencias Naturales Bernardino Rivadavia, and Dr. Ronderos from the Museo de La Plata, are responsible for my interest in entomology and aquatic insects. Don Johnston taught me a lot about how to conduct research, but the most valuable impression he left on me was his devotion to teaching; his continuous search for a level of excellence made him a great inspiration, and one of the better teachers I have ever known. Finally My most sincere thanks go to my parents, my sister, and the rest of my family for their support, patience and understanding. VITA November 11, 1961 Bom - Buenos Aires, Argentina 1982 - 1986 B.A. Biology C.A.E.C.E. College Buenos Aires, Argentina 1987 - 1989 Researcher, Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina 1990 - present Graduate Teaching and Research Associate, The Ohio State University PUBLICATIONS Refereed Publications Archangelsky, M. 1985 (1987). Nota distribucional sobre Tropistemus (Tropisternus) scutellaris Castelnau 1840 (Coleoptera, Hydrophilidae). Revta. Soc. ent. Arg. 44(2): 168. Archangelsky, M. 1989. A new species of the genus Phaenonotum Sharp, 1882 (Coleoptera, Hydrophilidae, Sphaeridiinae) from Venezuela. Aquat. Insects. 11(4): 217-220. Archangelsky, M. 1990 (1991). El gdnero Phaenonotum Sharp en la Argentina. I. Redescripcidn de Phaenonotum regimbarti Bruch, 1915 y Ph. argentinense Bruch, 1915 (Coleoptera, Hydrophilidae, Sphaeridiinae). Revta. Soc. ent. Arg. 49(1-4): 157-164. Archangelsky, M. 1992. El g6nero Phaenonotum Sharp en la Argentina II (Coleoptera, Hydrophilidae, Sphaeridiinae). Ubicacidn de Hydroglobus puncticollis Bruch, 1915 dento del gdnero Phaenonotum. Revta. Soc. ent. Arg. 51(1-4): 47-51. vii Archangelsky, M. and M. E. Durand. 1992a. Description of the preimaginal stages of Dibolocelus ovatus (Gemminger and Harold, 1868) (Coleoptera, Hydrophilidae: Hydrophilinae). Aquat. Insects. 14(2): 107-116. Archangelsky, M. andM. E. Durand. 1992b. Description of the preimaginal stages of Derallus angustus Sharp, 1882 (Coleoptera: Hydrophilidae, Hydrophilinae). Aquat. Insects. 14(3): 169-178. Archangelsky, M. and M. E. Durand. 1992c. Description of the immature stages and biology of Phaenonotum exstriatum (Say, 1835) (Hydrophilidae: Sphaeridiinae). Coleopts. Bull. 46(3): 209-215. Archangelsky, M. 1994a. Description of the preimaginal stages of Dactylostemum cacti (Coleoptera: Hydrophilidae, Sphaeridiinae). Ent. scand. 25: 121-128. Archangelsky, M. 1994b. Description of the immature stages of three Nearctic species of the genus Berosus Leach (Coleoptera: Hydrophilidae). Int.