INFORMATION TO USERS This manuscript has been reproduced from the microfihn master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter frice, 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 afreet 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 NorthZeeb Road, Ann Arbor MI 48106-1346 USA 313/761-4700 800/521-0600 SYSTEMATICS OF THE NEW WORLD PTININAE: (COLEOPTERA: ANOBIIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Thomas Keith Philips, M.S. ***** The Ohio State University 1997 Dissertation Committee: Approved by Dr. Norman F. Johnson, Adviser Dr. Charles A. Triplehorn Dr. Woodbridge A. Foster Dr. Brian J. Armitage Department of Entomology UMI Number: 9731694 UMI Microform 9731694 Copyright 1997, by UMI Company. Ail 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 Thomas Keith Philips 1997 ABSTRACT 1. The biology of the subfamily Ptininae (Coleoptera: Anobiidae), with emphasis on the New World fauna, is reviewed and includes the following topics: adult lifestyles, biology, development and immatures, economic importance, evolution of myrmecophily, and systematics. A list of all the currently accepted generic names of the Ptininae with a sublist of the native New World genera is included. A key to all native and introduced genera encountered in the New World is also given. 2, A phylogenetic analysis of the New World Ptininae, based on the principles of cladistics is presented. One hundred and two characters (43 multistate) from 34 taxa (22 species of ptinines, eight additional anobiids, three bostrichids, and one endecatomid) were utilized. The single tree shows both the Ptininae and its sister, the remaining Anobiidae, as monophyletic. The phylogeny indicates that wing loss has occurred at least three times within the Ptininae, and myrmecophily in the New World ptinines has also probably evolved on at least three separate occasions. The evolution of feeding habits, myrmecophily, and wing loss are discussed. The classifications of the Ptininae and the remaining Anobiidae are also examined. 3. Cubaptinus Zayas is synonymized with Fabrasia Martinez and Viana. Known only from Cuba, Cubaptinus shares in common several characteristic morphological features of Fabrasia that necessitate it being placed as a junior synonym. 4. Lachnoniptus iindae New Genus, New Species from the Virgin Islands, is described. It appears most similar to Trigonogenius, and characters differentiating the two genera are given. The habitat and probable biology of this new species are discussed. 5. Coieoaethes tetralobus New Genus, New Species from Panama, is described. It is the third myrmecophilous genus of ptinine known from the New World. It is characterized by the presence of four lobes near the elytral apices, each with patches of brush-like setae or trichomes. The relationship of Coieoaethes tetralobus to other myrmecophilous ptinines and the diversity and distribution of this latter group are also discussed. 6. A revision of the genus Niptinus Fall is presented. Niptinus grandimacufatus is described as new , and Trigonogenius niveus trinotatus Pic is elevated to specific status. Trigonogenius niveus Gorham and T. arcuatus Gorham are transferred to Niptinus, raising the total number of species in this genus to six. The genus and all III previously described species are redescribed, and a key is provided. The relationship of Niptinus to other closely related genera and the known biology are discussed. 7. The New World genus Prosternoptinus is revised. Thirteen new species are described: P. aeneus, P. argosomos, P. assumentum, P. be/iesi, P. cafiieguanensis, P. castaneus, P. expandipronotus, P. glabrihumeralis, P. kryptoambon, P. obrienorum, P. ocuiigrandis, P. guadrimaculatus, and P. tricoratus, raising the total number of known species to 24. The genus and most previously known species are redescribed, and a key to all species is provided. A discussion of the biology and relationships of the genus is also included. 8. The first larval leaf mining and live-leaf feeding by adults is reported in the Anobiidae and the more inclusive superfamily Bostrichoidea. Larvae of Pitnus antiilanus Bellés bore in leaves of Tournefortia gnaphaiodes (Linnaeus) (Boraginaceae) and adults feed on leaves of the larval host plant. Larvae complete their development and metamorphosis within a single leaf. It is suggested that leaf mining is a recently derived trait in Pitnus. IV ACKNOWLEDGMENTS I wish to thank a large number of individuals, and the order of my list is not indicative of the importance of the great number of people who helped make this dissertation possible. To those 1 possibly and accidentally excluded, my apologies. I thank my advisor, Norman F. Johnson, and committee composed of Brian J. Armitage, Woodbridge A. Foster, and Charles A. Triplehorn for their guidance and support throughout my studies. All of them spent a great deal of time reviewing this thesis and other various documents. Norman was particularly supportive throughout my tenure here with equipment needs and financial assistance for professional travel. His expertise in systematics, vision for the future of our field, and high standards have hopefully been at least partially acquired by me. Michael Ivie, my former advisor, has continued to support and encourage me and is one of the few with whom I can discuss anobiine evolution. He passed on his broad interest of Coleoptera to me, for which 1 am grateful. My gratitude is owed to John Wenzel for his musings and teachings on phylogenetic theory, techniques, and help with cladistic programs and data analysis. My appreciation is extended to Barry Valentine for his interesting discussions and wealth of knowledge on collections, museums, and other aspects of systematics, especially regarding the Coleoptera. He also reviewed several chapters of this dissertation. Xavier Bellés was of great help in suggestions on particular groups and in general encouragement in my studies on spider beetles. I thank him for his generosity in allowing me to work on "his" group. Additionally his reprints, and a set from Richard W hite, saved my search time and were of great use during these studies. My gratitude also extends to the following curators and individuals, who gave, loaned, or made available material for these and other studies: Rolf Aalbu, Robert Anderson, Fred Andrews, Miguel Archangelsky, Cheryl Barr, Richard Baumann, Jane Beard, Nicole Berti, Yves Bousquet, Roberta Brett, Edward Brodie, Jorge Carvajal, John Chemsak, Shawn Clark, William Clark, José Clavijo, Robert L. Davidson, Greg Forbs, David Furth, Terry Galloway, François Génier, Bruce Gill, Wilfred Hanson, Lee Herman, Richard Hoebeke, Gloria House, Michael Ivie, Norman Johnson, Luis Joly, David Kavanaugh, Peter Kovarik, Will Lanier, Jean-Michel Maes, Vicky Moseley, Alfred Newton, Charles O'Brien, Mark O'Brien, Norman Penny, Philip Perkins, Darren Pollock, Edward Riley, Terence Schiefer, Catherine Seibert, Derek Sikes, Paul Skelley, Cecil Smith, Angel Solis, Michael Thomas, William Warner, and Keti Zanol. I am grateful to George Keeney who supplied colonies of two species and media for rearing. Thanks to Derrick Sikes and Richard Miller, who made the original observations of the association of Pitnus antiilanus with Tournefortia gnaphaiodes, and to Mike and LaDonna Ivie, who made the study on the biology of this species possible. Michael also collected most of the specimens of Lachnoniptus iindae. vi I thank K. Roach, D. Hanmer, and D. Hennessey (USDA/ APHIS/ PRO) for providing a permit to import spider beetles and host plant material from the Virgin Islands and Kathleen Akin (USDA/ APHIS/ VS) for one to import more spider beetles from Venezuela. Thanks to the following individuals for their help to my coauthors (Chapter 8) in the Virgin Islands: K, Woody for help with field work in the Virgin Islands; D. Nellis for help locating St. Thomas populations; Z.-M. Hillis of the National Park Service and E. Towle and S. R. Tate of Island Resources Foundation for logistic support during field work on St. Croix and St. Thomas. I also thank John Furlow for his assistance with identification of the host plant and for use of the herbarium library. Thanks to the staff of Lewis and Clark Caverns. Montana, for permission to collect spider beetles in the caves. I thank the librarians at the OSU Biological-Pharmacy library: Susan Birky, Bruce Leach, and Susan Ward for their patience in helping me find the many volumes of obscure references needed to do taxonomic work. My gratitude to Ken Chamberlain (OSU Communications and Technology) for allowing me to use his great Nikon macro camera equipment, which made it relatively easy to take photos of my tiny beetles. Miguel Archangelsky also was a great deal of help for his photographic skills on numerous occasions.