Mites (Acari) Associated with the Ants (Formicidae) of Ohio and the Harvester Ant
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Mites (Acari) Associated with the Ants (Formicidae) of Ohio and the Harvester Ant, Messor pergandei, of Arizona THESIS Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University By Kaitlin A. Uppstrom Graduate Program in Evolution, Ecology and Organismal Biology The Ohio State University 2010 Master's Examination Committee: Associate Professor Hans Klompen, Advisor Associate Professor Susan Jones Professor Steven Rissing Copyright by Kaitlin A. Uppstrom 2010 Abstract Ants (Formicidae) have long been an insect group of great interest to the scientific world, whether for their ecological roles, feeding strategies, or social behaviors. They form complex colonies, harboring resources that can potentially be exploited by myrmecophiles (organisms living in association with ants). Myrmecophily has been studied in detail for Coleoptera, but mites (Acari), the most frequent of ant guests, remain largely unstudied. Previous work has focused primarily on descriptions and has provided little ecological information. The first study is an effort to provide a more robust list of the often overlooked inhabitants of ant nests focusing on Ohio, a state that has yet to be mentioned in any myrmecophilous mite studies. A general survey of common Ohio ants was conducted from April 2008-March 2010. Phoretic mites were individually removed from ants and debris in 273 colonies. Mite collections totaled 198 species: 151 species phoretic and at least 47 mite species in non-phoretic relationships within the ant nests. Phoretic mites consisted of representatives of the cohort Astigmata (Histiostomatidae and Acaridae), the cohort Heterostigmatina (Scutacaridae, Pygmephoridae, and Microdispidae), and the suborder Mesostigmata (Laelapidae, Antennophoridae, and Uropodina). Many mite species were host specific and attachment site specific. An unusually large number of mite species was found to be associated with the ant genus Lasius, possibly the result of social parasitism. Post hoc statistical analyses show significantly greater mite diversity in ii colonies when 1) in the ant subfamily Formicinae, 2) the colony is in the woods, 3) the nest substrate is wood, 4) the colony is populous, 5) the ants are large, and 6) the ant species establishes its nest parasitically. A second study focused on the seed harvester ant Messor pergandei and its acarine associates. At least seven mite species are phoretically associated with M. pergandei: Armacarus sp., Lemanniella sp., Petalomium sp., Forcellinia sp., Histiostoma sp., Unguidispus sp., and Cosmoglyphus sp. Most of these species show preference for specific phoretic attachment sites and most preferentially board female alates rather than male alates. Five mite species were found in low numbers inhabiting the chaff piles: Tydeidae sp., Procaeculus sp., Anystidae sp., Bakerdania sp., and Tetranychidae sp. The phoretic Petalomium sp. was observed consuming fungus on a dead dealate, but the roles of the other species are still unclear. iii Dedicated to Jana, Kevin, Ariel and Jennifer iv Acknowledgments I would like to thank my advisor, Dr. Hans Klompen, for his inspired ideas, constant support, enthusiasm, and patience, for the freedom to mold my own unique project, and for the career and life changing moment which began the moment I stepped into the Acarology lab as an undergraduate student. I would like to thank my committee member, Dr. Susan Jones, for her thoughtful guidance, assistance in reducing a growing project into something managable, and for careful editing which allowed me to write what I really intended. I would like to thank my committee member, Dr. Steven Rissing for his great assistance in providing information and guidance allowing me to arrive safely in Arizona and collect Messor pergandei, for corrections and input on my thesis and for asking the hard questions. I would like to thank Dr. Joseph Raczkowski for advice and insight concerning Messor pergandei and later the Lasius genus and its habits. Many thanks to Dr. Dave Walter, Dr. Ron Ochoa, Dr. Cal Welbourn, and Dr. Barry OConnor, of the Acarology Summer Program, who have provided ecological and taxonomic guidance throughout my project. I would like to thank the people who assisted with the Messor pergandei project: Dr. Bob Johnson for information, ideas, and the generous use of his shovel for digging v ant craters, Clint Penick for teaching me how to search and rear dealates, Shellie and Robb Hjellum for welcoming me and my ant specimens into their home. I would like to thank Bob Bryant and Brenda Bryant for permitting me to collect numerous ant colonies on their property giving me the opportunity to find Antennophorus and Trichocylliba mites. I would like to thank the members of the Acarology Lab past and present, Monica Farfan, Luke Kapper, and Sam Bolton for friendship, suggestions, and advice throughout my project. I am indebted to The Ant Course which allowed me to meet so many interesting people and acquire the knowledge in ant taxonomy, collecting, and ecology that has been invaluable to my project. I thank Gary and Holly Coovert for their hospitality, assistance with identification of difficult species groups, and for the invaluable reference, The Ants of Ohio. Special thanks to my family and close friends: My parents Jana Morse, and Kevin Uppstrom, for supporting me in all things, school and otherwise, nurturing my love of nature, and for having so many rocks in the backyard that house ants and their mites, my sisters Jennifer Alford and Ariel Uppstrom for support, advice, and for listening to me talk incessantly about bugs, to Gabe Campbell for endless patience, support, the technical skills I lack, and for his mutual interest in ants, to Joshua Bryant, for his company and bravery (particularly for Formica exsectoides) on collecting trips, for his advice on my project, and his patient ear. vi Vita January, 24, 1985 ...........................................Born, Hamilton, Ohio, USA 2007................................................................B.S. Zoology, The Ohio State University 2007-2009 ......................................................Graduate Teaching Associate, The Ohio State University 2009-2010 ......................................................Metro School Fellow, The Ohio State University 1980 to present Publications Uppstrom, K. and H. Klompen. (2005). A new species of Julolaelaps (Acari: Iphiopsididae) from African millipedes. International Journal of Acarology, 31: 143-147. Corriveau, M., K. Uppstrom, and H. Klompen. (2010). Effect of eight storage modes on DNA preservation. In: Sabelis, M. W., Bruin, J. (Eds), Trends in Acarology, Proceedings of the 12th International Congress. Springer Science, Dordrecht, pp. 553-556. Fields of Study Major Field: Evolution, Ecology and Organismal Biology vii Table of Contents Abstract ............................................................................................................................... ii Acknowledgments............................................................................................................... v Vita .................................................................................................................................... vii List of Tables ................................................................................................................... xiii List of Figures .................................................................................................................. xix Chapter 1: Ants, Mites, and Current Knowledge of Interspecific Associations ................ 1 Family Formicidae: The Ants ......................................................................................... 1 Ants as Social Insects .................................................................................................. 1 The Ant Nest ................................................................................................................ 2 Communication in Ants ............................................................................................... 3 Interactions among Ants .............................................................................................. 4 Myrmecophiles ................................................................................................................ 6 Mites and Myrmecophily............................................................................................. 7 Phoresy in Mites .......................................................................................................... 8 Mites Documented from Ant Nests ........................................................................... 10 viii Scope of these Studies ................................................................................................... 22 Literature Cited ............................................................................................................. 24 Chapter 2: Mites Associated with Ants in Ohio ............................................................... 38 Introduction: Goals, Questions, and Scope of this Project ............................................ 38 Materials and Methods .................................................................................................. 40 Collection Dates and Localities ................................................................................. 40 Colony Sampling Methods .......................................................................................