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Phylogeny of Lasius (Acanthomyops

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Phylogeny of Lasius (Acanthomyops PHYLOGENY AND FOUNDING STAGE OF LASIUS (ACANTHOMYOPS) (HYMENOPTERA: FORMICIDAE) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By Joseph Martin Raczkowski, M. S. **** The Ohio State University 2008 Dissertation Committee: Approved by Professor John W. Wenzel, Co-Advisor Professor Steve Rissing, Co-Advisor ______________________________ Professor Norman F. Johnson ______________________________ Professor Joan Herbers Co-Advisors Graduate Program in Evolution, Ecology, and Organismal Biology ABSTRACT The evolution of social parasitism in ants remains a controversial topic despite nearly 200 years of study and discussion. A lack of the details of socially parasitic behavior and phylogenies of socially parasitic groups are the greatest hindrances to understanding the evolution of this behavior. In this dissertation I address these problems for a group of North American ants, Lasius (Acanthomyops). I review social parasitism in ants, summarizing what we know and identifying the problems with the available data including distributional data. I present a study of socially parasitic behavior for my group of interest, as well as a phylogeny based on morphology and molecules. Finally, I discuss the implications of my findings on the evolution of social parasitism within the context of Emery's Rule. ii Dedicated to Michell, Ava, and Dean Raczkowski. iii ACKNOWLEDGMENTS This dissertation would not have been possible without the help of many people. I would like to thank my committee members John Wenzel, Steve Rissing, Norm Johnson, and Joan Herbers for improving my work. I thank all Wenzel Lab people, past and present, including: Kurt Pickett, Mary Daniels, Dave Rosenthal, Sibyl Bucheli, Hojun Song, Todd Gilligan, Sarah Mominee, Gloria Luque, Giancarlo Lopez-Martinez, Erin Morris, Ryan Caesar, and Glené Mynhardt. I am grateful to the following people for sharing their knowledge: Chris Johnson, Larry Phelan, Meg Daly Lab, John Freudenstein Lab, and Hans Klompen. I thank all people of the Museum of Biological Diversity for providing a pleasant work environment. I thank the following people from outside Ohio State University: Edward O. Wilson, Stefan Cover, Gary Alpert, Jim Carpenter, Roberto Keller, Ted Schultz, John LaPolla, Mike Sharkey Lab, Earl Werner, Justin Congdon, and Joshua Capps. I thank Michell, Ava, Dean, Russell, Carl, Paul, Mary, Bill, and Jody Raczkowski for their support. I thank Ralph Martin, Gail Martin, and Brandon Martin for their support. Lastly I thank Robert Stieferman, Jeff Latham, Jonathon Tanner, Steve and Tiffany Graham, and John and Laura Riffle for their friendship. iv TABLE OF CONTENTS Abstract .............................................................................................................................. ii Dedication ......................................................................................................................... iii Acknowledgments ............................................................................................................ iv Vita ..................................................................................................................................... v List of Figures ................................................................................................................. viii List of Tables .................................................................................................................... xi Chapters 1. Introduction..................................................................................................................... 1 2. Review of social parasitism in Formicidae (Insecta: Hymenoptera) with an emphasis on the genus Lasius ....................................................................................... 5 Introduction ............................................................................................................ 5 Associations of Ants .............................................................................................. 6 Host characteristics in the different forms of social parasitism ............................. 8 Temporary social parasitism in Lasius ants ........................................................... 9 Dependence of measures of diversity upon taxonomic effort ............................. 22 Predictors of species richness among ants ........................................................... 24 Limitations of the data ......................................................................................... 26 Taxonomic effort, lumping and splitting ............................................................. 27 v Importance of revisionary systematics ................................................................. 29 Conclusion ........................................................................................................... 31 3. Founding Stage of Lasius (Acanthomyops) and the incidence of social parasitism ..................................................................................................................... 52 Introduction .......................................................................................................... 52 Methods ................................................................................................................ 55 Results .................................................................................................................. 58 Discussion ............................................................................................................ 63 Conclusions .......................................................................................................... 71 4. Phylogenetic Analysis of Lasius (Acanthomyops) ....................................................... 90 Introduction ......................................................................................................... 90 Taxonomic history .............................................................................................. 92 Materials and methods ....................................................................................... 95 Results and discussions ....................................................................................... 98 Conclusions ........................................................................................................ 111 5. Appendix A : Cytochrome Oxidase Subunit 1 Sequences Aligned by MUSCLE .... 123 6. Appendix B: wingless sequence data aligned by MUSCLE ..................................... 132 7. Appendix C : Taxonomic Descriptions....................................................................... 136 8. Bibliography .............................................................................................................. 161 vi LIST OF FIGURES Figure Page 2.1 Number of ant taxa (species and subspecific ranks) described from different nations, according to Antbase. Nations specifically indicated are, by region, Canada (41), the United States (1013), Nicaragua (13), Costa Rica (248), Colombia (233), Brazil (1266), Switzerland (91), Algeria (206), Egypt (57), Congo (95), Zaire (455),South Africa (515), Russia (237), China (169), India (505), Indonesia (903), New Guinea (657), and Papua New Guinea (41) ........... 42 2.2 Relationship between number of taxa described from a region and (a) the region’s area (square kilometers; r2= 0.201), (b) the mean temperature of the region (degrees Celsius; r2= 0.001), (c) the region’s average annual precipitation (millimeters; r2 = 0.059), and (d) the number of taxonomists describing taxa in the region (r2 = 0.592). The numbers in the figure panels refer to the following countries: (1) Brazil, (2) United States, (3) Indonesia, (4) New Guinea, (5) India, (6) Costa Rica, (7) Colombia, (8) Switzerland, (9) China, (10) Canada, (11) Russia, (12) South Africa, (13) Algeria, (14) Egypt, (15) Papua New Guinea, and (16) Zaire ....................................................................................................... 43 2.3 The number of taxa described from Colombia, by author ................................... 48 2.4 Classifications of taxa Forel described from Colombia ....................................... 49 2.5 Number of taxa described from Congo, by author .............................................. 50 2.6 Classifications of taxa Santschi described from Congo ....................................... 51 3.1 Reorganization of Wing’s (1968) hypotheses for the founding stage of Lasius (Acanthomyops) according to overall strategy (non-parasitic v parasitic), number of queens founding a colony, and details of the non-parasitic and parasitic habit ....................................................................................................... 84 3.2 Grouping experiment arenas. Four L. interjectus gynes were placed in each arena. Each arena contained four perches (rocks), four shelters (5 x 5 cm piece of glass with one end propped up by a cap from a 1.5 mL plastic vial), and four sources of food (piece of tin foil with Bhatkar’s diet) ...................................................... 85 vii 3.3 Survivorship of individual queens that started either singly (P < 0.03, Kaplan-Meier, Tarone-Ware), in groups of two (P < 0.04, Kaplan-Meier, Tarone-Ware) and in groups of four............................ 86 3.4 Number of flagging events recorded within the first hour of observations in
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