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Phylogeny and Evolution of Myrmecophily in Beetles, Based on Morphological Evidence (Coleoptera: Ptinidae, Scarabaeidae)

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Phylogeny and Evolution of Myrmecophily in Beetles, Based on Morphological Evidence (Coleoptera: Ptinidae, Scarabaeidae) Phylogeny and Evolution Of Myrmecophily In Beetles, Based On Morphological Evidence (Coleoptera: Ptinidae, Scarabaeidae) DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Glené Mynhardt Graduate Program in Evolution, Ecology and Organismal Biology The Ohio State University 2012 Dissertation Committee: Johannes Klompen, Advisor Marymegan Daly Norman Johnson T. Keith Philips Copyright by Glené Mynhardt 2012 Abstract Ant-associated behavior has evolved rampantly among various groups of Arthropoda, and has arisen in at least 34 families of beetles. Due to the amazing morphological modifications and different kinds of interactions that occur within myrmecophilous (ant-associated) beetles, authors have predicted that myrmecophily has evolved in a step-wise fashion from casual, facultative associations to closely integrated, obligate interactions. In this dissertation, myrmecophily within the Coleoptera is reviewed, and known behaviors, ant-beetle interactions, and associated morphological adaptations are discussed. In order to better understand how myrmecophily has evolved, two groups of beetles are studied in a phylogenetic context. A cladistic analysis of 40 species of the myrmecophilous scarab genus, Cremastocheilu s Knoch is presented. Characters related to a myrmecophilous habit are largely informative, especially those characters related to the glandular trichomes (clusters of setae typically associated with exocrine glands). Two of the five previously recognized subgenera, C. (Myrmecotonus ) and C. (Anatrinodia ) are synonymized with the subgenus C. (Cremastocheilus ). Even though behavioral information is only known for a few species, the resulting phylogeny indicates that monophyletic subgenera are largely associated with the same ant hosts, although specific interactions with ant hosts can vary even in closely-related taxa. In addition, a separate cladistics analysis of the spider beetles based on morphology is ii presented. The monophyly of previously proposed suprageneric groups are investigated, and eight tribes of spider beetles (four unnamed basal tribes, plus more derived Gibbiini, Ptinini, Sphaericini) are recognized to capture three unique monophyletic groups of spider beetles. Of eight myrmecophilous spider beetle genera, only one genus ( Gnostus ) can be placed, all other myrmecophilous representatives remain unplaced in phylogeny. Based on this analysis, myrmecophily has evolved independently in four lineages, with nearly all genera appearing basal in spider beetle phylogeny. Based on these findings, obligate myrmecophily has evolved in four different groups. Among the Old World taxa (South African, Australian), and based on morphology, antennae may have evolved in a step-wise fashion, from less specialized (normal, 11-segmented antennae) to more highly specialized (reduced, with fusion of segments); however, pronotal trichomes are present in all related taxa. Among New World taxa, the presence of unique myrmecophilous adaptations indicates rapid evolution of obligate ant-associated behavior and morphology, rather than step-wise evolution from casual or facultative to obligate associations. Finally, two new genera of Dominican amber spider beetles are described. The new genus Electrognostus may indicate a transition from a normal Ptinus -like spider beetle to a myrmecophilous type like Gnostus , but based on the phylogenetic analysis it can currently be placed within the Ptinini tribe. iii This document is dedicated to the many people who gave me strength to believe in myself. iv Acknowledgments I have a lot of people to thank for guiding and supporting me through this interesting venture called graduate school. I have to thank my family first, for giving me the opportunity to study biology, fall in love with insects, and pursue the study of some of the coolest beetles on Earth. Thanks, Anna Mynhardt, my mom, for being there on days that were less than great, and for supporting me and pushing me on days when doing all this felt right. To my dad, Hendrik Mynhardt, for instilling the drive and passion in me to always strive for more. And to my brother, Gavin Mynhardt, for being strong for me and for you. Within the first year of graduate school I met Jason Kolenda, a man who has stood by me through all the ups and downs and has carried me through some of the hardest days of my life. Thank you for making me laugh. Thank you to Jane, Dave, and Aaron Kolenda for making me see what life is really about – having the right people by your side. Thanks to “Grandpa” Herman Lubertazza, Laurie Parham, the Meserini family, Carney, Cheryl, Pat, Nathan, and Danny Lubertazza. Thanks to all my friends, Ashley Kulhanek, Meaghan Ventura, Joshua Bryant, Kaitlin Uppstrom, Monica Farfan, Erin Morris, Chelsea Korfel, the Cary family, The Meehl family, the Rinas family, the Spilker family, and Heather Stephens – you were all there for me at different times during this journey. Thank you Kathy Horava, Judith Cusin, Joelle Fenger, Reni Ayachitula, Lynn Healy and Joanne Strunk for your constant v encouragement and for making me feel like I could conquer the world. I would have been much less of a person without you. Thanks to my many friends at the Museum of Biological Diversity – Abby Reft, Ryan Caesar, Joe Raczkowski, Charuwat Taekul, Brandon Sinn, Ryan Folk, Paul Larson, Jason Macrander, Sam Bolton, Luciana Musetti, Mesfin Tadesse, John Freudenstein, and Steve Passoa for helping me with so many little things that I couldn’t have done without. I also have to thank some of the most important mentors and advisors who have led me to become a better scientist, a better thinker, a better writer, and a more confident person. Even though you left for something much bigger and better, John Wenzel, thanks for making me believe in me, and for letting me grow as a teacher. Thanks for letting me find myself, even if it meant struggling to get there. Thanks to one of the most efficient and kind people I know - Hans Klompen, who kindly adopted me and gave me the discipline and the guidance to finish. Thank you also to my other committee members, Meg Daly and Norm Johnson, for making me stronger, even if it meant questioning myself on a daily basis. I couldn’t have done any of this without my good friend, mentor, and colleague, Keith Philips. You are a true example of what it means to be a good scientist. Thank you for sharing your passion for spider beetles with me. You gave me the ideas and motivation to keep going. And to Linda Gerofsky for being like an academic mother when I needed it most. Thank you to Judy Ridgway, from the bottom of my heart, for making me become the teacher that I am, and for giving me the support I needed to pursue my teaching goals. You have been a mentor, a confidant, a friend, and an inspiration in so many ways. vi Thanks also to the newest friends I have made at the University Center for the Advancement of Teaching: Alan Kalish, Kathryn Plank, Teresa Johnson, Stephanie Rohdieck, Laurie Maynell, Jerry Nelms, Jennie Williams, Christy Anandappa, and to my fellow graduate consultants, Sharon Ross, Lindsay Bernhagen, Spencer Robinson, and Monica Kowalski. We all took different paths to end up together in a great place. Thank you also to all the many people who loaned me specimens, especially James Harrison, and to all of those who successfully talked me into studying myrmecophiles – Gary Alpert and Bill Warner. And to another person who knows how to share his love for knowledge, Xavier Bellés, thanks for promoting spider beetle research. Finally, thank you to all my students who made me become the teacher and the scientist that I’ve become. You are the reason I finally got to this stage. I was a student once, who didn’t quite know where I would end up. Follow your dreams. I did, and even though I’m still searching for answers and for my place in life, I never stop trying. vii Vita May 2000 .......................................................James Bowie High School 2004................................................................B.S. Biology, University of Texas at Austin 2006................................................................M.S. Entomology, Texas A&M University 2006 - 2010 ....................................................Graduate Teaching Associate, Center for Life Sciences Education, The Ohio State University 2010 - present .................................................University Center for the Advancement of Teaching, The Ohio State University Publications Mynhardt G. 2011. Growing into teaching: A graduate student’s journey. Talking about Teaching 5:26-29. Philips T. K. & Mynhardt G. 2011. Description of Electrognostus intermedius , the first spider beetle from Dominican amber with implications for spider beetle phylogeny (Coleoptera Ptinidae). Entomapeiron 4:37-51. viii Mynhardt G. & Wenzel J. W. 2010. Phylogenetic analysis of the myrmecophilous Cremastocheilus Knoch (Coleoptera: Scarabaeidae: Cetoniinae) based on external adult morphology. Zookeys 34:129-140. Abbott J. C. & Mynhardt, G. (2007). Description of the larva of Somatochlora margarita (Odonata: Corduliidae). International Journal of Odonatology 10:129-136. Mynhardt G., Cognato A.I & Harris M. K.. 2007. Population genetics of the pecan weevil, Curculio caryae Horn (Coleoptera: Curculionidae), based on mitochondrial DNA data. Annals of the Entomological Society of America 100:582-590. Fields of Study Major Field: Evolution,
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