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Coleoptera: Cucujoidea) Matthew Immelg Louisiana State University and Agricultural and Mechanical College, [email protected]

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Coleoptera: Cucujoidea) Matthew Immelg Louisiana State University and Agricultural and Mechanical College, Phalacrid@Gmail.Com Louisiana State University LSU Digital Commons LSU Doctoral Dissertations Graduate School 2011 Revision and Reclassification of the Genera of Phalacridae (Coleoptera: Cucujoidea) Matthew immelG Louisiana State University and Agricultural and Mechanical College, [email protected] Follow this and additional works at: https://digitalcommons.lsu.edu/gradschool_dissertations Part of the Entomology Commons Recommended Citation Gimmel, Matthew, "Revision and Reclassification of the Genera of Phalacridae (Coleoptera: Cucujoidea)" (2011). LSU Doctoral Dissertations. 2857. https://digitalcommons.lsu.edu/gradschool_dissertations/2857 This Dissertation is brought to you for free and open access by the Graduate School at LSU Digital Commons. It has been accepted for inclusion in LSU Doctoral Dissertations by an authorized graduate school editor of LSU Digital Commons. For more information, please [email protected]. REVISION AND RECLASSIFICATION OF THE GENERA OF PHALACRIDAE (COLEOPTERA: CUCUJOIDEA) A Dissertation Submitted to the Graduate Faculty of the Louisiana State University and Agricultural and Mechanical College in partial fulfillment of the requirements for the degree of Doctor of Philosophy in The Department of Entomology by Matthew Gimmel B.S., Oklahoma State University, 2005 August 2011 ACKNOWLEDGMENTS I would like to thank the following individuals for accommodating and assisting me at their respective institutions: Roger Booth and Max Barclay (BMNH), Azadeh Taghavian (MNHN), Phil Perkins (MCZ), Warren Steiner (USNM), Joe McHugh (UGCA), Ed Riley (TAMU), Mike Thomas and Paul Skelley (FSCA), Mike Ivie (MTEC/MAIC/WIBF), Richard Brown and Terry Schiefer (MEM), Andy Cline (CDFA), Fran Keller and Steve Heydon (UCDC), Cheryl Barr (EMEC), Norm Penny and Jere Schweikert (CAS), Mike Caterino (SBMN), Michael Wall (SDMC), Don Arnold (OSEC), Zack Falin (SEMC), Arwin Provonsha (PURC), Cate Lemann and Adam Slipinski (ANIC), and Harold Labrique (MHNL). I thank Richard Thompson (BMNH) and Zdeněk Švec (Prague, Czech Republic) for their support and discussing with me some tedious details concerning phalacrid taxonomy. I thank Roger Booth and Al Newton for taking time to assist me with clarification of the type status of certain George Champion specimens. Many individuals generously donated fresh specimens to help me in my research, but I especially thank Andy Cline (CDFA) and Jim Cornell (Charlotte, North Carolina) for their particularly generous contributions. A tremendous thanks goes to Stephanie Gross (Louisiana State University) for her help, guidance, and work ethic with regard to the molecular portion of this work; that section would not have been possible without her. Hector Urbina (LSU) also donated his time and expertise to issues regarding the molecular analysis. I benefitted daily from scientific and other conversations with my museum-mates, Michael Ferro, Jong-Seok Park, Alexey Tishechkin, Igor Sokolov, and Stephanie Gil. Victoria Bayless has shown me, by her own example at the LSAM, what a proper insect museum should look like. I owe her an enormous thanks for all of her assistance over the years and for being a voice of sanity during my time here. I thank John Lawrence (Gympie, Australia) and Adam Ślipiński (CSIRO, Canberra, Australia) for their encouragement and advice. I thank Rich Leschen (Landcare Research, Auckland, New Zealand) for insisting that I examine Phalacridae with the widest possible lens, and for his lively criticisms of my morphological analysis. I thank Warren Steiner (USNM) for sharing his drawings and conclusions from an unfinished manuscript on Acylomus, and for being so generous with his time and skill while passing me the torch of phalacrid knowledge. I thank the members of my committee: Dorothy Prowell, Meredith Blackwell, and Frederick Sheldon, for their helpful advice and challenging me when necessary. Most of all, I thank my major professor, Chris Carlton, for his friendship and for his unique non-meddling but enthusiastic and knowledgeable style of advising, which provided inspiration for and meshed extremely well with my own working style. I am a much better scientist as a result of my interactions with Chris. On a personal note, I would like to thank Crystal Maier for her love, support, encouragement, and patience during the time leading up to the completion of this life-consuming project. I thank my parents, Dennis Gimmel and Susan Allison, for providing me every opportunity in their power, and allowing me just the right amount of freedom, to pursue my interest in natural history. Thanks, Mom and Dad. This research was partially funded by NSF DEB-0516311 to Chris Carlton and Victoria Bayless. Travel to foreign museums was partially funded by an Ernst Mayr Travel Grant from Harvard University. ii TABLE OF CONTENTS ACKNOWLEDGMENTS .............................................................................................................. ii LIST OF FIGURES .........................................................................................................................v ABSTRACT ................................................................................................................................. viii CHAPTER 1 INTRODUCTION ...........................................................................................................1 1.1 PHYLOGENETIC POSITION OF PHALACRIDAE.......................................1 1.2 TAXONOMIC HISTORY OF PHALACRIDAE .............................................2 1.3 IMMATURE STAGES AND HABITS OF PHALACRIDAE .........................4 1.4 PHALACRIDAE IN THE FOSSIL RECORD ..................................................5 2 MATERIALS AND METHODS .....................................................................................6 2.1 SPECIMENS......................................................................................................6 2.2 GUIDELINES FOR GENERIC REVISION .....................................................8 2.3 INTRODUCTION TO PHYLOGENETIC ANALYSIS ...................................9 2.4 MORPHOLOGICAL ANALYSIS ....................................................................9 2.5 MOLECULAR ANALYSIS ............................................................................20 2.6 COMBINED ANALYSIS ...............................................................................21 2.7 OTHER METHODS ........................................................................................23 3 PHYLOGENETIC RESULTS .......................................................................................24 3.1 RESULTING TREES ......................................................................................24 3.2 DISCUSSION ..................................................................................................24 3.3 LITERATURE CITED (FOR CHAPTERS 1–3) ............................................26 4 SYSTEMATIC TREATMENT .....................................................................................36 4.1 FAMILY PHALACRIDAE LEACH, 1815 .....................................................36 4.2 REVISED CHECKLIST OF HIGHER TAXA OF PHALACRIDAE ............39 4.3 KEY TO WORLD GENERA OF PHALACRIDAE .......................................43 4.4 SUBFAMILY PHAENOCEPHALINAE MATTHEWS, 1899 ......................53 4.5 SUBFAMILY EUSTILBINAE GUILLEBEAU, 1892 ...................................60 4.6 SUBFAMILY BIOPHYTINAE GUILLEBEAU, 1894 ..................................75 4.7 SUBFAMILY PHALACRINAE LEACH, 1815 .............................................81 4.8 OLIBROPORINAE, SUBFAM. NOV. ...........................................................87 4.9 SUBFAMILY OCHROLITINAE GUILLEBEAU, 1894 ...............................98 4.10 SUBFAMILY OLIBRINAE GUILLEBEAU, 1892 ....................................102 4.11 OLIBROSOMINAE, SUBFAM. NOV. ......................................................111 4.12 PHALACRIDAE INCERTAE SEDIS ..........................................................118 4.13 NOMEN INQUIRENDUM .........................................................................146 4.14 TAXA REMOVED FROM PHALACRIDAE ............................................147 5 SUMMARY AND CONCLUSIONS ..........................................................................230 iii REFERENCES (FOR CHAPTER 4 AND APPENDIX B) .........................................................231 APPENDIX A: MORPHOLOGICAL CHARACTER MATRIX ...............................................260 APPENDIX B: WORLD CATALOGUE OF PHALACRIDAE .................................................261 VITA ............................................................................................................................................459 iv LIST OF FIGURES Figure 1. Phalacrid species-group taxa described over time, 1767–2006. ...................................... 4 Figure 2. Strict consensus tree from maximum parsimony analysis of morphological matrix. ... 29 Figure 3. Strict consensus tree from maximum parsimony analysis of 18S rDNA sequence data ................................................................................................................................................ 30 Figure 4. Resulting tree from Bayesian analysis of 18S rDNA sequence data.. ........................... 31 Figure 5. Best ML tree from maximum likelihood analysis of 18S rDNA sequence data.. ......... 32 Figure 6. Strict consensus tree (of 86 MP trees) from maximum parsimony analysis of concatenated morphological + 18S rDNA sequence data. ..........................................................
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