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Global Phylogeny of the Water Penny Beetles Using Both Molecular and Morphological Evidence (Co Mathew Vincent Wood Western Kentucky University, [email protected]

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Global Phylogeny of the Water Penny Beetles Using Both Molecular and Morphological Evidence (Co Mathew Vincent Wood Western Kentucky University, Maviwood@Gmail.Com Western Kentucky University TopSCHOLAR® Masters Theses & Specialist Projects Graduate School Spring 2016 Global Phylogeny of the Water Penny Beetles Using Both Molecular and Morphological Evidence (Co Mathew Vincent Wood Western Kentucky University, [email protected] Follow this and additional works at: http://digitalcommons.wku.edu/theses Part of the Biology Commons, Ecology and Evolutionary Biology Commons, and the Zoology Commons Recommended Citation Wood, Mathew Vincent, "Global Phylogeny of the Water Penny Beetles Using Both Molecular and Morphological Evidence (Co" (2016). Masters Theses & Specialist Projects. Paper 1560. http://digitalcommons.wku.edu/theses/1560 This Thesis is brought to you for free and open access by TopSCHOLAR®. It has been accepted for inclusion in Masters Theses & Specialist Projects by an authorized administrator of TopSCHOLAR®. For more information, please contact [email protected]. GLOBAL PHYLOGENY OF THE WATER PENNY BEETLES USING BOTH MOLECULAR AND MORPHOLOGICAL EVIDENCE (COLEOPTERA: PSEPHENIDAE) A Thesis Presented to The Faculty of the Department of Biology Western Kentucky University Bowling Green, Kentucky In Partial Fulfillment Of the Requirements for the Degree Master of Science By Matthew Vincent Wood May 2016 I dedicate this thesis to my parents, Markeeta and Barry Wood, who have always been there for me and supported me throughout my education and my life. ACKNOWLEDGMENTS First off I would like to thank my advisor Professor T. Keith Philips, who has helped me at every turn in completing this thesis. I would also like to thank the Western Kentucky University Biotechnology Center for access to equipment and space for my research and analysis. Thanks to Naomi Rowland for her help with optimizing molecular protocols. Thank you to the Western Kentucky University Biodiversity Center and Graduate Studies for their financial support. Thanks to Colleen and Rick Olson for providing encouragement during the final stretch of research. Thanks to James Dexter Wood for his help in collecting local specimens. Thanks to John Andersland for helping me hone my microscopic techniques. Finally I would like to thank all of the collaborators across the globe who provided the insect specimens used in this analysis, including Chi-Feng Lee (Institute of Biodiversity, National Cheng Kung University, Taiwan), William Sheppard (Essig Museum of Entomology, University of California, Berkeley), Andrew Short (Biodiversity Institute & Natural History Museum, University of Kansas), Ming- Luen Jeng (National Museum of Natural Science in Taiwan), Jiři Hájek (Department of Entomology, National Museum, Cirkusová, Czech Republic), Darren Mann (Oxford University Museum of Natural History, United Kingdom), Arlene Butwell (Griffith School of Environment, Griffith University Australia), and Masakazu Hayashi (Hoshizaki Green Foundation, Okinoshima 1659-5, Sono-chô, Izumo-shi, Shimane Pref., Japan). This work was supported by a National Science Foundation Biological Surveys and Inventories grant (DEB 0430132) awarded to T. Keith Philips. iv CONTENTS Introduction ......................................................................................................................... 1 Materials and Methods ........................................................................................................ 4 Results ................................................................................................................................. 9 Discussion ......................................................................................................................... 32 Appendix A ....................................................................................................................... 35 Appendix B ....................................................................................................................... 46 Literature Cited .............................................................................................................. 67 v LIST OF FIGURES Figure 1. CO1 Bayesian Analysis......................................................................................12 Figure 2. CO1 Parsimony Analysis...................................................................................13 Figure 3. Wingless Bayesian Analysis..............................................................................15 Figure 4. Wingless Parsimony Analysis...........................................................................16 Figure 5. Bayesian Combined Molecular Analysis..........................................................18 Figure 6. Parsimony Combined Molecular Analysis.......................................................19 Figure 7. Parsimony Combined Molecular Analysis Simplified.....................................20 Figure 8. Bayesian Morphological Analysis....................................................................22 Figure 9. Strict Consensus Parsimony Morphological Analysis......................................23 Figure 10. Majority Rule Parsimony Morphological Analysis........................................24 Figure 11. Lee et al. (2007) Morphological Analysis......................................................25 Figure 12. Total Evidence Bayesian Analysis 1..............................................................27 Figure 13. Total Evidence Bayesian Analysis 2..............................................................28 Figure 14. Total Evidence Parsimony Analysis Strict Consensus...................................29 Figure 15. Total Evidence Parsimony Analysis Majority Rule.......................................30 vi LIST OF TABLES Table 1. List of Analyzed Genera. .................................................................................. 10 Table 2. Monophyly of Proposed Subfamilies. .............................................................. 31 vii GLOBAL PHYLOGENY OF THE WATER PENNY BEETLES USING BOTH MOLECULAR AND MORPHOLOGICAL EVIDENCE (COLEOPTERA: PSEPHENIDAE) Matthew V. Wood May 2016 68 Pages Directed by: Keith Philips, Scott Grubbs, and Lawrence Alice Department of Biology Western Kentucky University The Psephenidae is a family of freshwater beetles usually found in swift streams worldwide. Their unique disc shaped and flattened larvae have made this a group of interest for scientists for centuries. Morphologically, this family has been relatively well researched, and systematically the family is fairly well known and supported as monophyletic. One issue with Psephenidae, and with many other insect groups, is the lack of the molecular phylogenetic analyses to test morphology hypothesizes. For this study, the relationships among the genera of this family were studied with both molecular and morphological data as well as combined in a total evidence analysis. DNA from specimens was extracted, amplified, and sequenced for all available genera that could be acquired locally and abroad through collaborators and their contacts in other countries. The nuclear gene Wingless (Wg) and the mitochondrial gene Cytochrome Oxidase 1 (CO1) were utilized in this study; amplification of several other nuclear genes was attempted but the results were poor and they were excluded from the analysis. After successfully sequencing these two genes from species representing nearly all of the known genera, the data were analyzed using both Bayesian and parsimony methods. Analyses were performed individually for each gene, a combined molecular analysis, using just morphological data, and a total evidence analysis using both molecular and morphological data. After analyzing the trees, definite inconsistencies viii were discovered between the current data set and the previous studies performed using only morphological characters. Individual gene analysis showed low support for the monophyly of proposed subfamilies within the psephenids, but combined molecular and total evidence analysis showed much more resolution as well as support for most but not all of the proposed subfamilies. ix INTRODUCTION Although insects are the most diverse class of animals on the planet, with over one million described species and counting, the documentation of species diversity and the hypothesized evolution of many groups are still very incomplete and includes the Psephenidae. This family of aquatic beetles (Coleoptera) is commonly referred to as the water penny beetles. Their name is derived from the larval appearance which often resembles a penny in both shape and coloration (Triplehorn & Johnson 2005). Currently there are 32 known genera and, based on phylogenetic evidence, includes 4 that are undescribed and over two hundred documented species (Lee et al. 2007). It is generally agreed that Psephenidae belongs to the infraorder Elateriformia. Insects of this infraorder tend to have a much longer larval life in comparison to their adult lives (Grimaldi & Engel, 2005). Elateriformia consists of six super-families; one of these is known as the Byrrhoidea, and including the Psephenidae, consists of most of the aquatic beetle species whose larval lifestyle is either fully aquatic or semi-aquatic. This evolutionary history is reasonably well supported by the fossil record as are many beetle phylogenies, because of their hard outer covering that fossilizes quite well (Grimaldi & Engel, 2005). The intriguing insects that make up Psephenidae have undergone a very interesting evolutionary history. It has been learned from their adult characters reflecting terrestrial habits that they had secondarily
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