Hymenoptera: Pompilidae)
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Utah State University DigitalCommons@USU All Graduate Theses and Dissertations Graduate Studies 5-2014 Molecular Systematics, Historical Biogeography, and Evolution of Spider Wasps (Hymenoptera: Pompilidae) Juanita Rodriguez Utah State University Follow this and additional works at: https://digitalcommons.usu.edu/etd Part of the Geology Commons Recommended Citation Rodriguez, Juanita, "Molecular Systematics, Historical Biogeography, and Evolution of Spider Wasps (Hymenoptera: Pompilidae)" (2014). All Graduate Theses and Dissertations. 2790. https://digitalcommons.usu.edu/etd/2790 This Dissertation is brought to you for free and open access by the Graduate Studies at DigitalCommons@USU. It has been accepted for inclusion in All Graduate Theses and Dissertations by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. MOLECULAR SYSTEMATICS, HISTORICAL BIOGEOGRAPHY, AND EVOLUTION OF SPIDER WASPS (HYMENOPTERA: POMPILIDAE) by Juanita Rodriguez A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in Biology Approved: ________________________________ _______________________________ James P. Pitts Carol D. von Dohlen Major Professor Major Professor ________________________________ _______________________________ Kimberly A. Sullivan Edward W. Evans Committee Member Committee Member ________________________________ _______________________________ James Strange Steve Larson Committee Member Committee Member _________________________ Mark R. McLellan Vice President for Research and Dean of the School of Graduate Studies UTAH STATE UNIVERSITY Logan, Utah 2014 ii Copyright © Juanita Rodriguez 2014 All Rights Reserved iii ABSTRACT Molecular Systematics, Historical Biogeography, and Evolution of Spider Wasps (Hymenoptera: Pompilidae) by Juanita Rodriguez, Doctor of Philosophy Utah State University, 2014 Major Professors: James P. Pitts and Carol D. von Dohlen Department: Biology Spider wasps are solitary parasitoids that use one spider to lay a single egg. Even though their behavior seems homogeneous, the features pertaining to nesting and hunting behavior are diverse for different species. There are approximately 5,000 described species, in 120 genera, but there are probably many undescribed species. The systematics of Pompilidae has been studied in recent years, but only morphological data have been used for this purpose. Because of the morphological homogeneity of spider wasps, molecular data may prove promising for understanding the systematics of the group. Furthermore, dated molecular phylogenies calibrated with fossil data may allow studying the historical biogeography and evolution of the group. I used the nuclear molecular markers elongation factor–1 α F2 copy (EF1), long–wavelength rhodopsin (LWRh), wingless (Wg), RNA polymerase II (Pol2), the D2–D3 regions of the 28S ribosomal RNA (28S), and the mitochondrial Cytochrome C Oxidase I (COI) in a Bayesian and Maximum Likelihood framework, to reconstruct the phylogenies of four main iv Pompilidae groups: the subfamily Pompilinae, the tribe Aporini, the genus Psorthaspis, and the genus Drepanaporus. I also studied the fossil Pompilidae, and used those results to produce time-calibrated phylogenies of Pompilinae, Aporini, and Psorthaspis. Molecular phylogenetic results support the utility of the use of molecular markers for species delimitation and sex-associations in Pompilidae. In addition, the use of dated phylogenies supports the correlation of host use with diversification rate-shifts, the coevolution of mimicry between pompilids and velvet ants, and various biogeographical hypotheses never tested before for spider wasps. (240 pages) v PUBLIC ABSTRACT Molecular Systematics, Historical Biogeography, and Evolution of Spider Wasps (Hymenoptera: Pompilidae) by Juanita Rodriguez, Doctor of Philosophy Utah State University, 2014 Major Professors: James P. Pitts and Carol D. von Dohlen Department: Biology The study of the diversity and classification of any group of organisms provides a foundation for further scientific studies in ecology, evolution, and conservation. Insects are among the most diverse organisms that inhabit the planet, but knowledge of their diversity and classification is still limited. One understudied group of insects is spider wasps. These are solitary parasitoids that use one spider to lay a single egg. There are approximately 5,000 described species, and many more to be described. Unfortunately, fewer than 10 scientists worldwide study these insects. One reason the group has not been very well studied is the difficulty in telling species apart. This makes their classification troubling. With the advent of molecular genetics methods, the use of molecular data to understand the classification and evolution of various groups is now possible. My dissertation uses molecular data to understand the classification of spider wasps, as well as their evolutionary relations. The evolutionary trees produced by these analyses are helpful to study the causes of current distributions of species, the diversification and the vi evolution of the group. Molecular phylogenetic results support the utility of the use of molecular markers for species delimitation and sex-associations in Pompilidae, the correlation of host use with diversification rate-shifts, the coevolution of mimic pompilids with velvet ants, and various biogeographical hypotheses never tested before for spider wasps. (240 pages) vii ACKNOWLEDGMENTS I would like to thank Carol von Dohlen and James Pitts, my academic advisors, for providing all the grounds, support, and knowledge necessary for my dissertation research. I would also like to thank Cecilia Waichert for being my friend, sister, co- author, and collaborator. I would like to thank my committee members; Kim Sullivan, Ted Evans, Jamie Strange, and Steve Larson, for all their support. I would like to thank Fernando Fernandez, my advisor for my master’s program and collaborator, for all the efforts he made through my undergraduate and graduate years to make me a successful scientist. I would also like to thank past and present lab mates Joseph Wilson, Emily Sadler, Cecilia Vieira, Sarah Clark, Catherine Clark, Nicole Boehme, and Dave Tanner for all the encouragement and support; especially Joe Wilson for being a great teacher, mentor, and collaborator, and Emily Sadler for being the best travel buddy and supporting me through all those stressful times. I would like to thank Dr. Marius Wasbauer for sharing all his knowledge with our lab. I would like to thank Wilford Hanson for funding international collecting expeditions. A NSF grant, two women and gender research grants, and various UAES publishing grants funded this research. I would like to thank the Graduate School for the dissertation completion grant. I would like to thank my Logan family, especially Jonathan Koch, Armen Armaghanyan, Rodrigo Ferreira, Mercedes Roman and Jane Li, Filomena Arrieta, and baby Artur. They kept me sane, well fed, loved, and happy. Finally and more importantly, I would like to thank my parents, Iomara Arrieta and Francisco Rodriguez; and my brother, Manuel Rodriguez, without whom this would viii not have been possible. I owe huge thanks to Jaime Florez for being the best partner/cook/statistitian ever. Juanita Rodriguez ix CONTENTS Page ABSTRACT....................................................................................................................... iii PUBLIC ABSTRACT .........................................................................................................v ACKNOWLEDGMENTS ................................................................................................ vii LIST OF TABLES............................................................................................................. xi LIST OF FIGURES .......................................................................................................... xii CHAPTER 1. INTRODUCTION.......................................................................................................1 2. TWO NEW GENERA AND THREE NEW SPECIES OF FOSSIL POMPILIDAE FROM AMBER AND THEIR EVOLUTIONARY IMPLICATIONS....................23 3. REVIEW OF FOSSIL SPIDER WASPS (HYMENOPTERA: POMPILIDAE) WITH THE DESCRIPTION OF A NEW SPECIES................................................44 4. ASSESSING SPECIES BOUNDARIES AND SEX-ASSOCIATIONS IN THE GENUS DREPANAPORUS (HYMENOPTERA: POMPILIDAE), WITH COMPARISON OF THE UTILITY OF CYTOCHROME C OXIDASE I AND A NUCLEAR MOLECULAR MARKER, AND THE DESCRIPTION OF A NEW SPECIES OF DREPANAPORUS .............................................................................80 5. HISTORICAL BIOGEOGRAPHY OF THE WIDESPREAD SPIDER WASP TRIBE APORINI (HYMENOPTERA: POMPILIDAE) .......................................114 6. MÜLLERIAN MIMICRY AS A RESULT OF CODIVERGENCE BETWEEN VELVET ANTS AND SPIDER WASPS...............................................................149 7. MOLECULAR PHYLOGENY OF POMPILINAE (HYMENOPTERA: POMPILIDAE): EVIDENCE FOR RAPID DIVERSIFICATION AND HOST SHIFTS IN SPIDER WASPS.................................................................................171 8. SUMMARY AND CONCLUSIONS......................................................................211 x APPENDICES .................................................................................................................214 CURRICULUM VITAE .................................................................................................220