Diptera: Blephariceridae) with an Emphasis on the Eastern Blepharicera Tenuipes Group Hogue
Total Page:16
File Type:pdf, Size:1020Kb
University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 12-2010 Phylogenetic analysis of the Nearctic Blepharicera Macquart (Diptera: Blephariceridae) with an emphasis on the eastern Blepharicera tenuipes group Hogue Amanda Jane Jacobson [email protected] Follow this and additional works at: https://trace.tennessee.edu/utk_graddiss Part of the Entomology Commons Recommended Citation Jacobson, Amanda Jane, "Phylogenetic analysis of the Nearctic Blepharicera Macquart (Diptera: Blephariceridae) with an emphasis on the eastern Blepharicera tenuipes group Hogue. " PhD diss., University of Tennessee, 2010. https://trace.tennessee.edu/utk_graddiss/889 This Dissertation is brought to you for free and open access by the Graduate School at TRACE: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of TRACE: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Amanda Jane Jacobson entitled "Phylogenetic analysis of the Nearctic Blepharicera Macquart (Diptera: Blephariceridae) with an emphasis on the eastern Blepharicera tenuipes group Hogue." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the equirr ements for the degree of Doctor of Philosophy, with a major in Plants, Soils, and Insects. John K. Moulton, Major Professor We have read this dissertation and recommend its acceptance: Ernest C. Bernard, Juan Luis Jurat-Fuentes, Randall L. Small Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official studentecor r ds.) To the Graduate Council: I am submitting herewith a dissertation written by Amanda Jane Jacobson entitled “Phylogenetic analysis of the Nearctic Blepharicera Macquart (Diptera: Blephariceridae) with an emphasis on the eastern Blepharicera tenuipes group Hogue.” I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Plants, Soils and Insects. John K. Moulton, Major Professor We have read this dissertation and recommend its acceptance: Ernest C. Bernard Juan Luis Jurat-Fuentes Randall L. Small Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official student records.) Phylogenetic analysis of the Nearctic Blepharicera Macquart (Diptera: Blephariceridae) with an emphasis on the eastern Blepharicera tenuipes group Hogue A Dissertation Presented for the Doctor of Philosophy Degree The University of Tennessee, Knoxville Amanda Jane Jacobson December 2010 © 2010 AMANDA J. JACOBSON ii ACKNOWLEDGEMENTS This dissertation, and the work it represents, would not have been possible without the assistance and guidance of my major professor, Dr. John K. Moulton. I also thank my other committee members: Drs. Ernest C. Bernard, Juan Luis Jurat-Fuentes, and Randall L. Small, for their time, suggestions, and encouragement. I would especially like to thank Dr. Gregory W. Courtney for serving as an external advisor and sparking my interest in such a fascinating group of insects. I am indebted to the many individuals who had a part in making this research possible: Ken Kingsley at Zion National Park whose interest and many efforts provided DNA-quality specimens of Blepharicera zionensis for the molecular aspects of this paper. Dr. Peter Zwick took time out of his collecting schedule for attempts to locate B. fasciata. Dr. John K. Moulton collected B. tenuipes in Canada and located many ideal blepharicerid habitats in the southeast. Dr. John Dunlap at the Bioimaging and Electron Microscopy Facility at the University of Tennessee served as a mentor for acquiring the techniques associated with scanning electron microscopy. Drs. Randall L. Small, P. Brandon Matheny, and Brian O’Meara were helpful obtaining and learning the various phylogenetic programs. I am most grateful for my loving parents, Larry and Sandy Jacobson. They taught me the power of hard work and compassion, and I am truly a better person because of their support and encouragement. Finally, I would like to thank Dr. Gregory R. Curler. He is a knowledgeable colleague and a reliable partner. I appreciate his assistance and excitement during numerous collecting trips. Greg’s talent in biological illustration provided the illustrations for the six new eastern Nearctic Blepharicera and his collecting intuition ultimately led to the discovery of B. crista in Virginia. iii ABSTRACT Abstract. The eastern Nearctic fauna of Blepharicera Macquart (Diptera: Blephariceridae) is revised to include twenty-three species, six of which are new to science. Descriptions of the larvae, pupae, and adults of B. amnicula n. sp., B. conifera n. sp., B. crista n. sp., B. enoristera n. sp., B. hillabee n. sp., and B. opistera n. sp. are presented. Keys to instar IV larvae, pupae, and adults of all eastern Blepharicera (except B. caudata Courtney) are provided. Phylogenetic studies were conducted to determine the relationships between eastern and western Nearctic Blepharicera and among species within these groups. Larvae, pupae, and adults were available for all known Nearctic species except B. caudata and B. micheneri Alexander. Molecular data acquired from two genes and morphological data for 44 characters were used to test previous phylogenetic hypotheses on the historical relationships and biogeography of Nearctic Blepharicera. Analyses using maximum parsimony, maximum likelihood, and Bayesian inference were conducted. Resulting phylogenies support monophyly of the B. tenuipes and B. micheneri groups and suggest that multiple species complexes may exist within the B. tenuipes group. Keywords: net-winged midges, United States, new species, lotic, phylogeny, Big Zinc Finger 2, BZF2, NADH dehydrogenase 2, ND2 iv TABLE OF CONTENTS CHAPTER 1. GENERAL INTRODUCTION .................................................................. 1 Dissertation organization ....................................................................................... 7 Literature cited....................................................................................................... 9 CHAPTER 2. NEW SPECIES OF NET-WINGED MIDGES OF THE GENUS BLEPHARICERA MACQUART (DIPTERA: BLEPHARICERIDAE) FROM EASTERN NORTH AMERICA........................................................................................................ 12 Abstract ............................................................................................................... 13 Introduction ......................................................................................................... 13 Methods............................................................................................................... 16 Taxonomy of eastern Blepharicera ...................................................................... 17 Blepharicera Macquart 1843.......................................................................... 17 Blepharicera tenuipes group Hogue 1978....................................................... 19 Keys to species of eastern Nearctic Blepharicera ........................................... 20 Species diagnoses and descriptions ................................................................ 32 Discussion ........................................................................................................... 74 Acknowledgements.............................................................................................. 76 Literature cited..................................................................................................... 77 CHAPTER 3. PHYLOGENETIC ANALYSIS OF NEARCTIC BLEPHARICERA MACQUART (DIPTERA: BLEPHARICERIDAE) ........................................................ 81 Abstract ............................................................................................................... 82 Introduction ......................................................................................................... 82 Methods............................................................................................................... 86 Results............................................................................................................... 105 Discussion ......................................................................................................... 123 Acknowledgements............................................................................................ 125 Literature cited................................................................................................... 126 CHAPTER 4. GENERAL CONCLUSIONS.................................................................. 130 Literature cited................................................................................................... 133 VITA............................................................................................................................. 134 v LIST OF TABLES Table 1.1. Location and species names for Nearctic Blepharicera..................................... 5 Table 2.1. Location and species names for Nearctic Blepharicera..................................