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The Tingidae (Hemiptera: Heteroptera) of Southern Central America

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The Tingidae (Hemiptera: Heteroptera) of Southern Central America THE TINGIDAE (HEMIPTERA: HETEROPTERA) OF SOUTHERN CENTRAL AMERICA (WITH AN EMPHASIS ON COSTA RICA) A Thesis Submitted to the Graduate Faculty of the North Dakota State University of Agriculture and Applied Science By Alexander Harris Knudson In Partial Fulfillment of the Requirements for the Degree of MASTER OF SCIENCE Major Department: Entomology April 2018 Fargo, North Dakota North Dakota State University Graduate School Title THE TINGIDAE (HEMIPTERA: HETEROPTERA) OF SOUTHERN CENTRAL AMERICA (WITH AN EMPHASIS ON COSTA RICA) By Alexander Harris Knudson The Supervisory Committee certifies that this disquisition complies with North Dakota State University’s regulations and meets the accepted standards for the degree of MASTER OF SCIENCE SUPERVISORY COMMITTEE: David A. Rider Co-Chair Janet J. Knodel Co-Chair Rebecca B. Simmons Mark A. Boetel Approved: April 12, 2018 Frank Casey Date Department Chair ABSTRACT The genera of Tingidae of the neotropics are herein diagnosed and a key for their identification is also provided. Five new genera are described from Central America, two from Panama, two from Costa Rica, and one new genus is described from Mexico. This brings the total of neotropical genera to 74. One new species, Mexibyrsa woolleyi is described from Mexico. The Tingidae of southern Central America (Nicaragua, Costa Rica, and Panama) are reviewed in detail. There are now 153 described species spread among 43 genera. Thirty-two new species from southern Central America are described. Identification keys are provided for all species found in the study region. iii ACKNOWLEDGMENTS Over the course of my Master’s degree I have made many contacts and friends in and outside of the field of entomology. There are literally countless people that have made this work possible and I wish to thank all who have participated, assisted, and provided for this project. Whenever possible, I have tried to record all individuals with their contributions, and to the best of my knowledge, they are listed below. First of all I would like to express sincere gratitude towards my advisors Dr. David A. Rider and Dr. Janet J. Knodel; without their support, knowledge, and sage advice, I would not have had this wonderful opportunity to work on this group of fascinating insects. I would also like to thank my committee members Dr. Rebecca Simmons and Dr. Mark A. Boetel for their encouragement and advice. Additionally, I thank David A. Rider, for his patience and support of my endeavors and for continuously pushing me to be my best. For additional support and career advice I thank Dr. Thomas J. Henry: United States Department of Agriculture, Systematic Entomology Laboratory, Dr. Theodore “Jim” Lewis: Instituto National de Biologia (INBio); Dr. Dennis D. Kopp: United States National Museum of Natural History (USNM); Dr. Joseph Schaffner: Texas A&M University (TAMU), and Dr. Alfred G. Wheeler Jr.: Clemson University. I also thank Laura Torres Miller: West Virginia Department of Agriculture, for her willingness to help and her enthusiastic support. I am indebted to all individuals who provided specimens for this large project and acknowledge them in the following orations. Despite contacting numerous museums and receiving many specimens on loan from multiple collections, I owe the greatest thanks to Max Barclay and Mick Webb: Natural History Museum of London (BMNH); Thomas J. Henry: iv USNM; Jim Lewis: INBio; Edward G. Riley, John D. Oswald, and Joseph Schaffner: TAMU for providing the majority of specimens used in this study. Irrespective of the holdings of each collection, all specimens provided were of upmost importance; Craig Brabant: University of Wisconsin Madison; Christopher Dietrich: Illinois Natural History Survey, Champaign, IL; Gerald Fauske: North Dakota State University, Fargo; Susan Halbert: Florida State Collection of Arthropods, Gainesville; Paul Hanson: University of Costa Rica, San José; Steve Heydon: Bohart Museum of Entomology University of California Davis; E. Richard Hoebeke and Joseph V. McHugh: University of Georgia, Athens; John M. Leavengood Jr.: United States Department of Agriculture, Animal Plant Health Inspection Service, Miami, FL; Jean Michel Maes: Entomological Museo de León, Nicaragua; Crystal Maier: Field Museum of Natural History, Chicago, IL; José Montero Ramírez: University of Georgia, Athens, Costa Rica Campus; late Norm Penny: California Academy of Sciences, San Francisco; John Rawlins: Carnegie Museum of Natural History, Pittsburgh, PA; David A. Rider: Personal Collection, Fargo, ND; Ruth Salas: American Museum of Natural History, New York, NY; Barb Sharanowski: University of Manitoba, Winnipeg, Canada: Robert Sites: University of Missouri, Columbia; and Jefferson Vaughn: University of North Dakota, Grand Forks. A project this massive would be incomplete without photographs, as such technical assistance with taking photographs was generously provided by Patrick Beauzay and Gerald Fauske: North Dakota State University, as well as Juan Mata Lorenzen and Jim Lewis: INBio. I also graciously appreciate receiving photographs of type specimens from Bo Delling: Swedish Museum of Natural History, Stockholm; Thomas Henry: USNM; Gabriel Mejdalani: v Universidade Federal do Rio de Janeiro, Brazil; Amoret Spooner: Oxford University, Oxford, England; Vanessa Verdecia: Carnegie Museum of Natural History; and Mic Webb: BMNH. Assistance acquiring collecting permits was generously provided by Barbara Lewis and Francisco Campos Rivera: Organization of Tropical Studies. Ground transportation was provided by Luis F. Delgado M. and Luis of Natural Car Rental. Partial funding for this project was generously provided by the NDSU Entomology Scholarship committee in the form of two Knippling thesis enhancement grants, one of which was used to visit Costa Rica. Additional funding was provided by Dr. John D. Oswald of TAMU. vi DEDICATION I wish to dedicate this work to my best friend and amor, my wife Veronica. Thank you for your compassionate support and tender care through this arduous journey we call life. Without your continual devotion, this project would have never come to fruition. ¡Besitos y abrazos del tu osito polar del norte! vii DISCLAIMER This manuscript in its entirety should not be recognized as a valid publication. Any species names proposed herein are only manuscript names and should not be regarded as published until they appear elsewhere in a refereed taxonomic journal. viii TABLE OF CONTENTS ABSTRACT ............................................................................................................................. iii ACKNOWLEDGMENTS ..........................................................................................................iv DEDICATION ......................................................................................................................... vii DISCLAIMER ........................................................................................................................ viii LIST OF FIGURES .................................................................................................................... x CHAPTER ONE: A KEY TO THE GENERA OF TINGIDAE (HEMIPTERA: HETEROPTERA) OF THE NEOTROPICS ................................................................................ 1 Abstract .................................................................................................................................. 1 Introduction ............................................................................................................................ 1 Materials and Methods ............................................................................................................ 3 Results .................................................................................................................................... 7 Literature Cited .................................................................................................................... 73 CHAPTER TWO: THE TINGIDAE (HEMIPTERA: HETEROPTERA) OF SOUTHERN CENTRAL AMERICA (NICARAGUA, COSTA RICA, AND PANAMA) .............................. 88 Abstract ................................................................................................................................ 88 Introduction .......................................................................................................................... 90 Materials and Methods ......................................................................................................... 92 Taxonomic Review of the Tingidae of Southern Central America ......................................... 96 Discussion .......................................................................................................................... 296 Literature Cited ................................................................................................................... 299 CHAPTER THREE: CONCLUSIONS .................................................................................... 312 ix LIST OF FIGURES Figure Page 1.1: Map of the Neotropical realm ............................................................................................... 4 1.2: General tingid anatomy, dorsal aspect .................................................................................. 5 1.3: General tingid anatomy, lateral aspect .................................................................................. 6 1.4: Mexibyrsa woolleyi, new species .......................................................................................
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