A Taxonomic Revision of the Nearctic Species of Apateticus Dallas and Podisus Herrich-Schaeffer (Heteroptera: Pentatomidae: Asopinae)
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AN ABSTRACT OF THE THESIS OF Kathryn Ann Phillips for the degree of Doctor of PhilosoPhY in Entomology presented on December 22, 1982 Title: A Taxonomic Revision of the Nearctic Soecies of Apateticus Dallas and Podisus Herrich-Schaeffer (Heteroptera: Pentatomidae: Asopinae) Redacted for privacy Abstract approved: John D. Lattin The Nearctic species of the pentatomid genera Apateticus Dallas and Podisus Herrich-Schaeffer were revised taxonomically. Approximately 8500 Nearctic specimens plus 500 extralimital examples were examined. Five species of Apateticus and 9 of Podisus, including 1 incidental Nearctic species, are redescribed. Important characters for species discrimination include shape of the male claspers, length of the ventral abdominal spine, and characteristics of the pronotum. Female genitalic characters are not reliable for species-level distinctions within each genus. Re-evaluation of previously used characters and additional male and female genitalic traits support the idea that these genera should remain separate. A cladistic analysis was performed on the species of each genus. One new species, Podisus brevispinus, is described. The latter, heretofore known as P. modestus (Dallas), has long been associated with a type specimen which is a synonym of P. maculiventris (Say). Examination of Rhaphigaster aggressor Walker, listed as synonymous with P modestus, showed the former to be a Neotropical Podisus. The following synonymies are suggested (junior synonym first): A. crocatus (Uhler) = A. bracteatus (Fitch), P. pallens (Sega) = P. maculiventris (Say) (reaffirmed), P. modestus (Dallas) = P. maculiventris (Say), and P. rubromaculatus Distant = P. acutissimus Stk. Podisus maculiventris, P. brevispinus, P. placidus Uhler, P. serieventris Uhler, and hEattLizas bracteatus are widely distributed in the United States and Canada. A. cynicus (Say) occurs from northeastern North America south to Florida and west to Arizona. P. acutissimus, P. sagitta (F.), and A. lineolatus (H.-S.) occur in the southern U. S., Mexico, Central America, and the Caribbean. P. mucronatus Uhler is confined to Florida and Cuba, while P. congrex (Stil) occurs from central Mexico into South America. Localized distributions are found in A. anatarius Van Duzee, A. marginiyentris (Stg.1), and P. fretus, Olsen. Apateticus and Podisus prey on a wide variety of insects, notably larval Lepidoptera, Hymenoptera, and Coleoptera. Podisus maculiventris is being investigated as a biological control agent of lepidopterous pests of soybean in the United States and has been introduced into Europe and Asia with varying success. Lists of known prey are presented and major biological literature for each species is reviewed. A Taxonomic Revision of the Nearctic Species of Apateticus Dallas and Podisus Herrich-Schaeffer (Heteroptera: Pentatomidae: Asopinae) by Kathryn Ann Phillips A THESIS submitted t Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Commencement June 1983 APPROVED: Redacted for privacy Professor/6f Entomology in charge ofmajor Redacted for privacy Head of Department of Entomolo Redacted for privacy Dean of Gradua scnool ci Date thesis is presented_ December 22, 1982 Typed by Kathryn Ann Phillins ACKNOWLEDGEMENTS I express my sincere appreciation to the many people who assisted and encouraged me throughout this undertaking. Financial support was provided through a Resident Instruction graduate teaching assistantship from September, 1979 through December 1980 and by the Science Technical Advisory Fund through a research assistantship from January 1981 through December 1982. I received support for computer services for unsponsored research from the Oregon State University Computer Center. Type specimens were lent by the American Museum of Natural History, New York, (AMNH), Dr. R. T. Schuh; British Museum (Natural History), London (BM), Dr. W. R. Dolling; California Academy of Sciences, San Francisco, (CAS), Dr. P. H. Arnaud, Jr.; Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, (MCZ), Mr. Alfred F. Newton, Jr.; Naturhistorisches Museum, Vienna, Dr. A. Kaltenbach; Naturhistoriska Riksmuseet, Stockholm, Dr. P. Lindskog; United States National Museum, Washington, D. C., (USNM), Mr. T. J. Henry. The following institutions and individuals kindly made loans of specimens: American Museum of Natural History, New York (AMNH), Dr. R. T. Schuh; Arizona State University, Tempe (ASU), Dr. F. F. Hasbrouck; California Academy of Sciences, San Francisco, (CAS), Dr. P. H. Arnaud, Jr.; Carnegie Museum of Natural History, Pittsburgh, PA, (CMNH), Dr. G. Ekis; Colorado State University, Fort Collins, (CSU), Dr. W. D. Fronk; Cornell University, Ithaca, NY, (CU), Dr. L. L. Pechuman; Joe E. Eaer Collection, Tampa, FL, (JE), Dr. J. E. Eger; Illinois Natural History Survey, Urbana, (INH), Mr. D. Voegtlin; Illinois State Museum, Springfield, (ISM), Dr. E. D. Cashatt; Iowa State University, Ames, (ISU), Dr. R. E. Lewis; Louisiana State University, Baton Rouge, (LSU), Dr. J. B. Chapin; Michigan State University, East Lansing, (MIS), Dr. R. L. Fischer; Montana State University, Bozeman, (MSU), Ms. S. D. Rose; Museum of Comparative Zoology, Harvard University, Cambridge, MA, (MCZ), Ms. M. Hathaway; Natural History Museum of Los Angeles County, CA, (LACM), Dr. C. L. Hogue; North Carolina State University, Raleigh, (NCSU), Ms. C. Parron; North Dakota State University, Fargo, (NDSU), Dr. E. U. Balsbaugh, Jr.; Northern Arizona University, Flagstaff, (NAU), Dr. C. D. Johnson; Ohio State University, Columbus, (OHS), Dr. C. A. Triplehorn; Oklahoma State University, Stillwater, (OKS), Dr. W. A. Drew; Oregon State University, Corvallis, (OSU), Dr. J. D. Lattin; Peabody Museum, Yale University, New Haven, CT, (YLU), Mr. D. G. Furth; Pennsylvania State University, University Park, (PSU), Dr. K. C. Kim; Purdue University, West Lafayette, IN, (PDU), Mr. A. V. Provonsha; San Diego Society of Natural History Museum, CA, (SDCM), Mr. D. K. Faulkner; Gary M. Stonedahl Collection, Corvallis, OR, (GMS), Dr. G. M. Stonedahi; Texas A & M University, College Station, (TAM), Dr. J. C. Schaffner; Texas Tech University, Lubbock, (TTU), Dr. O. F. Francke; Donald B. Thomas, Jr. Collection, Lincoln, NE, (DBT), Dr. D. B. Thomas, Jr.; University of Alberta, Edmonton, (UAL), Dr. G. E. Ball, Mr. D. Shpeley; University of Arizona, Tucson, (UAZ), Dr. F. G. Werner; University of British Columbia, Vancouver, (UEC), Mr. S. G. Cannings; University of California, Berkeley, (UCB), Dr. J. A. Powell, Mr. J. B. Whitfield; University of California, Riverside, (UCR), Mr. S. I. Frontier; University of Connecticut, Storrs, (UCT), Ms. J. E. O'Donnell; University of Georgia, Athens, (UGA), Dr. C. L. Smith; University of Idaho, Moscow, (UID), Dr. W. F. Barr; University of Kansas, Lawrence, (KU), Dr. P. D. Ashlock; University of Kentucky, Lexington, (UKY), Dr. P. H. Freytag; University of Michigan, Ann Arbor, (UMI), Dr. T. E. Moore, Mr. M. F. O'Brien; University of Minnesota, St. Paul, (UNN), Dr. P. E. Clausen; University of Missouri, Columbia, (UMO), Mr. E. G. Riley; University of Nebraska, Lincoln, (UNL), Dr. B. C. Ratcliffe; University of Wisconsin, Madison, (UWI), Mr. S. Krauth; University of Wyoming, Laramie, (UWY), Dr. R. J. Lavigne; Virginia Polytechnic Institute and State University, Blacksburg, (VPI), Dr. M. Kosztarab; Washington State University, Pullman, (WSU), Dr. R. S. Zack. I also thank Dr. Kenneth V. Yeargan, University of Kentucky, Lexington, for providing live eggs and preserved nymphs of Podisus maculiventris. Dr. Donald B. Thomas, Jr. generously allowed me the use of his notes on Podisus, and offered useful suggestions. Dr. Joe E. Eger also gave advice and suggestions, and supplied slides of Apateticus. To my colleagues, Dr. Gary M. Stonedahi and Mr. Jerry Cassis I am grateful for ideas, many discussions, and help. Jerry also prepared the scanning electron micrograph of Anateticus bracteatus shown in Figure 16. To the members of my thesis committee, Drs. Arnold P. Appleby, Kenton L. Chambers, Paul L. Farber, Jeffrey C. Miller, and Paul W. Oman, I express ray sincere thanks. My major professor, Dr. John D. Lattin, suggested this project and allowed me full use of his excellent library. He also provided encouragement, advice, and friendship throughout this project's duration. To him I owe special gratitude. Special appreciation goes to Mr. Michael E. Szmania for encouragement and moral support over the last 3 years. To my parents, Richard B. and Frances K. Phillips, who throughout my life have taught me the value of knowledge and provided me overy opportunity to pursue it, I dedicate this thesis. TABLE OF CONTENTS INTRODUCTION LITERATURE REVIEW 6 MATERIALS AND METHODS 9 BIOLOGY AND ECONOMIC IMPORTANCE 12 DISTRIBUTION 17 METHOD OF CLASSIFICATION 20 DISCUSSION OF TAXONOMIC CHARACTERS 23 TAXONOMY 31 Genus Apateticus Dallas 34 Key to the Species of Apateticus 36 Subgenus Apateticus Dallas 38 Apateticus lineolatus (Herrich - Schaeffer) 38 Apateticus marainiventris (Stga) 43 Subgenus Apoecilus St&l. 46 Apateticus bracteatus (Fitch) 47 Apateticus anatarius Van Duzee 56 Apateticus cynicus (Say) 9 Genus Podisus Herrich - Schaeffer 67 Key to the Nearctic Species of Podisus 70 Subgenus Tylospilus St1.1 72 Podisus acutissimus Stcal 72 Subgenus Podisus Herrich - Schaeffer 82 Podisus mucronatus Uhler 83 Podisus maculiventri (Say) 87 Podisus serieventris Uhier 114 Podisus concfrex (St&l) 122 Podisus brevispinus, new species 125 Podisus fretus