AN ABSTRACT OF THE THESIS OF Gary Michael Stonedahl for the degree of Doctor of Philosophy in Entomology presented on December 9, 1982 Title: A systematic study of the genus Phytocoris Fallen (Heteroptera: Miridae) in western North America Redacted for Privacy Abstract approved: John D. Lattin The western Nearctic species of the genus Phytocoris Fallen were taxonomically revised. Over 15,000 specimens were examined. Information concerning host plant associations and distributions was compiled and summarized for use by future researchers. External morphological features and characters of the male genitalia were used to distinguish species. The genital tubercles above the clasper bases, claspers, and sclerotized process of the vesica were especially useful in this effort. These structures have been illustrated for each of the included species. One-hundred and sixty-two species of Phytocoris are now recognized from western North America. These have been arranged into 20 species-groups ranging in size from two to twenty-two species. The formation of species-groups was based primarily on characters of the male genitalia. Relationships between groups were cladistically analyzed. Forty-four characters of the male genitalia and external morphology were used to determine the final arrangement of species-groups. Forty-six new species are described, and the following synonymies are proposed (senior synonym first): acaciae Knight = minuendus Knight; calli Knight = laticeps Knight; deserticola Knight lineatellus Knight; empirensis Knight m rinconae Knight; fraterculus Van Duzee = westwoodi Bliven; fuscipennis Knight = longirostris Knight; geniculatus Van Duzee = blackwelli Bliven; interspersus Uhler = eurekae Bliven; juniperanus Knight = chiricahuae Knight, flaviatus Knight, santaritae Knight; ketinelbi Bliven = kahtahbi Bliven; knowltoni Knight = albertae Knight, elongatus Knight; maritimus Van Duzee = sequoiae Bliven; neglectus Knight = yuroki Bliven; nigrifrons Van Duzee = abiesi. Knight, tehamae Bliven; palmeri Reuter = hesperellus Knight, hesperius Knight, vittatus Reuter; politus Reuter = rusticus Van Duzee; relativus Knight = albiscutellatus Knight; stellatus Van Duzee = arcatae Bliven, tinctus Knight; stitti Knight = albiceps Knight, merinoi Knight; strigosus Knight = flavellus Knight; sublineatus Knight = subcinctus Knight, tanneri Knight; vanduzeei Reuter = nigripubescens Knight; ventralis Van Duzee = contrastus Knight, ephedrae Knight, quadricinctus Knight; vinaceus Van Duzee = hyampon Bliven; yollabollae Bliven = albiclavus Knight, montanae Knight, taos Knight. The majority of western Nearctic Phytocoris species inhabit trees and shrubs; a few breed on grasses and herbaceous plants. Diversity is greatest in shrub and steppe communities of the southwestern United States. Most species are associated with one host plant or several related plant species; many appear to be inhabitants of bark. Most Phytocoris species are believed to be predaceous both as nymphs and adults, feeding on small, soft-bodied insects and mites. Several Nearctic species have been identified as predators of forest and orchard pests. A systematic study of the genus PhytocorisFall4n (Heteroptera: Miridae) in western North America by Gary Michael Stonedahl A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Commencement June 1983 APPROVED: Redacted forPrivacy Professor of eomology in charge of major ,Redacted for Privacy Head of Department of Entomology45/ Redacted for Privacy Dean of Gradua_ Date thesis is presented December 9, 1982 Typed by Julie A. Stonedahl for GARY MICHAEL STONEDAHL ACKNOWLEDGEMENTS I extend my thanks collectively to all persons involved with various aspects of this study; without their contributions much of the research could not have been completed. I express my warmest appreciation to Dr. John D. Lattin, my major professor, who suggested the study and provided guidance for its duration. His professional advice, constant encouragement and friendship are sincerely appreciated. I thank the members of my committee, Drs. Kenton L. Chambers, Robert E. Frenkel, James D. Hall, and William P. Stephen, for their comments on the thesis. Special thanks is given to Dr. Paul W. Oman for his valuable advice during the final stages of preparation of the thesis, and for his friendship during the course of this study. His ideas and comments concerning many aspects of my research and systematics in general were especially helpful. I am grateful to Mrs. Bonnie B. Hall for her comments regarding illustration techniques; to Dr. Ernst J. Dornfeld, Zoology Department, Oregon State University, for spending many hours with me translating German literature; and to Mrs. LaRea D. Johnston, Herbarium, Oregon State University, for providing plant identifications. Special thanks are extended to Mr. Thomas J. Henry, National Museum of Natural History, Washington, D.C. and to Dr. Randall T. Schuh, American Museum of Natural History, New York, for graciously hosting me in their homes during visits to their respective museums. I am grateful for their advice and encouragement during the course of this study, and for their assistance in collecting numerous specimens on field trips in the western United States. I especially thank the following individuals: Dr. Vincent P. Razafimahatratra for instructing me in the art of mirid genitalic dissection; Mr. Michael D. Schwartz for acquainting me with the operation of the Wagner 78 cladistics program, and for his advice and sincere encouragement during the course of this study; Ms. Kathy Phillips, Mr. Gerasimos Cassis, and Mr. Gary M. Cooper for their valuable advice and moral support; and my parents, who have provided wisdom and continuous encouragement throughout my academic career. Finally, to my wife Julie, I am indebted for recording and organizing the locality data and for her patience and constant encouragement throughout my graduate studies. I also thank Julie for her superb typing of the entire manuscript. Type specimens were borrowed from the California Academy of Sciences, San Francisco (CAS), Dr. Paul H. Arnaud, Jr.; the Swedish Museum of Natural History, Stockholm, Sweden (SMNH), Dr. Per Lindskog; the National Museum of Natural History, Washington, D.C. (USNM), Mr. Thomas J. Henry and Dr. Richard C. Froeschner; and the Zoological Museum of the University, Helsinki, Finland (ZMH), Dr. Antti Jansson. In addition to the above, the following institutions or private collections kindly loaned specimens for study: American Museum of Natural History, New York (AMNH), Dr. Randall T. Schuh; Arizona State University, Tempe (ASU), Dr. Frank F. Hasbrouck; Brigham Young University, Provo, Utah (BYU), Dr. Stephen L. Wood; California State Department of Food and Agriculture, Sacramento (CAF&A), Dr. Alan R. Hardy; Canadian National Collection, Ottawa (CNC), Dr. Leonard A. Kelton; Colorado State University, Fort Collins (CSU), Dr. Howard E. Evans; James Entomological Collection, Pullman, Washington (WSU), Dr. William J. Turner; Los Angeles County Museum of Natural History, California (LACM), Dr. Charles L. Hogue; Montana State University, Bozeman (MSU), Ms. Sharon D. Rose; Northern Arizona University, Flagstaff (NAU), Dr. Clarence D. Johnson; New Mexico State University, Las Cruces (NMS), Dr. James R. Zimmerman; Oregon State Department of Agriculture, Salem (OSDA), Mr. Kenneth Goeden; Oregon State University, Corvallis (OSU), Dr. John D. Lattin, Dr. Andrew P. Moldenke, Dr. Paul W. Oman, Dr. Loren K. Russell, Mrs. Becky L. Fichter, Mr. Gary M. Cooper, Mr. Michael D. Schwartz; Polhemus Collection, Englewood, Colorado (JTP), Dr. John T. Polhemus; San Diego Natural History Museum, California (SDNH), Mr. David K. Faulkner; Snow Entomological Collection, Lawrence, Kansas (KU), Dr. Peter D. Ashlock and Dr. George W. Byers; Texas A & M University, College Station (T.A&M and SHF), Dr. Joseph C. Schaffner; University of Arizona, Tucson (UAZ), Dr. Floyd G. Werner; University of British Columbia, Vancouver (UBC), Dr. Geoffrey G.E. Scudder; University of California, Berkeley (UCB), Dr. Jerry A. Powell; University of California, Davis (UCD), Dr. Robert 0. Schuster; University of California, Riverside (UCR), Dr. John D. Pinto and Mr. Saul I. Frommer; University of Idaho, Moscow (UID), Dr. William F. Barr; Utah State University, Logan (USU), Dr. Wilford J. Hanson; Western Washington University, Bellingham (WWU), Dr. Gerald F. Kraft. Financial support was provided through a research assistantship in the Oregon State University Systematic Entomology Laboratory with funds made available by the Science Technical Advisory Fund. The Oregon State University Computer Center granted funds for computer services in unsponsored graduate student research. Funding for visits to the United States National Museum of Natural History and the American Museum of Natural History was provided by a grant from the Theodore Roosevelt Memorial Fund, American Museum of Natural History, New York. Additional support for museum visits and field work was provided by the Oregon State University Foundation (Systematic Entomology Laboratory Fund). TABLE OF CONTENTS Page INTRODUCTION 1 LITERATURE REVIEW 4 METHODS AND MATERIALS 8 Materials and general procedures 8 Methods of classification 10 BIOLOGY 14 DISCUSSION OF TAXONOMIC CHARACTERS 22 CLASSIFICATION 30 DISTRIBUTION 38 TAXONOMY 43 Description of genus Phytocoris Fallen 43 Key to species-group 46 Cunealis Species-Group 62 Phytocoris rufoscriptus Van