A Revision of the Nearctic Species Of
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AN ABSTRACT OF THE THESIS OF John D. Oswald for the degree of Master of Science in Entomology presented on 11 April 1985. Title: A Revision of the Nearctic species of Sympherobius Banks (Neuroptera: Hemerobiidae). Abstract approved: Redacted for Privacy z / John D. Lattin The Nearctic species of the genus Sympherobius Banks are revised. Seventeen species are recorded from the region. Two new species, S. constrictus and S. quadricuspis are described from California and Arizona respectively. Four new specific synonyms are proposed (junior synonym(s) first): S. brunneus Nakahara and S. stangei Nakahara = S. angustus (Banks). S. gracilis Carpenter = S. umbratus (Banks), and S. texanus Nakahara = S. killingtoni Carpenter. Five species are newly recorded from Mexico. An annotated bibliography of relevant literature and key to the species is presented. A list of world Sympherobius species and phylogenetic notes on the Nearctic species are also included. A Revision of the Nearctic species of Sympherobius Banks (Neuroptera: Hemerobiidae) by John David Oswald A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Presented 11 April 1985 Commencement June 1985 APPROVED: Redacted for Privacy Professoz ¢f Entomology in charge of major Redacted for Privacy Head of department of Ento ogy Redacted for Privacy Dean of Gradua School Date thesis is presented 11 April 1985. ACKNOWLEDGEMENTS I gratefully acknowledge the following institutional collections and their curatorial staffs for loaning Sympherobius specimens for study; institutions loaning primary types are asterisked: (AMNH) American Museum of Natural History; Randall T. Schuh (ASUT) Arizona State University, Tempe; Frank F. Hasbrouck (BMNH*) British Museum (Natural History); P. C. Barnard (CAS*) California Academy of Sciences; Paul Arnaud (CMP) Carnegie Museum, Pittsburg; Chen W. Young (CSU) Colorado State University; W. D. Fronk (CU) Cornell University; James K. Liebherr (FEM) Frost Entomological Museum (Pennsylvania State University); K. C. Kim (INHS) Illinois Natural History Survey; Donald W. Webb (ISU) Iowa State University; Robert E. Lewis (LACM) Los Angeles County Museum; Charles L. Hogue (LSU) Louisiana State University; Cheryl B. Barr (MCZ*) Museum of Comparative Zoology (Harvard University); A. F. Newton (MSU) Mississippi State University; Patricia R. Miller (NCSR) North Carolina State University, Raleigh; Carol Parron (OKS) Oklahoma State University; W. A. Drew (OSU) Ohio State University; Charles A. Triplehorn (OSUC) Oregon State University, Corvallis; John D. Lattin (PMY) Peabody Musuem (Yale University); Charles Remington (SMEK*) Snow Museum of Entomology (University of Kansas); George Byers (TAMU) Texas A & M University; H. R. Burke (UAF) University of Arkansas, Fayetteville; Chris Carlton (UAT) University of Arizona, Tucson; Floyd G. Werner (UCB) University of California, Berkeley; J. A. Powell (UCD) University of California, Davis; Robert 0. Schuster (UCR) University of California, Riverside; Saul Frommer (UGA) University of Georgia, Athens: Cecil L. Smith (UIM) University of Idaho, Moscow; Frank Merickel (UMAA) University of Michigan, Ann Arbor; Barry M. O'Connor (UMC) University of Missouri, Columbia; R. L. Blinn (UMSP) University of Minnesota, St. Paul; Philip J. Clausen (UNH) University of New Hampshire; Donald S. Chandler (USNM*) United States National Museum; Oliver S. Flint, Jr. (UWL) University of Wyoming, Laramie; Robert Lavigne (UWM) University of Wisconsin, Madison; Steven Krauth (VPI) Virginia Polytechnic Institute; Michael Kosztarab (WSU) Washington State University; Richard S. Zack Acronyms used for institutional collections are those of Heppner and Lamas (1982), except Colorado State University (CSU), which they do not list. I especially thank Dr. John D. Lattin for his professional guidance and friendship during my graduate studies at Oregon StateUniversity. I thank also the members of my graduate committee, Drs. Kenton L. Chambers, Jefferson J. Gonor and Paul Oman. Fellow graduate studens Gary M. Stonedahl and Kathryn A. Phillips tutored me in museum techniques and offered encouragement and assistance during my first year of study. Financial support for this study was provided through a research assistantship in the Oregon State University Systematic Entomology Laboratory. Computer services were made possible through a grant for unsponsored graduate research by the Oregon State University Computer Center. TABLE OF CONTENTS page I. INTRODUCTION 1 II. TERMS AND ABBREVIATIONS 3 III. MATERIALS AND METHODS 4 IV. BIOLOGY 6 V. TAXONOMY OF THE NEARCTIC SPECIES OF SYMPHEROBIUS BANKS GENUS SYMPHEROBIUS BANKS 8 KEY TO THE NEARCTIC SPECIES OF SYMPHEROBIUS . 20 Sympherobiusamiculus (Fitch, 1856) . 29 Sympherobiusangustus (Banks, 1904) . 32 Sympherobiusarizonicus Banks, 1911 . 38 Sympherobiusbarberi (Banks, 1903) . 40 Sympherobiusbeameri Gurney, 1948 . 43 Sympherobiusbifasciatus Banks, 1911 . 45 Sympherobiuscalifornicus Banks, 1911 . 47 Sympherobiusconstrictus new species . 50 Sympherobiusdistinctus Carpenter, 1940 . 52 Sympherobiuskillingtoni Carpenter, 1940 . 54 Sympherobiuslimbus Carpenter, 1940 . 57 Sympherobiusoccidentalis (Fitch, 1856) . 59 Sympherobiusperparvus (McLachlan, 1869) . 62 Sympherobiuspictus (Banks, 1904) . 65 Sympherobiusquadricuspis new species . 68 Sympherobiussimilis Carpenter, 1940 . 69 Sympherobiusumbratus (Banks, 1903) . 71 VI. PHYLOGENETIC CONSIDERATIONS 75 VII. SYNOPSIS OF THE NEARCTIC SPECIES OF SYMPHEROBIUS 84 LITERATURE CITED 85 APPENDIX A. FIGURES 1 - 106 94 APPENDIX B. LIST OF THE WORLD SPECIES OF SYMPHEROBIUS BANKS 109 INDEX TO THE SPECIES OF SYMPHEROBIUS 115 A REVISION OF THE NEARCTIC SPECIES OF SYMPHEROBIUS BANKS (NEUROPTERA: HEMEROBIIDAE) I. INTRODUCTION The brown lacewings (Hemerobiidae) are a cosmopolitan family of the order Neuroptera, Planipennia. The generic classification of this family has not yet been stabilized. Authors who apply conservative generic concepts recognize approximately 45 recent genera while authors adhering to more restrictive generic concepts recognize about 60 genera. The family contains approximately 550 recognized species. The genus Sympherobius Banks contains 46 species distributed throughout the Nearctic (17 species), Neotropical (15), Palearctic (14) and Ethiopian (1) faunal regions (see appendix B). Revisional treatments of regional hemerobiid faunas which contain Sympherobius species may be found in Carpenter (1940) [North America], Aspbck et al.(1980) and Killington (1936, 1937) [Europe], Kozhantshikov (1956) [Asia], Kuwayama (1962)[Japan] and Tjeder (1961) [SouthernAfrica]. The species of Central and South America have been listed by Penny (1977), but the taxonomy of these species is in need of review. The generic epithet Symphercbius (from the Greek "sympheron", useful) originally appeared without 2 description in a list of neuroptercid insects published by Nathan Banks (1904). Banks (1906) included seven North American species in the genus and provided the first generic description. Subsequently, fifteen additional species have been described from the Nearctic region by Banks (1911), Nav&s (1912a, 1914a), Carpenter (1940), Gurney (1948) and Nakahara (1960, 1965a). Three of these species were placed in synonymy by Carpenter (1940). The present revision recognizes 17 valid Nearctic species, describes two new species and places four additional names in synonymy. 3 II. TERMS AND ABBREVIATIONS Morphological terminology follows Tjeder (1954, 1961, 1970) or Morse (1931). Illustrations of gross planipennian morphology may he found in these works. Where this terminology has proven inadequate in describing details of male genitalic structures, new descriptive terms have been introduced. States and provinces are abbreviated with standard two-letter United States Postal Service abbreviations. Annotations used in synonymical listings are as follows: A = Adult description or characterization Bic = Biocontrol Bio = Biology C = Cocoon Dst = Distribution E = Egg F = Female description or oharacterization ID = Incorrect determination Key = Key, keyed L = Larvae (number indicates instar) Lst = List, listed MG = Male genitalia Mor = Morphology (other than taxonomic characters) OD = Original description P = Pupa Par = Parasites PP = Prepupa RD = Redescription Tax = Taxonomy (including synonymy, homonomy, type data etc.) W = Wing * = indicates a figure of the item it follows 4 III. MATERIALS AND METHODS More than 2100 specimens of Nearctic Sympherobius were examined during the course of the present study. Species decisions are based on structures of the male genitalia. For dissection and examination, male genitalia were cleared for 5 to 10 minutes in a subboiling solution of 10 percent NaOH, neutralized in dilute acetic acid, rinsed in water, dried on a paper blotter and transfered to glycerin. When dissected male genitalia are temporarily mounted in glycerin on depression slides, the fine structural details are easily observed with a compound microscope. Following examination, genitalia were stored in glycerin in microvials pinned below the specimen. Forewings were mounted on slides in Hoyer's solution and photographed using Kodak Panatomic X film. Forewing lengths were measured from the anterior margin of the tegula to the wing apex. All drawings and measurements were made using a calibrated ocular grid reticle. Figures of the following structures are given for each species:(1) male ninth tergite and ectoproct (lateral view),(2) gonarcus