american beetles, volume ii: polyphaga: scarabaeoidea through curculionoidea

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  • AMERICANBEETLES

    VOLUME 2

    Polyphaga:Scarabaeoidea through Curculionoidea

  • Boca Raton London New York Washington, D.C.CRC Press

    AMERICANBEETLES

    VOLUME 2

    Polyphaga:Scarabaeoidea through Curculionoidea

    Edited bythe late Ross H. Arnett, Jr., Ph.D.

    Michael C. Thomas, Ph.D.Paul E. Skelley, Ph.D.

    andJ. Howard Frank, D. Phil.

  • COVER FIGURES: Center - Coccinellidae,

    Harmonia axyridus

    (Palles) [Photo by Fred J. Santana]. Outer rim, clockwise from top:Ripiphoridae,

    Macrosiagon cruentum

    (Germar) [by Fred J. Santana]; Meloidae,

    Lytta magister

    Horn [by Charles L. Bellamy];Carabidae,

    Rhadine exilis

    (Barr and Lawrence) [by James C. Cokendolpher]; Melyridae,

    Malachius mirandus

    (LeConte) [by MaxE. Badgley]; Lampyridae,

    Microphotus angustus

    LeConte [by Arthur V. Evans].

    This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, andsources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information,but the authors and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use.

    Neither this book nor any part may be reproduced or transmitted in any form or by any means, electronic or mechanical, includingphotocopying, microfilming, and recording, or by any information storage or retrieval system, without prior permission in writing from thepublisher.

    The consent of CRC Press LLC does not extend to copying for general distribution, for promotion, for creating new works, or for resale.Specific permission must be obtained in writing from CRC Press LLC for such copying.

    Direct all inquiries to CRC Press LLC, 2000 N.W. Corporate Blvd., Boca Raton, Florida 33431.

    Trademark Notice:

    Product or corporate names may be trademarks or registered trademarks, and are used only for identification andexplanation, without intent to infringe.

    Visit the CRC Press Web site at www.crcpress.com

    © 2002 by CRC Press LLC

    No claim to original U.S. Government worksInternational Standard Book Number 0-8493-0954-9

    Library of Congress Card Number 00-050809Printed in the United States of America 1 2 3 4 5 6 7 8 9 0

    Printed on acid-free paper

    Library of Congress Cataloging-in-Publication Data

    American beetles / edited by Ross H. Arnett and Michael C. Thomas.p. cm.

    Contents: v. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia.Includes bibliographical references (p.).ISBN 0-8493-0954-9 (alk. paper : v. 2))1. Beetles—North America. I. Arnett, Ross H. II. Thomas, M. C. (Michael Charles).

    1948–QL581 .A43 2002595.76

    ¢

    097—dc21 00-050809 CIP

  • iv

    To Ross H. Arnett, Jr.1919-1999

    and

    Mary Arnett1919-2002

  • v

  • vii

    PrefaceIt has been nearly 40 years since Ross H. Arnett, Jr. published the first fascicle of The Beetles of the United States: A Manual for

    Identification. It quickly became an indispensable tool for professional and amateur coleopterists, general entomologists, and natural-ists. Although there were four additional printings it has long been out of print and difficult to obtain. It was prepared to replaceBradleys A Manual of the Genera of Beetles of America, North of Mexico, which itself was some 30 years out of date in 1960. AmericanBeetles is, in turn, designed to replace The Beetles of the United States. It is hoped that it will prove to be as useful as its predecessor.

    Ironically, much of the preface to the original edition applies today as well as it did 40 years ago:

    Many genera have since been described and reported within the area concerned, and many families have beenrevised. Extensive changes have been made in the family classification of the beetles of the United States during thisperiod.

    The aim of this series of fascicles is to provide a tool for the identification of adult beetles of the United Statesto family and genus with the aid of illustrations, keys, descriptions, and references to sources for keys and descrip-tions of the species of this area. All of the genera known to inhabit this area are included in the keys and lists ofgenera which follow.

    The design and format of this work follow closely that of the original edition, but the way it was put together was quite different.Its predecessor was very much the work of one man, Ross H. Arnett, Jr. With a few exceptions (George Ball wrote the carabid treatmentfor both the 1960 edition and for this one), Dr. Arnett wrote the family treatments of The Beetles of the United States. Many specialistsreviewed those chapters, but they were almost entirely Dr. Arnetts work.

    When Dr. Arnett announced plans to prepare a work to replace The Beetles of the United States, coleopterists literally lined up tovolunteer their time and expertise in preparing the family treatments. Ultimately, more than 60 coleopterists participated in thepreparation of American Beetles. This has truly been a community project.

    Due to the size of the ensuing work, American Beetles is being printed in two volumes. Volume 1 includes the introductorymaterial, and family treatments for the Archostemata, Myxophaga, Adephaga, and Polyphaga: Staphyliniformia. The remainder of thePolyphaga and the keys to families appear here in Volume 2.

    Sadly, although Dr. Arnett initiated this project and was instrumental in its planning, he did not live to see its fruition. He becameseriously ill in late 1998 and died on July 16, 1999 at the age of 80. We hope he would be pleased with the outcome.

    Michael C. Thomas, Ph.D.Gainesville, Florida

    April 3, 2002

  • viii

    Table of Contents

    Preface ................................................................................................................................................................................................................... vii

    Acknowledgments ............................................................................................................................................................................................... xi

    Contributors to Volume 2 ................................................................................................................................................................................. xii

    Series SCARABAEIFORMIA ............................................................................................................................................................................. 1Superfamily SCARABAEOIDEA.............................................................................................................................................................. 1

    Family 23. LUCANIDAE .................................................................................................................................................................... 6Family 24. DIPHYLLOSTOMATIDAE ........................................................................................................................................ 10Family 25. PASSALIDAE .................................................................................................................................................................. 12Family 26. GLARESIDAE ................................................................................................................................................................ 15Family 27. TROGIDAE .................................................................................................................................................................... 17Family 28. PLEOCOMIDAE............................................................................................................................................................ 20Family 29. GEOTRUPIDAE ............................................................................................................................................................ 23Family 30. OCHODAEIDAE........................................................................................................................................................... 28Family 31. HYBOSORIDAE ............................................................................................................................................................ 32Family 32. CERATOCANTHIDAE ............................................................................................................................................... 34Family 33. GLAPHYRIDAE ............................................................................................................................................................ 37Family 34. SCARABAEIDAE .......................................................................................................................................................... 39

    Series ELATERIFORMIA ................................................................................................................................................................................ 82Superfamily SCIRTOIDEA ...................................................................................................................................................................... 82

    Family 35. EUCINETIDAE ............................................................................................................................................................. 82Family 36. CLAMBIDAE .................................................................................................................................................................. 85Family 37. SCIRTIDAE ..................................................................................................................................................................... 87

    Superfamily DASCILLOIDEA ................................................................................................................................................................. 90Family 38. DASCILLIDAE ............................................................................................................................................................... 90Family 39. RHIPICERIDAE ............................................................................................................................................................ 92

    Superfamily BUPRESTOIDEA ................................................................................................................................................................ 95Family 40. SCHIZOPODIDAE ....................................................................................................................................................... 95Family 41. BUPRESTIDAE .............................................................................................................................................................. 98

    Superfamily BYRRHOIDEA .................................................................................................................................................................. 113Family 42 BYRRHIDAE ................................................................................................................................................................. 113Family 43. ELMIDAE ..................................................................................................................................................................... 117Family 44. DRYOPIDAE ................................................................................................................................................................ 121Family 45 LUTROCHIDAE ........................................................................................................................................................... 123Family 46. LIMNICHIDAE............................................................................................................................................................ 125Family 47. HETEROCERIDAE .................................................................................................................................................... 127Family 48. PSEPHENIDAE ........................................................................................................................................................... 133Family 49. PTILODACTYLIDAE ................................................................................................................................................. 135Family 50. CHELONARIIDAE ..................................................................................................................................................... 139Family 51. EULICHADIDAE ........................................................................................................................................................ 142Family 52. CALLIRHIPIDAE ........................................................................................................................................................ 144

    Superfamily ELATEROIDEA ............................................................................................................................................................... 146Family 53. ARTEMATOPODIDAE............................................................................................................................................. 146Family 54. BRACHYPSECTRIDAE ............................................................................................................................................. 148Family 55. CEROPHYTIDAE ....................................................................................................................................................... 150Family 56. EUCNEMIDAE............................................................................................................................................................ 152Family 57. THROSCIDAE.............................................................................................................................................................. 158Family 58. ELATERIDAE .............................................................................................................................................................. 160Family 59. LYCIDAE ...................................................................................................................................................................... 174Family 60. TELEGEUSIDAE ........................................................................................................................................................ 179

  • ix

    Family 61. PHENGODIDAE ........................................................................................................................................................ 181Family 62. LAMPYRIDAE ............................................................................................................................................................. 187Family 63. OMETHIDAE .............................................................................................................................................................. 197Family 64. CANTHARIDAE.......................................................................................................................................................... 202

    Series BOSTRICHIFORMIA .......................................................................................................................................................................... 219Family 65. JACOBSONIIDAE ...................................................................................................................................................... 219

    Superfamily DERODONTOIDEA ...................................................................................................................................................... 221Family 66. DERODONTIDAE ..................................................................................................................................................... 221

    Superfamily BOSTRICHOIDEA........................................................................................................................................................... 224Family 67. NOSODENDRIDAE .................................................................................................................................................. 224Family 68. DERMESTIDAE .......................................................................................................................................................... 228Family 69. BOSTRICHIDAE ......................................................................................................................................................... 233Family 70. ANOBIIDAE ................................................................................................................................................................ 245

    Series CUCUJIFORMIA .................................................................................................................................................................................. 261Superfamily LYMEXYLOIDEA............................................................................................................................................................ 261

    Family 71. LYMEXYLIDAE .......................................................................................................................................................... 261Superfamily CLEROIDEA ..................................................................................................................................................................... 263

    Family 72. TROGOSSITIDAE ...................................................................................................................................................... 263Family 73. CLERIDAE .................................................................................................................................................................... 267Family 74. MELYRIDAE ................................................................................................................................................................ 281

    Superfamily CUCUJOIDEA ................................................................................................................................................................... 305Family 75. SPHINDIDAE .............................................................................................................................................................. 305Family 76. BRACHYPTERIDAE .................................................................................................................................................. 309Family 77. NITIDULIDAE ............................................................................................................................................................ 311Family 78. SMICRIPIDAE.............................................................................................................................................................. 316Family 79. MONOTOMIDAE ....................................................................................................................................................... 319Family 80. SILVANIDAE ............................................................................................................................................................... 322Family 81. PASSANDRIDAE ......................................................................................................................................................... 327Family 82. CUCUJIDAE ................................................................................................................................................................. 329Family 83. LAEMOPHLOEIDAE ................................................................................................................................................ 331Family 84. PHALACRIDAE ........................................................................................................................................................... 335Family 85. CRYPTOPHAGIDAE ................................................................................................................................................. 338Family 86. LANGURIIDAE ........................................................................................................................................................... 343Family 87. EROTYLIDAE ............................................................................................................................................................. 348Family 88. BYTURIDAE ................................................................................................................................................................ 354Family 89. BIPHYLLIDAE ............................................................................................................................................................. 356Family 90. BOTHRIDERIDAE ..................................................................................................................................................... 358Family 91. CERYLONIDAE .......................................................................................................................................................... 363Family 92. ENDOMYCHIDAE .................................................................................................................................................... 366Family 93. COCCINELLIDAE ...................................................................................................................................................... 371Family 94. CORYLOPHIDAE ....................................................................................................................................................... 390Family 95. LATRIDIIDAE ............................................................................................................................................................. 395

    Superfamily TENEBRIONOIDEA ...................................................................................................................................................... 399Family 96. MYCETOPHAGIDAE ............................................................................................................................................... 399Family 97. ARCHEOCRYPTICIDAE .......................................................................................................................................... 401Family 98. CIIDAE .......................................................................................................................................................................... 403Family 99. TETRATOMIDAE ...................................................................................................................................................... 413Family 100. MELANDRYIDAE .................................................................................................................................................... 417Family 101. MORDELLIDAE ....................................................................................................................................................... 423Family 102. RIPIPHORIDAE ........................................................................................................................................................ 431Family 103. COLYDIIDAE ............................................................................................................................................................ 445Family 104. MONOMMATIDAE ................................................................................................................................................. 454Family 105. ZOPHERIDAE........................................................................................................................................................... 457Family 106. TENEBRIONIDAE .................................................................................................................................................. 463Family 107. PROSTOMIDAE ........................................................................................................................................................ 510

  • x

    Family 108. SYNCHROIDAE ........................................................................................................................................................ 512Family 109. OEDEMERIDAE ...................................................................................................................................................... 514Family 110. STENOTRACHELIDAE.......................................................................................................................................... 520Family 111. MELOIDAE ................................................................................................................................................................ 522Family 112. MYCTERIDAE........................................................................................................................................................... 530Family 113. BORIDAE .................................................................................................................................................................... 534Family 114. PYTHIDAE ................................................................................................................................................................. 537Family 115. PYROCHROIDAE ..................................................................................................................................................... 540Family 116. SALPINGIDAE .......................................................................................................................................................... 544Family 117. ANTHICIDAE............................................................................................................................................................ 549Family 118. ADERIDAE ................................................................................................................................................................ 559Family 119. SCRAPTIIDAE ........................................................................................................................................................... 564

    Superfamily CHRYSOMELOIDEA ...................................................................................................................................................... 568Family 120. CERAMBYCIDAE ..................................................................................................................................................... 568Family 121. BRUCHIDAE .............................................................................................................................................................. 602Family 122. MEGALOPODIDAE ................................................................................................................................................ 609Family 123. ORSODACNIDAE ..................................................................................................................................................... 613Family 124. CHRYSOMELIDAE .................................................................................................................................................. 617

    Superfamily CURCULIONOIDEA ....................................................................................................................................................... 692Family 125. NEMONYCHIDAE .................................................................................................................................................. 692Family 126. ANTHRIBIDAE ......................................................................................................................................................... 695Family 127. BELIDAE .................................................................................................................................................................... 701Family 128. ATTELABIDAE ......................................................................................................................................................... 703Family 129. BRENTIDAE .............................................................................................................................................................. 711Family 130. ITHYCERIDAE ......................................................................................................................................................... 720Family 131. CURCULIONIDAE ................................................................................................................................................... 722

    Family Key ......................................................................................................................................................................................................... 816Taxonomic Index .............................................................................................................................................................................................. 836

  • xi

    Acknowledgments for Volume II

    Originally, Ross Arnett was to have authored many of the family treatments, especially for those families with no specialistsavailable. His death in 1999 left many families without an author. Several volunteers stepped forward, but Dan Young of the Universityof Wisconsin took responsibility for more than his fair share and got several of his enthusiastic graduate students involved in theproject also. The members of the Editorial Board, listed in the Introduction, provided guidance, advice, and constructive criticism, butJ. Howard Frank of the University of Florida has been outstanding in his unwavering demands for scholarship and proper English,and joined Paul E. Skelley and Michael C. Thomas, both of the Florida Department of Agriculture and Consumer Services, as an editorof Volume II. John Sulzycki of CRC Press has been more than helpful throughout some trying times.

    Many of the excellent habitus drawings beginning the family treatments were done by Eileen R. Van Tassell of the University ofMichigan for The Beetles of the United States, and for Volume 2 of American Beetles she produced excellent new ones for families 100, 108,and 119.

    Authors of the family treatments often have acknowledgments in their respective chapters throughout the body of the text.Ross Arnett's widow, Mary, was always his support staff throughout his long and productive career. After Ross' death, she helped

    by providing free and gracious access to Ross' files, and by her steady encouragement and quiet conviction that we would indeed be ableto finish this, Ross Arnett's last big project. Unfortunately, Mary Arnett did not live to see Volume II published. She became ill in thefall of 2001 and died on January 3, 2002.

    And I would like to again acknowledge my wife, Sheila, for her patience and forbearance during the long and sometimes difficultpath that led to this volume.

    Michael C. Thomas, Ph.D.Gainesville, Florida

    April 3, 2002

  • xii

    Authors

    Rolf L. Aalbu, Ph.D.Department of EntomologyCalifornia Academy of SciencesGolden Gate ParkSan Francisco, CA 94118-4599106. Tenebrionidae.

    Robert S. Anderson, Ph.D.Canadian Museum of Nature/Entomol-

    ogyP.O. Box 3443, Station DOttawa, ON K1P 6P4 CANADA125. Nemonychidae; 127. Belidae; 129.Brentidae; 130. Ithyceridae; 131. Curculionidae.

    Fred G. Andrews, Ph.D.California Department of Food and Ag-

    riculturePlant Pest Diagnostics Laboratory3294 Meadowview Rd.Sacramento, CA 95832-144895. Latridiidae.

    Ross H. Arnett, Jr., Ph.D.Senior Editor

    C. L. Bellamy, D.Sc.EntomologyNatural History Museum900 Exposition BlvdLos Angeles CA 90007, U.S.A.40. Schizopodidae; 41. Buprestidae.

    Marco A. Bologna, Ph.D.Dipartimento di BiologiaUniversità degli studi ROMA TREViale G. Marconi, 446I-00146 ROMA111. Meloidae.

    Yves Bousquet, Ph.D.Eastern Cereal and Oilseed Research CentreAgriculture and Agri-Food CanadaOttawa, ON K1A 0C6, CANADA79. Monotomidae.

    Stanley Bowestead, Ph.D.Department of EntomologyThe Manchester Museum, The UniversityManchester M12 9PL England94. Corylophidae.

    Kirby W. Brown, Ph.D.P.O. Box 1838Paradise, CA 95967106. Tenebrionidae.

    J. Milton Campbell, Ph.D.420 Everetts LaneHopkinsville, KY 42240106. Tenebrionidae.

    David C. Carlson, Ph.D.4828 Dauntless WayFair Oaks, CA 9562830. Ochodaeidae; 33. Glaphyridae.

    Donald S. Chandler, Ph.D.Department of ZoologyUniversity of New HampshireDurham, NH 03824117. Anthicidae; 118. Aderidae.

    Shawn M. Clark, Ph.D.Monte L. Bean MuseumBrigham Young UniversityProvo, UT 84602122. Megalopodidae; 123. Orsodacnidae; 124.Chrysomelidae.

    Arthur V. Evans, Ph.D.Department of EntomologyNational Museum of Natural HistorySmithsonian Institutionc/o 1600 Nottoway Ave.Richmond, VA 2322734. Scarabaeidae: Melolonthinae.

    Zachary H. FalinDivision of EntomologyNatural History Museum and Biodiversity

    Research CenterSnow Hall, 1460 Jayhawk Blvd.Lawrence, KS 66045102. Ripiphoridae

    R. Wills Flowers, Ph.D.Center for Biological ControlFlorida A & M UniversityTallahassee, FL 32307124. Chrysomelidae.

    J. Joseph GierschDepartment of EntomologyMontana State UniversityBozeman, MT 5971765. Jacobsoniidae

    Arthur J. GilbertCalifornia Department of Food and

    Agriculture2889 N. Larkin St., Suite 106Fresno, CA 93727124. Chrysomelidae.

    Bruce D. Gill, Ph.D.4032 Stonecrest RoadWoodlawn, ON K0A 3M0, CANADA34. Scarabaeidae: Scarabaeinae.

    Michael A. Goodrich, Ph.D.Department of Biological SciencesEastern Illinois UniversityCharleston, IL 6192088. Byturidae; 89. Biphyllidae

    Robert D. Gordon, Ph.D.Northern Plains EntomologyP.O.Box 65Willow City, ND 5838434. Scarabaeidae: Aphodiinae.

    Dale H. Habeck, Ph.D.Entomology and Nematology Depart-

    mentUniversity of FloridaGainesville, FL 3261176. Brachypteridae; 77. Nitidulidae.

    Robert W. Hamilton, Ph.D.Department of BiologyLoyola University Chicago6525 North Sheridan RoadChicago, IL 60626128. Attelabidae.

    Henry A. Hespenheide, Ph.D.Department of Organismic BiologyUniversity of CaliforniaLos Angeles, CA 90095-1606131. Curculionidae: Conoderinae.

    Frank T. Hovore, Ph.D.14734 Sundance PlaceSanta Clarita, CA 91387-154228. Pleocomidae.

    Anne T. Howden, M.Sc.Canadian Museum of NatureP.O. Box 3443, Station DOttawa, ON K1P 6P4 Canada131. Curculionidae: Entiminae.

    Contributors to Volume 2 of American Beetles

  • xiii

    Michael A. Ivie, Ph.D.Department of EntomologyMontana State UniversityBozeman, MT 5971749. Ptilodactylidae; 50. Chelonariidae; 65.Jacobsoniidae; 67. Nosodendridae; 69.Bostrichidae; 90. Bothrideridae; 103. Colydiidae;104. Monommatidae; 105. Zopheridae; FamilyKey.

    John A. Jackman, Ph.D.Department of Entomology412 Heep CenterTexas A & M UniversityCollege Station, TX 77843101. Mordellidae.

    Mary L. Jameson, Ph.D.W436 Nebraska HallSystematics Research CollectionsUniversity of Nebraska State MuseumLincoln, NE 68588-0514Scarabaeoidea; 24. Diphyllostomatidae; 26.Glaresidae; 27. Trogidae; 29. Geotrupidae; 31.Hybosoridae; 32. Ceratocanthidae; 34.Scarabaeidae.

    Paul J. Johnson, Ph.D.Insect Research Collection, Box 2207-ASouth Dakota State UniversityBrookings, SD 5700742. Byrrhidae; 58. Elateridae; 55. Cerophytidae;57. Throscidae.

    Kerry KatovichDepartment of EntomologyUniversity of Wisconsin-MadisonMadison, WI 5370639. Rhipiceridae; 47. Heteroceridae.

    John M. Kingsolver, Ph.D.Florida State Collection of ArthropodsP. O. Box 147100Gainesville, FL 3261468. Dermestidae; 121. Bruchidae.

    David G. Kissinger, Ph.D.24414 University Avenue #40Loma Linda, CA 92354129. Brentidae: Apioninae.

    Boris A. Korotyaev, Ph.D.Zoological InstituteRussian Academy of SciencesUniversitetskaya nab 1.St. Petersburg, 199034, Russia131. Curculionidae: Ceutorhynchinae.

    Nadine L. KriskaDepartment of EntomologyUniversity of Wisconsin-MadisonMadison, WI 53706109. Oedemeridae.

    John F. Lawrence, Ph.D.12 Hartwig RoadMothar MountainGympie, QLD 4570 Australia98. Ciidae.

    Richard A.B. Leschen, Ph.D.New Zealand Arthropods CollectionLandcare Research, Private Bag 92170120 Mt. Albert Road, Mt. AlbertAuckland, New Zealand66. Derodontidae; 72. Trogossitidae; 85.Cr yptophagidae; 86. Languriidae; 92.Endomychidae; 94. Corylophidae.

    James E. Lloyd, Ph.D.Entomology and Nematology Depart-

    mentUniversity of FloridaGainesville, FL 3261162. Lampyridae.

    Wenhua LuWes Watkins Agricultural Research and

    Extension CenterOklahoma State UniversityP.O. Box 128Lane, OK 74555101. Mordellidae.

    Adriean J. Mayor, Ph.D.Department of Entomology and Plant

    Pathology205 Ellington Plant Sciences Bldg.University of TennesseeKnoxville, TN 3790174. Melyridae.

    Joseph V. McHugh, Ph.D.Department of EntomologyUniversity of GeorgiaAthens, GA 3060275. Sphindidae; 87. Erotylidae.

    Richard S. Miller, Ph.D.Department of EntomologyMontana State UniversityBozeman, MT 5971759. Lycidae; 60. Telegeusidae.

    Jyrki Muona, Ph.D.Division of EntomologyFinnish Museum of Natural HistoryP. O. Box 17, FIN-00014University of Helsinki, Finland56. Eucnemidae.

    Gayle H. Nelson, Ph.D.1308 N. W. Hawk CreekBlue Springs, MO 64015-178740. Schizopodidae; 41. Buprestidae.

    Sean T. OKeefe, Ph.D.Department of Biological and Environ-

    mental SciencesMorehead State UniversityMorehead, KY 4035138. Dascillidae; 61. Phengodidae.

    Weston Opitz, Ph.D.Department of BiologyKansas Wesleyan University100 Claflin AvenueSalina, KS 6740173. Cleridae.

    T. Keith Philips, Ph.D.Department of BiologyWestern Kentucky UniversityBowling Green, KY 42101-357665. Jacobsoniidae; 70. Anobiidae; 90.Bothrideridae.

    John D. Pinto, Ph.D.Department of EntomologyUniversity of CaliforniaRiverside, California 92521111. Meloidae.

    Darren A. Pollock, Ph.D.Department of BiologyEastern New Mexico UniversityPortales, NM 8813099. Tetratomidae; 100. Melandryidae; 112.Mycteridae; 113. Boridae; 114. Pythidae; 116.Salpingidae; 119. Scraptiidae.

    Michele B. PriceDepartment of EntomologyUniversity of Wisconsin-MadisonMadison, WI 5370678. Smicripidae.

  • xiv

    Robert J. Rabaglia, Ph.D.Maryland Department of Agriculture50 Harry S Truman ParkwayAnnapolis, MD 21401131. Curculionidae: Scolytinae.

    Alistair S. RamsdaleDepartment of EntomologyUniversity of Wisconsin-MadisonMadison, WI 5370663. Omethidae; 64. Cantharidae.

    Brett C. Ratcliffe, Ph.D.W436 Nebraska HallSystematics Research CollectionsUniversity of Nebraska State MuseumLincoln, NE 68588-051423. Lucanidae; 24. Diphyllostomatidae; 34.Scarabaeidae.

    Edward G. RileyDepartment of EntomologyTexas A and M UniversityCollege Station, TX 77843122. Megalopodidae; 123. Orsodacnidae; 124.Chrysomelidae.

    Jack C. Schuster, Ph.D.Universidad del Valle de GuatemalaAptdo. 82Guatemala, Guatemala25. Passalidae.

    William D. Shepard, Ph.D.Department of BiologyCalifornia State University Sacramento6000 J StreetSacramento, CA 9581943. Elmidae; 44. Dryopidae; 45. Lutrochidae;46. Limnichidae; 48. Psephenidae; 51.Eulichadidae.

    Paul E. Skelley, Ph.D.Third EditorFlorida State Collection of ArthropodsFlorida Department of Agriculture and

    Consumer ServicesP. O. Box 147100Gainesville, FL 32614-710034. Scarabaeidae: Aphodiinae; 85.Cr yptophagidae; 86. Languriidae; 87.Erotylidae; 92. Endomychidae.

    Andrew B. T. SmithW436 Nebraska HallSystematics Research CollectionsUniversity of Nebraska State MuseumLincoln, NE 68588-051434. Scarabaeidae.

    Ronald E. Somerby, Ph.D.Plant Pest Diagnostics BranchCalifornia Department of Food and

    Agriculture3294 Meadowview RoadSacramento, CA 95832106. Tenebrionidae.

    Warren E. Steiner, Jr.Department of Systematic Biology -

    EntomologyNational Museum of Natural HistorySmithsonian InstitutionWashington, DC 2056084. Phalacridae.

    Margaret K. Thayer, Ph.D.Field Museum of Natural History1400 S. Lake Shore DriveChicago, IL 6060598. Ciidae.

    Donald B. Thomas, Ph.D.USDA, ARS2301 S. International Blvd.Weslaco, TX 78596106. Tenebrionidae.

    Michael C. Thomas, Ph.D.Second EditorFlorida State Collection of ArthropodsFlorida Department of Agriculture and

    Consumer ServicesP. O. Box 147100Gainesville, FL 32614-710080. Silvanidae; 81. Passandridae; 82.Cucujidae; 83. Laemophloeidae; 91.Cerylonidae; 120. Cerambycidae.

    Charles A. Triplehorn, Ph.D.Museum of Biological DiversityOhio State University1315 Kinnear RoadColumbus, OH 43212106. Tenebrionidae.

    Robert H. Turnbow, Jr., Ph.D.ATZQ-DEL-PC1404 Dilly Branch RoadFort Rucker, AL 36362-5105120. Cerambycidae.

    Barry D. Valentine, Ph.D.2359 Eastcleft DriveColumbus, OH 43221126. Anthribidae.

    Natalia J. Vandenberg, Ph.D.Systematic Entomology LaboratoryPSI, ARS, USDAc/o National Museum of Natural His-

    toryWashington, DC 20560-016893. Coccinellidae.

    Daniel K. Young, Ph.D.Department of EntomologyUniversity of WisconsinMadison, WI 5370635. Eucinetidae; 36. Clambidae; 37. Scirtidae;52. Callirhipidae; 53. Artemetopodidae; 54.Brachypsectridae; 71. Lymexylidae; 96.Mycetophagidae; 97. Archeocrypticidae; 99.Tetratomidae; 107. Prostomidae; 108.Synchroidae; 110. Stenotrachelidae; 115.Pyrochroidae.

    Editorial Board

    J. Howard Frank, D. Phil.Fourth EditorEntomology and Nematology Depart-

    mentUniversity of FloridaGainesville, FL 32611

    David G. Furth, Ph.D.Department of EntomologySmithsonian InstitutionWashington, DC 20560-0165

    Michael A. Ivie, Ph.D.Department of EntomologyMontana State UniversityBozeman, MT 59717

    Brett C. Ratcliffe, Ph.D.Systematic Research CollectionsUniversity of NebraskaLincoln, NE 68588-0514

    Daniel K. Young, Ph.D.Department of EntomologyUniversity of WisconsinMadison, WI 53706

  • Superfamily Scarabaeoidea · 1

    Description. Antennalclub lamellate. Prothorax can behighly  modified  for  burrow-ing,  with  large  coxae  (mostwith concealed trochantins andclosed coxal cavities). Protibiaedentate with a single spur  inmost. Wing venation reducedand  with  a  strong  intrinsicspring mechanism for folding.Tergite 8  forming a  true py-gidium and not concealed bytergite 7. Four Malpighian tu-bules.  Larvae  scarabaeiform(cylindrical, C-shaped). Super-family classification: Lawrenceand  Britton  1991;  Lawrenceand Newton 1995.

    Status of the classifica-tion. The hierarchical level of

    families and subfamilies within the Scarabaeoidea is in disarrayand remains unresolved. In the previous rendition of this work(Arnetts The Beetles of the United States, 1968), the Scarabaeoideaincluded three families: Passalidae, Lucanidae, and Scarabaeidae.This three-family system of classification was the traditionalNorth American system and had several practical and conceptualadvantages. First, it recognized the shared, derived characters thatunite subfamilies within the family Scarabaeidae. Second, it pro-vided a classification system that allowed easy retrieval of hierar-chical information based on the fact that subfamilies were part ofthe family Scarabaeidae (e.g., life history, morphology, larval type).Phylogenetic research indicates that the family Scarabaeidae (in thetraditional sense) is not a monophyletic group. Therefore, wehave chosen to follow the 12-family system established by Browneand Scholtz (1995, 1999) and Lawrence and Newton (1995). Thissystem places emphasis on the differences that separate taxa ratherthan the similarities that unite them. Whereas families, subfami-lies, and tribes in the staphylinoids and curculionoids are beingcombined because of shared characters (thus increasing efficientdata retrieval), the scarabaeoids are being split into numerousfamilies because of  supposed differences (thus, in our view, de-creasing information retrieval). The debate concerning scarabaeoidclassification systems illustrates the weak phylogenetic founda-tion of  the superfamily. This problem is the result of  a numberof factors including (1) lack of thorough study of all scarabaeoidtaxa, (2) lack of diagnostic characters for all taxa, (3) lack of phylo-genetic study of all taxa, (4) prevailing philosophies regardingcategorical levels, and (5) emphasis in research on the less speciosegroups of scarabaeoids and lack of research on the more speciosegroups (such as the subfamilies of Scarabaeidae including theMelolonthinae,  Rutelinae,  Dynastinae,  Aphodiinae,  andCetoniinae).

    Series SCARABAEIFORMIA Crowson 1960

    (=  Lamellicornia)

    Superfamily  SCARABAEOIDEA  Latreille  1802

    INTRODUCTION

    by Mary Liz Jameson and Brett C. Ratcliffe

    Common name: The scarabaeoid beetles

    The superfamily Scarabaeoidea is a large, diverse, cosmopolitan group of  beetles. Scarabaeoids are adapted tomost  habitats,  and  they  are  fungivores,  herbivores,  necrophages,  coprophages,  saprophages,  and  some  arecarnivores. They are widely distributed, even living in the Arctic in animal burrows. Some scarabs exhibit paren-tal  care  and  sociality.  Some are myrmecophilous,  termitophilous, or  ectoparasitic. Many possess  extravagant horns,others are able to roll into a compact ball, and still others are highly armored for inquiline life. Some are agriculturalpests that may destroy crops while others are used in the biological control of  dung and dung flies. Scarabaeoids arepopular beetles due to their large size, bright colors, and interesting natural histories. Early Egyptians revered the scarabas a god, Jean Henri Fabre studied their behavior, and Charles Darwin used observations of  scarabs in his theory ofsexual selection.

    Acknowledgments.  Mary  Liz  Jameson  and  Brett  Ratcliffe  thankour  scientific  illustrators, Mark Marcuson and Angie Fox,  for  theirfine  work.  Al  Newton,  Margaret  Thayer,  Philip  Harpootlian,  andAndrew  Smith  are  especially  acknowledged  for  assistance  withresolving  authorship  of  some  taxa  and  numerous  grammaticalcorrections.  For  assistance  with  the  keys  and  other  aspects  of  themanuscript,  we  thank  Andrew  Smith,  Federico  Ocampo,  WilliamWarner,  and  Charles  Brodel.  Our  development  of   the  scarabaeoidchapters was supported by a NSF/PEET grant (DEB9712447) to B.C. Ratcliffe  and M. L.  Jameson. Parts of  the Dynastinae  treatmentwere  supported  by  a  NSF/BS&I  grant  (DEB9870202)  to  B.  C.Ratcliffe.

    FIGURE  1.  Anomala binotata(Gyllenhal)  (Scarabaeidae)  (Usedby  permission  of  University  ofNebraska State Museum)

  • 2 · Superfamily Scarabaeoidea

    Within the Scarabaeoidea there is a disparity in the knowl-edge between less speciose basal lineages and the more speciosegroups  of  higher  Scarabaeidae.  For  example,  the  familyTrogidae includes approximately 300 species in four genera. Ex-cellent revisionary, larval, and phylogenetic studies are availablefor this group (Baker 1968; Scholtz 1982, 1986, 1990, 1991, 1993;Scholtz and Peck 1990). Excellent monographs are also availablefor the approximately 600 species of Geotrupidae (Howden 1955,1964, 1979, 1985a-b, 1992; Howden and Cooper 1977; Howdenand Martínez 1978) and the Trogidae (Vaurie 1955), and thesewill provide the foundation for addressing relationships withinthis group. In comparison, the family Scarabaeidae (sensu Lawrenceand Newton 1995) includes approximately 91% of the species (ca27,800) of Scarabaeoidea. Within the Scarabaeidae, approximately21,000 species are in the subfamilies Melolonthinae, Dynastinae,Rutelinae, and Cetoniinae (the higher scarabs). Only a few phy-logenetic analyses have addressed relationships of pleurostictsubtribes, genera, or species (Ratcliffe 1976; Ratcliffe and Deloya1992; Jameson 1990, 1996, 1998; Jameson et al. 1994; Krell 1993),and only one analysis has been conducted to address tribal orsubfamilial relationships (Browne and Scholtz 1999).

    Historically, the superfamily Scarabaeoidea was divided intotwo generalized groups based on the position of the abdominalspiracles; the Laparosticti and Pleurosticti. Pleurostict scarabs werecharacterized by having most of the abdominal spiracles situatedon the upper portion of  the sternites (Ritcher 1969; Woodruff1973) and included taxa whose adults feed on leaves, flowers andpollen, and whose larvae feed primarily on roots and decayingwood. Laparostict scarabs, on the other hand, were characterizedby having most of the abdominal spiracles located on the pleuralmembrane between the tergites and sternites (Ritcher 1969) andincluded taxa whose adults and larvae feed on dung, carrion,hides, and feathers. The position of the spiracles, however, is nota consistent character (Ritcher 1969), and, in recent years, subfami-lies and tribes that were once included in the Laparosticti havebeen raised to higher taxonomic status (family and subfamily,respectively).

    The composition of the Scarabaeoidea remains a topic ofdebate. Lawrence and Newton (1995) proposed 13 families (12found in the Nearctic, Belohinidae are Madagascan), and Scholtzand Browne (1996) and Browne and Scholtz (1995, 1998, 1999)proposed  13  families  (all Nearctic,  including Bolboceratidae;Belohinidae were not addressed). In this work we follow, withsome hesitation, the system of Lawrence and Newton (1995)and treat the Scarabaeoidea as including 12 Nearctic families (eightor nine of which were previously considered subfamilies of thefamily Scarabaeidae, and one of which was previously considereda subfamily of the Lucanidae). Our reluctance to accept elevationof new families within the Scarabaeoidea stems from the factthat: 1) there have been no comprehensive taxonomic treatmentsof all higher categories of scarabaeoids (families and subfamilies)and, 2) there are few comprehensive, rigorous, phylogenetic analy-ses  of  higher  scarabaeoid  groups  and,  thus,  a  lack  ofsynapomorphic characters that establish a basis for uniform fa-milial and subfamilial levels. We prefer to see clades delimited by

    shared derived characters before the elevation of certain taxa tofamily level. Despite our reluctance to accept this classificationsystem, we have little basis for disputing the validity of currenttaxonomic conclusions other than the fact that some of thesetaxonomic conclusions have been based on narrow taxonomicframeworks (only scarab taxa from certain geographic regionsrather than all scarab groups) or based on few characters or suitesof characters.

    Underlying the classification problem is, of course, the factthat we are dealing with constructs that are 200 years old and thatpre-date evolutionary theory. Linnaean classifications were basedon overall morphological similarity rather than shared, derivedcharacters. Thus, some groups within the scarabaeoids are notmonophyletic lineages; instead, they are groups that were createdhistorically because they superficially resembled each other. Oursystem of classification needs to convey information and con-cepts and allow for easy retrieval of information. Whether a cer-tain taxon is classified at the level of family or subfamily may betrivial if  we can continue to convey the needed information. Weremain apprehensive that the trend of elevation to many familieswithin the Scarabaeoidea will result, at least in the short term, in anet loss in retrievability of information.

    Despite the considerable debate, phylogenetic analyses ofscarabaeoid higher categories are on-going and their results bringus closer to understanding relationships of the groups. A pre-liminary total evidence phylogenetic analysis of 13 families ofScarabaeoidea (excluding Belohinidae, including Bolboceratidae)and most of the subfamilies was conducted using 134 adult andlarval characters (Brown and Scholtz 1999). Results of  this analy-sis showed that the superfamily Scarabaeoidea is comprised ofthree major lineages: the glaresid lineage that consists of only thefamily Glaresidae; the passalid lineage that consists of two majorlines--a  glaphyrid  line  (containing  Glaphyridae,  Passalidae,Lucanidae, Diphyllostomatidae, Trogidae, Bolboceratidae, andPleocomidae),  and  a  geotrupid  line  (containing Geotrupidae,Ochodaeidae, Ceratocanthidae, and Hybosoridae); and the scarablineage  (containing  Aphodiinae,  Scarabaeinae,  Orphninae,Melolonthinae, Rutelinae, Dynastinae, and Cetoniinae).

    The series Scarabaeiformia is comprised exclusively of thesuperfamily  Scarabaeoidea.  Monophyly  of  the  group  is  wellfounded and undisputed (Lawrence and Britton 1991). The sis-ter group for the Scarabaeoidea, however, is not resolved andcontinues  to  be  debated.  Two  groups  are  considered:  theStaphyliniformia and the Dascilloidea. The Scarabaeoidea andStaphyliniformia share characters of the wing venation and theabdomen that are not present in the dascilloids (Kukalová-Peckand Lawrence 1993). The Scarabaeoidea and Dascilloidea sharesimilar larval characters (lack of  urogomphi that are present inStaphyliniformia, cribriform spiracles, separate galea and lacinia)and adult characters (form of the ommatidium, male genitalia,mouthparts) (Scholtz et al. 1994). Lawrence and Newton (1982)argued that similarities in the Dascilloidea and Scarabaeoidea areattributable to either plesiomorphic or convergent characters thatare associated with soil-dwelling habits.

  • Superfamily Scarabaeoidea · 3

    Distribution. The superfamily Scarabaeoidea is one of thelargest superfamilies in the Coleoptera and includes approximately2,200 genera and about 31,000 species worldwide (Dalla Torre1912-1913; Endrödi 1985; Hanski and Cambefort 1991; Krikken1984; Lawrence 1982; Machatschke 1972; Scholtz 1982). Whilesome  of  the  smaller  groups  are  well  known  worldwide  (e.g.,Geotrupidae and Trogidae), some other groups (e.g., Scarabaeidaethat comprises 91% of the Scarabaeoidea) cannot be identified toeven genus-level with reliability.

    In the Nearctic region, the taxonomy of most scarabaeoidsis now fairly well known although there remain a few areas ofuncertainty. For example, the phylogenetic position of  both thePleocomidae and the Hopliini needs to be addressed. In thiswork, there are numerous changes in the author and/or date ofmany genera and even some higher categories since Arnett (1968).These are the result of greater scrutiny of the original literaturerather than accepting at face value the often erroneus or incom-plete information provided in older catalogs and faunal works.

    There are approximately 170 genera and 2,000 species in theUnited States, Canada, and Nearctic Mexico. Smith (2001) pro-vided a checklist of Nearctic scarabaeoids. Regional works: Blatchley1910; Loding 1945; Saylor 1948; Edwards 1949; Helgesen andPost 1967; Hatch 1971; Woodruff  1973; Kirk and Balsbaugh1975; Shook 1978; Lago et al. 1979; Ratcliffe 1991; Downie andArnett 1996; Morón et al. 1997; Harpootlian 2001.

    SUMMARY OF THE FAMILIES AND SUBFAMILIESOF SCARABAEOIDEA OF THE UNITED STATES,

    CANADA, AND NEARCTIC MEXICO

    23. LucanidaeLucaninaeNicaginaeSyndesinae

    24. Diphyllostomatidae25. Passalidae26. Glaresidae27. Trogidae28. Pleocomidae29. Geotrupidae

    BolboceratinaeGeotrupinae

    30. OchodaeidaeOchodaeinaeChaetocanthinae

    31. Hybosoridae32. Ceratocanthidae33. Glaphyridae34. Scarabaeidae

    AphodiinaeScarabaeinaeMelolonthinaeRutelinaeDynastinaeCetoniinae

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    LAWRENCE, J. F. and A. F. NEWTON, Jr. 1982. Evolution andclassification of beetles. Annual Review of Entomology andSystematics, 13: 261-290.

    LAWRENCE, J. F. and A. F. NEWTON, Jr. 1995. Families andsubfamilies of Coleoptera (with selected genera, notes, refer-ences and data on family-group names). Pp. 779-1006. In: J.Pakaluk and S. A. Slipínski, eds. Biology, Phylogeny, andClassification of Coleoptera. Papers Celebrating the 80thBirthday of Roy. A. Crowson. Muzeum i Instytut ZoologiiPAN. Warsaw, 1092 pp.

    LODING, H. P. 1945. Catalogue of the beetles of Alabama.Alabama Geological Survey Monograph, 11: 1-172.

    MACHATSCHKE, J. W. 1972. Scarabaeoidea: Melolonthidae,Rutelinae. Coleopterorum Catalogus Supplementa, 66: 1-361.

    MORÓN, M. A., B. C. RATCLIFFE, and C. DELOYA. 1997.Atlas de los escarabajos de México. Coleoptera: Lamellicornia.Vol.  1.  Familia  Melolonthidae.  Subfamilias  Rutelinae,Dynastinae, Cetoniinae, Trichiinae, Valginae y Melolonthinae.Sociedad Mexicana de Entomología, Mexico 280 pp.

    RATCLIFFE,  B.  C.  1976.  A  revision  of  the  genus  Strategus(Coleoptera: Scarabaeidae). Bulletin of the University of Ne-braska State Museum, 10: 93-204.

    RATCLIFFE, B. C. 1991. The scarab beetles of Nebraska. Bulletinof the University of Nebraska State Museum, 12: 1-333.

    RATCLIFFE, B. C. and A. C. DELOYA. 1992. The biogeographyand phylogeny of Hologymnetis  (Coleoptera: Scarabaeidae:Cetoniinae) with a revision of the genus. Coleopterists Bul-letin, 46: 161-202.

    RITCHER, P. O. 1969. Spiracles of adult Scarabaeoidea (Co-leoptera) and their phylogenetic significance. I. The abdomi-nal spiracles. Annals of the Entomological Society of America,62: 869-880.

    SAYLOR, L. W. 1948. Contributions toward a knowledge of theinsect  fauna  of  lower  California.  No.  10.  Coleoptera,Scarabaeidae. Proceedings of the California Academy of Sci-ences, Series 4, 24: 337-374.

    SCHOLTZ, C. H. 1982. Catalogue of the world Trogidae (Co-leoptera: Scarabaeoidea). Republic of South Africa, Depart-ment of Agriculture and Fisheries, Entomology Memoire,54: 1-27.

  • Superfamily Scarabaeoidea · 5

    SCHOLTZ, C. H. 1986. Phylogeny and systematics of the Trogidae(Coleoptera: Scarabaeoidea). Systematic Entomology, 11:355-363.

    SCHOLTZ, C. H. 1990. Phylogenetic trends in the Scarabaeoidea(Coleoptera). Journal of Natural History, 24: 1027-1066.

    SCHOLTZ, C. H. 1991. Descriptions of larvae of AustralianOmorgus Erichson, with implications for the phylogeny of theTrogidae (Insecta: Coleoptera). Invertebrate Taxonomy, 5:827-835.

    SCHOLTZ, C. H. 1993. Descriptions of larvae of African Trogidae(Coleoptera), with  implications for  the phylogeny of  thefamily. African Entomology, 1: 1-13.

    SCHOLTZ, C. H., D. J. BROWNE and J. KUKALOVÁ-PECK1994. Glaresidae, archeopteryx of the Scarabaeoidea (Co-leoptera). Systematic Entomology, 19: 259-277.

    SCHOLTZ, C. H. and D. J. BROWNE. 1996. Polyphyly in theGeotrupidae (Coleoptera: Scarabaeoidea): a case for a newfamily Bolboceratidae. Journal of Natural History, 30: 597-614.

    SCHOLTZ, C. H. and S. PECK. 1990. Description of a PolynoncusBurmeister  larva, with  implications for phylogeny of the

    Trogidae (Coleoptera: Scarabaeoidea). Systematic Entomol-ogy, 15: 383-389.

    SHOOK, G. A. 1978. Records of  some  scarabs  from Idaho(Coleoptera: Scarabaeidae). Coleopterists Bulletin, 32: 52.

    SMITH, A. B. T. 2001. Checklist of the Scarabaeoidea of theNearctic Realm (Includes Canada, the continental UnitedStates, and the following states of northern Mexico: BajaCalifornia,  Baja  California  Sur,  Chihuahua,  Coahuila  deZaragoza,  Durango,  Nuevo  Leon,  Sinaloa,  Sonora,Tamaulipas,  and  Zacatecas).  URL:  http://www-museum.unl.edu/research/entomology/nearctic.htm.

    VAURIE, P. 1955. A revision of the genus Trox in North America.Bulletin of the American Museum of Natural History, 106: 1-89.

    WOODRUFF, R. E. 1973. The scarab beetles of Florida (Co-leoptera: Scarabaeidae). Part. I. The Laparosticti (subfamilies:Scarabaeinae,  Aphodiinae,  Hybosorinae,  Ochodaeinae,Geotrupinae, Acanthocerinae). Arthropods of Florida andNeighboring Land Areas, 8: 1-220.

  • 6 · Family 23. Lucanidae

    23. LUCANIDAE Latreille 1804

    by Brett C. Ratcliffe

    Family common name: The stag beetles

    Stag beetles range in size from less than 1 cm to 9 cm. The largest Nearctic species attains a length of 6 cm. Mostmales  possess  greatly  enlarged,  curving  mandibles  that  are  sometimes  used  in  combat  with  male  opponentsduring fights to establish dominance. Female lucanids lack enlarged mandibles. In most, male development of  themandibles is allometric, that is, the size of  the mandibles is proportional to the size of  the body. Those males with thelargest mandibles are referred to as male majors and those with the smallest mandibles are called male minors.

    Description. Length 8.0-60.0 mm. Shape usually weaklyconvex, subdepressed, or cylin-drical,  elongate.  Color  testa-ceous  to  reddish  brown  toblack.

    Head prognathus, not de-flexed. Antennae geniculate orstraight,  10-segmented,  with3-7  segmented  club  (allantennomeres  unopposableand  tomentose);  first  ant-ennomere  often  subequal  toremaining  antennomeres.Eyes with eucone or acone om-matidia; eye canthus present orabsent.  Clypeus  and  labrumfused to frons. Mandibles pro-duced beyond apex of labrum,prominent (males often withlarge,  curved,  elongate  man-

    dibles).  Maxillae  with  4-segmented  palpi;  labium  with  3-seg-mented palpi.

    Pronotum weakly convex, base narrower than elytral base,lacking tubercles, ridges, horns, or sulci. Elytra weakly convex,with or without impressed striae. Scutellum exposed, triangularor parabolic. Pygidium concealed by elytra or only weakly exposed.Legs with coxae transverse, mesocoxae separated; protibiae den-tate on outer margin, apex with one spur; meso- and metatibiawith ridges, apex with 2 spurs; tarsi 5-5-5; claws equal in size,simple;  empodium  present,  extending  weakly  beyond  fifthtarsomere or extending nearly one half claw length, with 2 toseveral setae. Profemora with tomentose patch anteriorly.

    Abdomen with 5 visible sternites; 8 functional abdominalspiracles situated in pleural membrane. Wings well developed,with M-Cu loop and two, apical, detached veins. Male genitaliatrilobed. References: Didier and Seguy 1953; Scholtz 1990.

    Larvae are scarabaeiform (c-shaped, subcylindrical). Colorcreamy-white or yellowish (except at caudal end which may bedarkened by accumulated feces). Cranium heavily sclerotized,lightly pigmented. Antennae 3-4 segmented, last segment greatly

    reduced  in  size.  Stemmata  absent  (present  in  Platycerus).Frontoclypeal suture present. Labrum at apex rounded or weaklylobed. Epipharynx rounded or lobed, with symmetrical tormae.Maxilla with galea and lacinia distinctly separate; maxillary stridu-latory teeth absent (present in Platycerus); maxillary palpus 4-seg-mented. Mandibles elongate, asymmetrical. Abdominal segments3-7 with 2 annuli, each with 1 or more transverse rows of shortsetae. Spiracles cribriform. Anal opening Y-shaped or longitudi-nal, surrounded by 2 fleshy lobes. Legs 4-segmented. Stridula-tory apparatus on meso- and metathoracic legs present; clawspresent. References: Ritcher 1966; Scholtz 1990.

    Habits and habitats. Lucanids are usually associated withdecaying wood and logs in coniferous and deciduous forest habi-tats. Adults of some species are attracted to lights at night andsome feed at sap flows from fluxing trees. Adults of some smallerspecies have been observed feeding on flowers. The eggs arecustomarily laid in crevices in bark or logs, and the larvae feed ondecaying wood. The larvae resemble those of  Scarabaeidae, but inlucanids the anal opening is longitudinal or Y-shaped transverseor Y-shaped in a few. References: Ratcliffe 1991.

    Status of the classification. The family Lucanidae has longbeen  considered  one  of  the  most  primitive  groups  in  theScarabaeoidea (Crowson 1967; Howden 1982; Ritcher 1966), andscarabaeoid classifications and evolutionary hypotheses have gen-erally regarded the Lucanidae as basal to all scarabaeoids (Howden1982; Iablokoff-Khnzorian 1977; Lawrence and Newton 1995).However, based on comparison with primitive scarabaeoidgroups, Scholtz et al. (1994) hypothesized that the scarabaeoidfamily Glaresidae, rather than the Lucanidae, is the most primi-tive scarabaeoid. According to this hypothesis, the Lucanidae is amember of a clade including the Passalidae, Diphyllostomatidae,Glaphyridae,  Trogidae,  Pleocomidae,  and  Bolboceratinae(Geotrupidae).

    Prior to the taxonomic elevation of the genus Diphyllostomato the family Diphyllostomatidae (Holloway 1972), the Lucanidaewas hypothesized to be most closely related to the Passalidae(Howden 1982). Based on shared characters, it is now thoughtthat  the  Lucanidae  is  most  closely  related  to  theDiphyllostomatidae (Caveney 1986; Browne and Scholtz 1995).

    The world Lucanidae (about 800 species) have been treatedin checklists by Benesh (1960) and Maes (1992) and in illustrated

    FIGURE  23.1.  Lucanus capreolus(Linnaeus) (Used by permission ofUniversity  of  Nebraska  StateMuseum)

  • Family 23. Lucanidae · 7

    catalogs by Didier and Seguy (1953) and Mizunuma and Nagai(1994). The latter is spectacular for its colored plates of the worldfauna. Benesh (1960) recognized eight subfamilies, four of whichoccurred in the United States. The classification of the NorthAmerican species had been relatively stable until Howden andLawrence (1974) proposed a significant rearranging of generawithin subfamilies based partly on the work of Holloway (1960,1968, 1969). This newer classification is followed here, with threesubfamilies  now  recognized  as  occurring  in  North  America:Nicaginae, Syndesinae, and Lucaninae.

    Distribution. The world fauna consists of about 800 spe-cies (Mizunuma and Nagai 1994) with more species found inAsia than in other areas. In North America there are three sub-families with eight genera and 24 species. Keys to adults: Benesh1946; Blatchley 1910; Howden and Lawrence 1974; Ratcliffe 1991.Keys to larvae: Ritcher 1966; Smith 2001. World catalog: Krajcik2001. Regional works: Edwards 1949; Hatch 1971; Kirk andBalsbaugh 1975; Ratcliffe 1991. Biology: Milne 1933; Hoffman1937.

    KEY TO THE NEARCTIC SUBFAMILIES, TRIBES,AND GENERA OF LUCANIDAE

    1. Antenna straight, not geniculate (Fig. 2); body formoval or cylindrical; prosternal process narrow, an-terior coxae nearly contiguous ....................... 2

    Antenna geniculate (Fig. 3); body form elongate,weakly flattened; prosternal process broad, an-terior coxae distinctly separated (Lucaninae) . 4

    2(1). Form short, oval, convex; elytra coarsely punctatewith conspicuous bristles; antenna with clublamellate (Nicaginae) ............................. Nicagus

    Form elongate, cylindrical; elytra striate and nearlyglabrous; antenna with club only weakly lamel-late (Syndesinae) .............................................. 3

    3(2). Head of male with long, median horn; female withmedian tubercle; mandibles in both sexes smalland inconspicuous (Sinodendrini) . Sinodendron

    Head in both sexes lacking median horn or tubercle;mandibles large, conspicuous, especially in male(Ceruchini) ............................................ Ceruchus

    4(1). Eye nearly entire, canthus absent or very short;antenna weakly geniculate, scape lacking grooveat apex; body length mostly less than 15 mm(Platycerini) ....................................................... 5

    Eye divided by canthus, canthus more than 1/4 aslong as eye; antenna strongly geniculate, scapewith apical groove; body length more than 15 mm......................................................................... 7

    5(4). Body robust, obovate, strongly convex; posteriortibia stout, expanded at apex, triangular in crosssection; apical spurs spatulate ..... Platyceropsis

    Body distinctly depressed; posterior tibia narrow,not expanded at apex, not triangular in cross sec-tion; apical spurs slender, not spatulate .......... 6

    6(5). Anterior margin of head semicircularly excised;clypeus mostly concealed; mandibles subequalto length of head; club of antenna with 4 seg-ments in male, 3 segments in female ........................................................................... Platycerus

    Anterior margin of head lacking excision on front;clypeus distinctly produced; mandibles about halflength of head; club of antenna 3-segmented inboth sexes .................................... Platyceroides

    7(4). Elytra nearly smooth; pronotum with lateral marginstrongly arcuate or angulate; posterior tibia with0, 2, or 3 spines along outer edge (Lucanini) .................................................................. Lucanus

    Elytra striate-punctate; pronotum with lateral mar-gins subparallel; posterior tibia with 1 spine onouter edge (Dorcini) ................................ Dorcus

    CLASSIFICATION OF THE NEARCTIC GENERA

    Lucanidae Latreille 1804

    Lucaninae Latreille 1804

    Characteristics: Eye partly or completely divided by canthus.Antenna  geniculate.  Body  form  elongate,  weakly  flattened.Pronotal process broad between procoxae, coxae distinctly sepa-rated. Internal sac of aedeagus permanently everted.

    Most of the lucanids of the world are found in this subfam-ily although many of the tribes are poorly or inconsistently char-acterized (Howden and Lawrence 1974). The classification of theNorth American genera seems now to be stable.

    Lucanini Latreille 1804

    Characteristics: Antenna strongly geniculate, scape with apicalgroove. Elytra nearly smooth or with minute and irregular punc-tation. Pronotum with lateral edges arcuate or angulate. Metatibialacking spines or with 2 or 3 spines along outer edge. Most withbody length greater than 25 mm.

    This tribe consists of  two genera in the New World. Onegenus, Cantharolethrus, is found from Mexico to South America.Keys to U.S. species: Fuchs 1882; Dillon and Dillon 1961. Keys tolarvae: Ritcher (1966) was unable to distinguish between the threeU.S. species he examined.

    FIGURES  2.23-3.23.  Right  antenna  dorsal  view  of:  2.  Ceruchuspiceus  and 3. Platycerus  sp.

    2 3

  • 8 · Family 23. Lucanidae

    States and Canada. Key to species: Benesh 1946. Larval descrip-tion: Ritcher 1966. Biology: Hoffman 1937.

    Nicaginae LeConte 1860

    Characteristics: Antenna not geniculate, instead straight, withstrongly lamellate club. Eye entire (lacking canthus). Pronotumwith lateral margins strongly crenulate. Elytra coarsely punctate,with conspicuous bristles. Prosternal process narrow so that an-terior coxae almost contiguous. Aedeagus with internal sac ever-sible. Body form short, oval, convex.

    According to Howden and Lawrence (1974), the Nicaginaecontain two genera: Ceratognathus from Australia and New Zealandand Nicagus from North America and Japan. Previously, thesegenera were included in the subfamily Aesalinae.

    Nicagus LeConte 1860One species, N. obscurus LeConte, occurs in the eastern half  of  theUnited States.

    Syndesinae MacLeay 1819

    Characteristics: Antenna straight, not geniculate; club weaklylamellate. Eye entire (lacking canthus). Elytra striate, nearly gla-brous. Prosternal process narrow, anterior coxae nearly contigu-ous. Aedeagus with internal sac eversible. Body form elongate,cylindrical.

    This subfamily is composed of three tribes: Syndesini withthe genera Syndesus (from Australia, Tasmania, New Caledonia,and Africa) and Psilodon (from South America); Sinodendrini withthe genus Sinodendron (from North America); and Ceruchini withthe single Holarctic genus Ceruchus.

    Sinodendrini Mulsant 1842

    Characteristics: Head of male with long, median horn; femalewith median tubercle. Mandibles in both sexes small and incon-spicuous.

    Sinodendron Hellwig 1894Ligniperda Fabricius 1801

    This genus contains two species and one, S. rugosum Mannerheim,is found in the Pacific Northwest. Generic overview: Hatch 1928.

    Ceruchini Jacquelin du Val 1857

    Characteristics: Head lacking median horn or tubercle in bothsexes. Mandibles large, conspicuous (especially in male).

    Ceruchus MacLeay 1819Seven species occur in this Holarctic genus (Benesh 1960), andthree species are found in North America. One species occurs insoutheastern Canada and the northeastern quadrant of the UnitedStates while the other two species are found from California to

    Lucanus Scopoli 1763Hexaphyllus Mulsant 1839Pseudolucanus Hope 1845

    The genus Lucanus (Fig. 1) contains about 50 species distributedin Asia, Europe, and North America (Benesh 1960). Most of thespecies occur in Asia while five species occur in the United Statesand northern Mexico. Three species are restricted to the easternUnited States while the other two are found in the Southwestand northern Mexico. Keys to species: Dillon and Dillon 1961.Biology: Milne 1933.

    Dorcini Parry 1864

    Characteristics: Antenna strongly geniculate, scape with apicalgroove.  Elytra  striate-punctate.  Pronotum  with  lateral  edgessubparallel. Metatibia with 1 spine on outer edge. Body lengthmore than 15 mm.

    There is only one genus of Dorcini represented in NorthAmerica: Dorcus. The tribe Dorcini has six genera, five of whichare endemic to Asia.

    Dorcus MacLeay 1819The genus Dorcus contains about 30 species; most occur in Asiaand India although two are also found in southern Europe, onein Mexico, and two in the United States (Benesh 1937). The twoU.S. species are found in the eastern half  of  the country, and onespecies occurs in Quebec. Key to species: Downie and Arnett1996.

    Platycerini Mulsant 1842

    Characteristics: Antenna weakly geniculate, scape lacking apicalgroove. Eye nearly entire, eye canthus absent or very short. Elytrastriate in most. Most with body length less than 15 mm.

    The  tribe  Platycerini  consists  of  three  genera  in  NorthAmerica, with Platycerus also being found in Asia, Europe, andNorth Africa. Key to genera: Benesh 1946; Howden and Lawrence1974.

    Platyceroides Benesh 1946This genus contains seven species, all of which are  found  inWashington, Oregon, and northern California. Key to species:Benesh 1946.

    Platyceropsis Benesh 1946This monobasic genus contains one species, P. keeni (Casey), whichoccurs from British Columbia to northern California.

    Platycerus Geoffroy 1762Systenocerus Weise 1883Systenus Sharp and Muir 1912

    Eight species are found in this genus, and five of these occur inthe United States. Four of  the U.S. species are found in the PacificNorthwest while the fifth species is found in the eastern United

  • Family 23. Lucanidae · 9

    Vancouver  Island. Larval  description: Ritcher  1966. Biology:Hoffman 1937.

    BIBLIOGRAPHY

    BENESH, B. 1937. Some notes on boreal American Dorcinae(Coleoptera: Lucanidae). Transactions of the American Ento-mological Society, 63: 1-15.

    BENESH, B. 1946. A systematic revision of the Holarctic genusPlatycerus Geoffroy. Transactions of the American Entomo-logical Society, 72: 139-202.

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