NUMBER 443 13 JUNE 1994 - REVISION OF PHANAEUS MACLEAY, A NEW WORLD GENUS OF SCARABAEINE DU~G BEETLES (COLEOPTERA: SCARABAEIDAE, SCARABAEINAE) -----------------------_..--------------------- w. D. EDMONDS N ATURAL HISTORY I\lUSEUM OF Los ANGELES COUNTY The scientific publications of the Natural History Mu­ SERIAL seum of Los Angeles County have been issued at irregular intervals in three major series; the issues in each series are PUBLICATIONS numbered individually, and numbers run consecutively, OF THE regardless of the su bject matter. • Contributions in Science, a miscellaneous series of tech­ NATURAL HISTORY nical papers describing original research in the life and earth sciences. MUSEUM OF • Science Bulletin, a miscellaneous series of monographs describing original research in the life and earth sci­ Los ANGELES ences. This series was discontinued in 1978 with the issue of Numbers 29 and 30; monographs are now COUNTY published by the Museum in Contributions in Science. • Science Series, long articles and collections of papers on natural history topics. Copies of the publications in these series are sold through the Museum Book Shop. A catalog is available on request. The Museum also publishes Technical Reports, a mis­ cellaneous series containing information relative to schol­ arly inquiry and collections but not reporting the results of original research. Issue is authorized by the Museum's Scientific Publications Committee; however, manuscripts do not receive anonymous peer review. Individual Tech­ nical Reports may be obtained from the relevant Section of the Museum. SCIENTIFIC PUBLICATIONS COMMITTEE Craig C. Black, Museum Director NATURAL HISTORY MUSEUM Daniel M. Cohen OF Los ANGELES COUNTY Kirk Fitzhugh 900 EXPOSITION BOULEVARD John M. Harris, Committee Chairman Charles L. Hogue Los ANGELES, CALIFORNIA 90007 Edward C. Wilson Richard C. Hink, Managing Editor Printed at Allen Press, Inc., Lawrence, Kansas Robin A. Simpson, Head of Publications ISSN 0459-8113 REVISION OF PHANAEUS MACLEAY, A NEW WORLD GENUS OF SCARABAEINE DUNG BEETLES COLEOPTERA: SCARABAEIDAE, SCARABAEINAE w. D. EDMONDS1 CONTENTS ABSTRACT • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 3 RESU MEN ........................................................................ 3 INTRODUCTION .................................................................. 3 HISTORY OF THE GENUS PHANAEUS .............................................. 3 PRESENT CLASSIFICATION ........................................................ 4 Taxonomic Characters ............................................................. 4 Primary Type Material ............................................................. 5 Taxonomic Summary ............................................................... 8 Phanaeus Macleay, 1819 . .. 8 Invalid Names Assigned to Phanaeus Macleay ........................................ 8 BIOLOGY, GEOGRAPHY, AND EVOLUTION OF PHANAEUS. .. 9 South American Patterns ................................................... .. 12 Middle American Patterns .......................................................... 13 N orrh American Patterns ........................................................... 13 KEY TO THE PHANAEINE GENERA ................................................ 17 SYSTEMATICS OF THE GENUS PHANAEUS MACLEAY ................................ 17 Key to the Subgenera of Phanaeus ................................................... 18 Notiophanaeus, New Subgenus .................................................... 18 Key to the Species Groups of N otiophanaeus ...................................... 19 The Splendidulus Group ....................................................... 19 Key to the Species of the Splendidulus Group .................................... 19 Phanaeus (N otiophanaeus) splendidulus (Fabricius) ............................... 22 Phanaeus (Notiophanaeus) dejeani Harold ...................................... 23 Phanaeus (N otiophanaeus) haroldi Kirsch ....................................... 24 Phanaeus (N otiophanaeus) melibaeus Blanchard .................................. 24 The Chalcomelas Group ....................................................... 25 Key to the Species of the Chalcomelas Group .................................... 27 Phanaeus (Notiophanaeus) chalcomelas (Perry) ................................... 28 Phanaeus (Notiophanaeus) achilles Boheman .................................... 30 Phanaeus (N otiophanaeus) cambeforti Arnaud ................................... 31 Phanaeus (N otiophanaeus) meleagris Blanchard .................................. 31 The Bispinus Group ........................................................... 33 Key to the Species of the Bispinus Group ....................................... 33 Phanaeus (N otiophanaeus) alvarengai Arnaud ................................... 34 Phanaeus (N otiophanaeus) bispinus Bates ....................................... 35 The Palaeno Group ........................................................... 35 Key to the Species of the Palaeno Group ........................................ 36 Phanaeus (Notiophanaeus) kirbyi Vigors ........................................ 37 Phanaeus (N otiophanaeus) palaeno Blanchard ................................... 38 1. Department of Biological Sciences, California State Polytechnic University, Pomona, 3801 Temple Avenue, Po­ mona, California 91768. Research Associate, Natural History Museum of Los Angeles County, Los Angeles, California 90007. Investigador Invitado, Instituto de Ecologia, Xalapa, Veracruz. Contributions in Science, Number 443, pp. 1-105 Natural History Museum of Los Angeles County, 1994 The Endymion Group .......................................................... 39 Key to the Species of the Endymion Group. .. 39 Phanaeus (N otiophanaeus) halffterorum Edmonds . .. 41 Phanaeus (Notiophanaeus) endymion Harold ..................................... 43 Phanaeus (Notiophanaeus) pyrois Bates .......................................... 44 Subgenus Phanaeus Macleay . .. 46 Key to the Species Groups of Phanaeus, sen. str. .................................... 46 The Hermes Group. .. 47 Key to the Species of the Hermes Group ......................................... 48 Phanaeus (Phanaeus) hermes Harold ............................................ 48 Phanaeus (Phanaeus) prasinus Harold ........................................... 49 The Tridens Group ............................................................. 49 Key to the Species and Subspecies of the Tridens Group ............................ 50 Phanaeus (Phanaeus) nimrod Harold ............................................ 52 Phanaeus (Phanaeus) tridens Laporte-Castelnau ................................... 53 Phanaeus (Phanaeus) eximius Bates ............................................. 54 P hanaeus (P hanaeus) furiosus Bates. .. 54 Phanaeus (Phanaeus) daphnis Harold ........................................... 55 The Triangularis Group ......................................................... 56 Key to the Species and Subspecies of the Triangularis Group ........................ 56 Phanaeus (Phanaeus) adonis Harold ............................................ 57 Phanaeus (Phanaeus) triangularis (Say) ........................................... 58 The Mexicanus Group. .. 60 Key to the Species and Subspecies of the Mexicanus Group. .. 60 P hanaeus (P hanaeus) flohri N evinson. .. 63 Phanaeus (Phanaeus) demon Laporte-Castelnau ............ .. 65 Phanaeus (Phanaeus) scutifer Bates ............................................. 67 Phanaeus (Phanaeus) lunaris Taschenberg . .. 67 Phanaeus (Phanaeus) wagneri Harold ........................................... 68 Phanaeus (Phanaeus) mexicanus Harold ......................................... 69 Phanaeus (Phanaeus) amithaon Harold .......................................... 69 The Beltianus Group ........................................................... 71 Key to the Species of the Beltianus Group. .. 71 P hanaeus (P hanaeus) beltianus Bates . .. 72 Phanaeus (Phanaeus) howdeni Arnaud ........................................... 73 Phanaeus (Phanaeus) sallei Harold .............................................. 73 The Amethystinus Group ........................................................ 74 Key to the Species and Subspecies of the Amethystinus Group ....................... 74 Phanaeus (Phanaeus) melampus Harold .......................................... 74 Phanaeus (Phanaeus) amethystinus Harold ....................................... 77 The Quadridens Group . .. 79 Key to the Species of the Quadridens Group. .. 79 Phanaeus (Phanaeus) palliatus Sturm. .. 79 Phanaeus (Phanaeus) damocles Harold .......................................... 81 Phanaeus (Phanaeus) quadridens (Say) ........................................... 82 The Vindex Group. .. 83 Key to the Species of the Vindex Group .......................................... 83 Phanaeus (Phanaeus) igneus Macleay ............................................ 85 Phanaeus (Phanaeus) difformis LeConte ......................................... 88 Phanaeus (Phanaeus) vindex Macleay ........................................... 89 I ncertae Sedis .. .. 91 Phanaeus labreae (Pierce) . .. 91 Phanaeus(?) antiquus Horn ...................................................... 92 ACKNOWLEDGMENTS ............................................................. 92 LITERATURE CITED. .. 93 APPENDIX. .. 99 2. Contributions in Science,