WHITE GRUBS AND THEIR ALLIES A Study of North American Scarabaeoid Larvae NUMBER FOUR : ENTOMOLOGY }``` ` .f -' eta STUDIES IN i, BY PAUL O. RITGHER Corvallis, Oregon OREGON STATE UNIVERSITY PRESS .- OREGON STATE MONOGRAPHS STUDIES IN ENTOMOLOGY JoHN D. LATTIN, Consulting Editor NUMBER ONE A Review of the Genus Eucerceris (Hymenoptera: Sphecidae) By HERMAN A. SCULLEN NUMBER TWO The Scolytoidea of the Northwest: Oregon, Washington, Idaho, and British Columbia By W. J. CHAMBERLAIN NUMBER THREE Stonefíies of the Pacific Northwest By STANLEY G. JEWITT, JR. NUMBER FOUR White Grubs and Their Allies By PAUL O. RITCHER © 1966 Oregon State University Press Library of Congress Catalog Card number: 66 -63008 Printed in the United States of America By the Department of Printing, Oregon State University Author's Acknowledgments THE INFORMATION published in this book represents Mrs. Patricia Vaurie, American Museum of Natural work done over the past thirty years while the History ; Bernard Benesh, Sunbright, Tennessee; E. C. writer was on the staffs of the Kentucky Agricul- Cole, University of Tennessee; W. A. Price, the late tural Experiment Station (1936- 1949), North Carolina H. H. Jewett, L. H. Townsend, and other members of State College (1949- 1952), and Oregon State Univer- the Kentucky Department of Entomology and Botany; sity (1952 -1966). I am especially indebted to the Ken- J. D. Lattin, Louis Gentner, and other entomologists at tucky Agricultural Experiment Station for permission Oregon State University; D. Elmo Hardy, University to reproduce much of the material contained in my Ken- of Hawaii ; W. F. Barr of the University of Idaho; tucky Bulletins 401, 442, 467, 471, 476, 477, 506, and Joe Schuh of Klamath Falls, Oregon; Kenneth Fender 537, which have long been out of print. and Dorothy McKay- Fender of McMinnville, Oregon; For loan or gift of specimens, for identification of William Gibson of the Rockefeller Foundation ; Ferd adults, for advice or for other favors in connection with Butt, Lopez Island, Washington ; Roger Friend and J. these studies, I am greatly indebted to the following Peter Johnson, Connecticut Agricultural Experiment H. J. people in this country : W. H. Anderson, the late A. G. Station ; C. B. Dominick, Chatham, Virginia; Böving, E. A. Chapin, O. L. Cartwright, and D. M. Franklin, Massachusetts Agricultural Experiment Sta- \, Anderson of the United States National Museum ; the tion ; George Gyrisco and Henry Dietrich, Cornell Uni- ti late T. R. Chamberlin of the Madison, Wisconsin, versity; C. E. Heit, New York ; Vincent Roth, Ameri- .. White Grub Laboratory; E. M. Searles and the late can Museum of Natural History, Southwest Research C. L. Fluke of the University of Wisconsin ; H. H. Ross Station; Robert Lauderdale, University of Nevada ; Ray and M. W. Sanderson of the Illinois State Natural His- Hutson, Michigan State University; H. R. Bryson, ; formerly at , tory Survey; J. J. Davis of Purdue University; the late Kansas State University Vernon Olney, Philip Luginbill of the Federal Bureau of Entomology The Dalles, Oregon, Mid -Columbia Experiment Sta- and Plant Quarantine; W. P. Hayes of the University tion ; Frank Beer of the General Science Department at ,. óf Illinois ; L. P. Wehrle, Floyd Werner, and George Oregon State University; Forrest Peifer of Hood utler of the University of Arizona ; C. H. Hadley of River, Oregon, and a host of others. the Federal Japanese Beetle Laboratory ; C. F. Smith, T. B. Mitchell, and the late B. B. Fulton of North Caro- Workers abroad to whom I am indebted for mate- lina State College ; H. G. Reinhard of the Texas Agri- rial or other help pertaining to this study include the cultural Experiment Station ; C. H. Hoffman, United late Fritz van Emden, the late Gilbert Arrow and E. B. States Department of Agriculture ; Melville Hatch, Uni- Britton of the British Museum of Natural History ; versity of Washington; Hugh Leech and E. S. Ross of E. S. Narayanan and T. V. Venkatraman of the Indian the California Academy of Science ; Larry Saylor, Los Agricultural Research Institute, New Delhi, India; Gatos, California; E. Gorton Linsley, Ray Smith, John Philip Carne of the C.S.I.R.O., Canberra, Australia; MacSwain, and Paul Hurd of the University of Califor- Niilo Virkki of Finland ; R. A. Crowson, University of nia; Mont Cazier of Arizona State College ; Robert Potts Glasgow; and Kenneth Rachie, of the Rockefeller and Peter Ting, California Department of Agriculture; Foundation, New Delhi, India. 3 A number of former graduate students have given The writer is especially indebted to his two capable me a great deal of help with my studies and their as- assistants, Charles Baker and Mrs. Nandini Raj adhy- sistance is gratefully acknowledged. These include the aksha, who have assisted in many ways. Most of the following, all of whom completed a thesis on some phase Figures on Plates IV to VII, XIV, XV, XX, XXXI, dealing with Scarabaeidae : Henry Howden, Science and XXXVI to XLI were drawn by Mrs. Rajadhyak- Service, Ottawa, Canada (biology and taxonomy of sha. Geotrupinae, and so forth) ; Manohar Jerath, Nigeria The entire manuscript has been reviewed critically (Aphodiinae); Sutharm Areekul, Thailand (compara- by John D. Lattin of Oregon State University and tive morphology of the alimentary canal and nervous Henry Howden of the Canadian Science Service, re- system of Scarabaeidae) ; Aphirat Arunin, Thailand sulting in many improvements and additions. (comparative musculature of scarabaeoid larvae) ; and Financial support from the Kentucky Agricultural the late Floyd Ellertson, Mid -Columbia Experiment Experiment Station, the Oregon Agricultural Experi- Station, Hood River, Oregon (biology of Pleocoma). ment Station, the General Research Fund of Oregon The assistance of Gerald Kraft, Western Washing- State University, and the National Science Foundation ton State College, Bellingham; Everett Burts, United (Grant NSF- G17935) is gratefully acknowledged. States Department of Agriculture, Wenatchee, Wash- ington; and David Fellin, United States Forest Service, Paul O. Ritcher Missoula, Montana, is also acknowledged. September 1965 4 Contents Introduction 7 Subfamily Troginae 71 Superfamily Scarabaeoidea, Key to the Families 11 Subfamily Melolonthinae 75 Family Scarabaeidae, Key to the Subfamilies 11 Subfamily Rutelinae 100 Subfamily Scarabaeinae (Coprinae) 13 Subfamily Dynastinae 128 Subfamily Aphodiinae 26 Subfamily Cetoniinae 161 Subfamily Hybosorinae 37 Family Lucanidae 185 Subfamily Geotrupinae 41 Family Passalidae 199 Subfamily Pleocominae 57 Bibliography 207 Subfamily Glaphyrinae 62 Index 215 Subfamily Acanthocerinae 67 FRONTISPIECE: Third-stage larva of Phyllophaga hirticula. List of Plates Plate 1, Figures 1 -20 20 -21 Plate VII, Figures 77 -88 38 -39 Plate II, Figures 21 -23 22 -23 Plate VIII, Figures 89 -97 48 -49 Plate III, Figures 34 -46 24 -25 Plate IX, Figures 98 -109 50 -51 Plate IV, Figures 47 -55 30 -31 Plate X, Figures 110 -124 52 -53 Plate V, Figures 56 -57 32-33 Plate XI, Figures 125 -136 54-55 Plate VI, Figures 68 -76 34 -35 Plate XII, Figures 137 -146 60 -61 5 Plate XIII, Figures 147- 155 64 -65 Plate XXVIII, Figures 332 -357 148 -149 Plate XIV, Figures 156 -165 68 -69 Plate XXIX, Figures 358 -365 150 -151 Plate XV, Figures 166 -175 72 -73 Plate XXX, Figures 366 -374 152 -153 Plate XVI, Figures 176 -189 90 -91 Plate XXXI, Figures 375 -381 154 -155 Plate XVII, Figures 190 -204 92 -93 Plate XXXII, Figures 382 -389 156 -157 Plate XVIII, Figures 205 -216 94 -95 Plate XXXIII, Figures 390 -396 158 -159 Plate XIX, Figures 217 -227 96 -97 Plate XXXIV, Figures 397 -424 178 -179 Plate XX, Figures 228 -235 98 -99 Plate XXXV, Figures 425 -440 180 -181 Plate XXI, Figures 236 -248 116 -117 Plate XXXVI, Figures 441 -451 182 -183 Plate XXII, Figures 249 -258 118 -119 Plate XXXVII, Figures 452 -461 190 -191 Plate XXIII, Figures 259 -273 120 -121 Plate XXXVIII, Figures 462 -468 192 -193 Plate XXIV, Figures 274 -282 122 -123 Plate XXXIX, Figures 469 -478 194 -195 Plate XXV, Figures 238 -302 124 -125 Plate XL, Figures 479 -487 196 -197 Plate XXVI, Figures 303 -316 126 -127 Plate XLI, Figures 488 -495 202 -203 Plate XXVII, Figures 317 -331 146 -147 Plate XLII, Figures 496 -501 204 -205 6 Introduction WRITE GRUBS" is the common name applied in inconspicuous brown or black color ; but some are many countries to larvae of beetles belonging to bright green, blue, or yellow, with metallic lustre. Many the family Scarabaeidae, and in particular to those of species are nocturnal and rarely seen by the average economic importance to agricultural crops. While most person except when the beetles are attracted to light. "white grubs" are white with yellowish or reddish Others are diurnal and may be found on flowers or brown heads, some have orange, yellow, or bluish overripe fruits. bodies. The superfamily Scarabaeoidea includes three fam- General biology. In general, grubs of Scarabaeidae, ilies, the Scarabaeidae, Lucanidae, and Passalidae. Ac- Lucanidae stout and of as well, are curled (C- shaped), cording to Peterson (1951) about 1,250 species occur bodied, and have three pairs of well- developed legs. in the United States and Canada. The bulk of our spe- Passalidae, and have Grubs of however, are elongate cies belong to the family Scarabaeidae. only two pairs of well-developed legs. The head has stout, downward projecting (hypognathous) mandibles The Lucanidae and Passalidae inhabit decaying and a pair of three- or four -segmented antennae. There wood. The family Scarabaeidae, which includes four- are three thoracic segments and ten abdominal seg- teen commonly recognized subfamilies, has often been ments. The thorax and the first eight abdominal seg- divided into the lamellicorn scavengers and the lamelli- ments bear spiracles on each side which are usually C- corn leaf chafers (Comstock, 1950). This is a generali- shaped or kidney shaped.