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United StatesDepartment ofAgriculture

AgriculturalResearchService

Technical

BulletinNumber 1912

November 2004

Handbook of the

Bruchidae of the United

States and Canada(Insecta, Coleoptera)

John M. Kingsolver

Volume I

Kingsolver was research entomologist, SystematicEntomology Laboratory, PSI, Agricultural Research

Service, U.S. Department of Agriculture. He is

presently research associate with the Florida State

Collection of Arthropods.


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Abstract

Kingsolver, John M. 2004. Handbook ofthe Bruchidae of the United States andCanada (Insecta, Coleoptera). U.S. Depart-ment of Agriculture, Technical Bulletin

1912, 2 vol., 636 pp.Distinguishing characteristics and diag-nostic keys are given for the 5 subfami-lies, 24 genera, and 156 species of theseed beetle family Bruchidae of the Unit-ed States and Canada (including Hawaii).

Associated data for each species descrip-tion include a history of the name, syn-onymical names, type specimen infor-mation, geographical distribution, hostplants, and parasitoids. Appendices givespecies, host, fossil, and parasitoid lists

as well as a glossary of terms and a bibli-ography.

Bruchidae are found on every major landmass except Antarctica and New Zea-land. Eggs are usually laid on the seed orfruit of a plant suitable for developmentof the larva. Immature stages are spentinside seeds that have been excavated bylarval feeding. Adults live free and feedon pollen and nectar.

Johnson (1970) estimates that approxi-

mately 84 percent of the known hosts ofBruchidae are in the plant family Legu-minosae. The remaining hosts are scat-tered among 31 other families. Sixteenplant families support larval feeding inthe United States and Canada. Severalspecies of Bruchidae, especially those

with a cosmopolitan distribution, are no-torious pests of stored leguminous seeds.

Keywords: Coleoptera, Bruchidae, Am-blycerinae, Bruchidiinae, Bruchinae, Ky-

torhininae, Megacerinae, United States,Canada, Hawaii, taxonomy, insect-plantinteractions.

This is part of a series of studies ofbruchid genera contributing to a com-prehensive database for this importantseed-feeding beetle family in the Western

Hemisphere. It provides the means toidentify these insects for taxonomists,students, museum curators, biodiver-sity workers, port identifiers, and ecolo-gists conducting studies in rangeland,pasture, and forest management in theUnited States and Canada.

Mention of commercial products in thispublication is solely for the purpose ofproviding specific information and doesnot imply recommendation or endorse-ment by the U.S. Department of Agricul-ture over others not mentioned.

While supplies last, single copies ofthis publication can be obtained at nocost from John M. Kingsolver, FloridaState Collection of Arthropods, P.O. Box147100, Gainesville, FL 326147100.

Copies of this publication may be pur-chased from the National Technical In-formation Service, 5285 Port Royal Road,Springfield, VA 22161. ARS has no addi-tional copies for free distribution.

November 2004

The U.S. Department of Agriculture (USDA)

prohibits discrimination in all its programs and

activities on the basis of race, color, national

origin, age, disability, and where applicable, sex,

marital status, familial status, parental status,religion, sexual orientation, genetic information,

political beliefs, reprisal, or because all or part of

an individuals income is derived from any public

assistance program. (Not all prohibited bases

apply to all programs.) Persons with disabilities

who require alternative means for communication

of program information (Braille, large print,

audiotape, etc.) should contact USDAs TARGET

Center at (202) 720-2600 (voice and TDD). To

file a complaint of discrimination, write to USDA,

Director, Office of Civil Rights, 1400 Independ-

ence Avenue, S.W., Washington, D.C. 20250-9410, or call (800) 795-3272 (voice) or (202) 720-

6382 (TDD). USDA is an equal opportunity

provider and employer.

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Contents

Volume 1

Acknowledgments ............................................................................................... ix

Abbreviations ....................................................................................................... x

Introduction ......................................................................................................... 1

Importance of the Bruchidae .....................................................................................2

Biology of the Bruchidae ........................................................................................... 3

Morphology of the Bruchidae.....................................................................................8

Materials and Methods ............................................................................................ 16

Taxonomy of the Bruchidae of the United States and Canada ............................. 19

Key to the Bruchidae of the United States and Canada, Including Hawaii ................ 21

Subfamily Pachymerinae Bridwell ....................................................................... 24

Tribe Caryedontini Bridwell ..................................................................................... 24Genus CaryedonSchoenherr ............................................................................... 24

Caryedon serratus(Olivier) ............................................................................... 24

Tribe Pachymerini Bridwell ..................................................................................... 26

Genus CaryobruchusBridwell .............................................................................. 26

Caryobruchus gleditsiae (Linnaeus)................................................................... 26

Subfamily Kytorhininae Bridwell ........................................................................ 28

Genus KytorhinusFischer von Waldheim ............................................................. 28

Kytorhinus prolixus(Fall) .................................................................................. 28

Subfamily Amblycerinae Bridwell ........................................................................ 30

Tribe Amblycerini Bridwell ...................................................................................... 30

GenusAmblycerusThunberg ............................................................................... 30

Amblycerus eustrophoides(Schaeffer) ............................................................... 32

Amblycerus ireriaeRomero, Johnson, and Kingsolver ....................................... 33

Amblycerus nigromarginatus(Motschulsky)....................................................... 34

Amblycerus obscurus(Sharp) ............................................................................ 34

Amblycerus robiniae (Fabricius) ........................................................................ 35

Amblycerus schwarziKingsolver ....................................................................... 37

Amblycerus vitis(Schaeffer) .............................................................................. 38

Tribe Spermophagini Borowiec ................................................................................ 39

Genus ZabrotesHorn ........................................................................................... 39

Zabrotes amplissimusKingsolver ...................................................................... 42

Zabrotes arenarius(Wolcott).............................................................................. 42

Zabrotes bexarensisKingsolver ......................................................................... 43

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Zabrotes chandleriKingsolver ........................................................................... 44

Zabrotes chavesiKingsolver .............................................................................. 44

Zabrotes crucigerHorn...................................................................................... 45

Zabrotes cynthiaeKingsolver ............................................................................ 46

Zabrotes densusHorn. ..................................................................................... 47

Zabrotes eldenensisKingsolver. ........................................................................ 47Zabrotes humboldtaeKingsolver ....................................................................... 48

Zabrotes obliteratusHorn ................................................................................. 49

Zabrotes planifronsHorn .................................................................................. 50

Zabrotes spectabilisHorn ................................................................................. 50

Zabrotes stephaniKingsolver ............................................................................ 51

Zabrotes subfasciatus(Boheman) ..................................................................... 52

Zabrotes subnitensHorn ................................................................................... 55

Zabrotes sylvestrisRomero and Johnson .......................................................... 55

Zabrotes victoriensisKingsolver ........................................................................ 56

Subfamily Bruchinae Pic ..................................................................................... 58

Tribe Megacerini Bridwell ........................................................................................ 58

Genus MegacerusFahraeus ................................................................................. 58

Megacerus (Pachybruchus) coryphae(Olivier) .................................................... 59

Megacerus (Megacerus) cubiculus(Casey) .......................................................... 60

Megacerus (Megacerus) cubicus(Motschulsky) .................................................. 61

Megacerus (Megacerus) discoidus(Say) ............................................................. 63

Megacerus (Megacerus) impiger(Horn) .............................................................. 64

Megacerus (Pachybruchus) leucospilus(Sharp) .................................................. 65

Megacerus (Serratibruchus) maculiventris(Fahraeus)......................................... 66

Megacerus (Megacerus) ripiphorus(Fahraeus).................................................... 68

Megacerus (Serratibruchus) schaefferianusBridwell .......................................... 68

Tribe Bruchini Bridwell ........................................................................................... 69

Genus BruchusLinnaeus ..................................................................................... 69

Bruchus brachialisFahraeus ............................................................................ 70

Bruchus pisorum(Linnaeus) ............................................................................. 72Bruchus rufimanusBoheman............................................................................ 74

Tribe Bruchidiini Bridwell ....................................................................................... 75

Genus BorowieciusAnton ....................................................................................75

Borowieciusademptus(Sharp) .......................................................................... 75

Genus BruchidiusSchilsky .................................................................................. 76

Bruchidius cisti(Fabricius)................................................................................ 77

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Bruchidius villosus(Fabricius) .......................................................................... 78

Genus CallosobruchusPic .................................................................................... 79

Callosobruchus chinensis(Linnaeus) ................................................................. 80

Callosobruchus maculatus(Fabricius) ............................................................... 82

Callosobruchus phaseoli(Gyllenhal) .................................................................. 86

Callosobruchus pulcherPic ............................................................................... 87Tribe Acanthoscelidini Bridwell ............................................................................... 88

GenusAbutiloneusBridwell ................................................................................. 88

Abutiloneus idoneusBridwell ............................................................................ 88

GenusAcanthoscelidesSchilsky .......................................................................... 89

Acanthoscelides aequalis(Sharp) ...................................................................... 96

Acanthoscelides alboscutellatus(Horn) ............................................................. 97

Acanthoscelides atomus(Fall) ........................................................................... 98

Acanthoscelides aureolus(Horn) complex .......................................................... 99

Acanthoscelides baboquivariJohnson ............................................................. 101

Acanthoscelides bisignatus(Horn) ................................................................... 102

Acanthoscelides biustulus(Fall) ...................................................................... 103

Acanthoscelides calvus(Horn) ......................................................................... 104

Acanthoscelides chiricahuae(Fall) ................................................................... 105

Acanthoscelides compressicornis(Schaeffer) .................................................... 106

Acanthoscelides comstockiJohnson ................................................................ 107

Acanthoscelides daleaeJohnson .................................................................... 108

Acanthoscelides desmanthiJohnson ............................................................... 109

Acanthoscelides distinguendus(Horn) ............................................................. 109

Acanthoscelides flavescens(Fahraeus) ............................................................ 111

Acanthoscelides floridae(Horn) ....................................................................... 112

Acanthoscelides fraterculus (Horn) .................................................................. 113

Acanthoscelides fumatus(Schaeffer) ............................................................... 114

Acanthoscelides griseolus(Fall). ...................................................................... 115

Acanthoscelides helianthemumBottimer ......................................................... 116

Acanthoscelides herissantitusJohnson ........................................................... 117Acanthoscelides inquisitus(Fall) ...................................................................... 118

Acanthoscelides kingsolveriJohnson .............................................................. 119

Acanthoscelides lobatus(Fall) ......................................................................... 120

Acanthoscelides longistilus(Horn). .................................................................. 121

Acanthoscelides macrophthalmus(Schaeffer) ................................................... 122

Acanthoscelides margaretaeJohnson. ............................................................ 123

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Acanthoscelides mixtus(Horn) ........................................................................ 124

Acanthoscelides modestus(Sharp) .................................................................. 125

Acanthoscelides mundulus(Sharp).................................................................. 126

Acanthoscelides napensisJohnson ................................................................. 127

Acanthoscelides obrienorumJohnson.............................................................. 128

Acanthoscelides obsoletus(Say) ...................................................................... 129Acanthoscelides obtectus(Say) ........................................................................ 130

Acanthoscelides oregonensisJohnson ............................................................. 134

Acanthoscelides pallidipennis(Motschulsky) ................................................... 135

Acanthoscelides pauperculus(LeConte) ........................................................... 136

Acanthoscelides pectoralis(Horn) .................................................................... 137

Acanthoscelides pedicularius(Sharp) .............................................................. 138

Acanthoscelides perforatus(Horn) ................................................................... 139

Acanthoscelides prosopoides(Schaeffer) .......................................................... 140

Acanthoscelides pullus(Fall) ........................................................................... 142

Acanthoscelides pusillimus(Sharp) ................................................................. 143

Acanthoscelides quadridentatus(Schaeffer) ..................................................... 144

Acanthoscelides rufovittatus(Schaeffer) .......................................................... 145

Acanthoscelides schaefferi(Pic) ....................................................................... 146

Acanthoscelides schrankiae(Horn) ................................................................. 147

Acanthoscelides seminulum(Horn) .................................................................. 148

Acanthoscelides speciosus(Schaeffer) ............................................................. 149

Acanthoscelides stylifer(Sharp) ...................................................................... 150

Acanthoscelides subaequalisJohnson ............................................................ 151

Acanthoscelides submuticus(Sharp) ............................................................... 152

Acanthoscelides tenuisBottimer ..................................................................... 154

Acanthoscelides tridenticulatusBottimer ......................................................... 155

GenusAlgarobiusBridwell ................................................................................. 155

Algarobius bottimeriKingsolver ....................................................................... 156

Algarobius prosopis(LeConte) ......................................................................... 157

GenusAlthaeusBridwell .................................................................................... 158Althaeus folkertsiKingsolver ........................................................................... 158

Althaeus hibisci(Olivier) ................................................................................. 160

Althaeus steineriKingsolver ............................................................................ 161

Genus CaryedesHummel .................................................................................. 163

Caryedes helvinus(Motschulsky) .................................................................... 164

Caryedes incensus(Sharp) ............................................................................. 165

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Genus GibbobruchusPic .................................................................................... 166

Gibbobruchus cristicollis(Sharp) ..................................................................... 167

Gibbobruchus divaricataeWhitehead and Kingsolver....................................... 168

Gibbobruchus mimus (Say) .............................................................................. 169

Genus LithraeusBridwell. .................................................................................. 170

Lithraeus atronotatus(Pic) .............................................................................. 171Genus MeibomeusBridwell ................................................................................ 172

Meibomeus desmoportheusKingsolver and Whitehead .................................... 172

Meibomeus musculus(Say) ............................................................................. 173

Meibomeus surrubresus(Pic) ........................................................................... 174

Genus MerobruchusBridwell .............................................................................. 175

Merobruchus insolitus(Sharp) ......................................................................... 176

Merobruchus julianus(Horn) ........................................................................... 177

Merobruchus knulli(White) .............................................................................. 179

Merobruchus lysilomaeKingsolver .................................................................. 180

Merobruchus major(Fall) ................................................................................. 181

Merobruchus placidus(Horn) .......................................................................... 182

Merobruchus teraniKingsolver ........................................................................ 183

Merobruchus vacillator(Sharp). ....................................................................... 184

Genus MimosestesBridwell ............................................................................... 185

Mimosestes acaciestesKingsolver and Johnson .............................................. 186

Mimosestes amicus(Horn) .............................................................................. 187

Mimosestes insularisKingsolver and Johnson................................................. 188

Mimosestes mimosae(Fabricius) ..................................................................... 189

Mimosestes nubigens(Motschulsky)................................................................ 191

Mimosestes protractus(Horn) .......................................................................... 192

Mimosestes ulkei (Horn) ................................................................................. 193

Genus NeltumiusBridwell .................................................................................. 194

Neltumius arizonensis(Schaeffer). ................................................................... 195

Neltumius gibbithorax(Schaeffer) .................................................................... 196

Neltumius texanus(Schaeffer) ......................................................................... 197Genus SenniusBridwell ..................................................................................... 198

Sennius abbreviatus(Say) ............................................................................... 200

Sennius cruentatus(Horn) .............................................................................. 201

Sennius discolor(Horn) ................................................................................... 202

Sennius fallax(Boheman) ............................................................................... 203

Sennius lebasi(Fahraeus) ............................................................................... 205

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Appendix I. Hawaiian Bruchidae ....................................................................... 223

Key to Hawaiian Bruchidae ................................................................................... 224

Host Associations of Hawaiian Bruchidae .............................................................. 225

Appendix II. Synonymical List of the Bruchidae of theUnited States and Canada ................................................................................. 227

Extant Species ...................................................................................................... 227

Extinct Species ..................................................................................................... 237

Appendix III. Hosts of the Bruchidae of the United States andCanada, by Bruchid .......................................................................................... 239

Appendix IV. Hosts of the Bruchidae of the United States andCanada, by Host Plant ...................................................................................... 246

Appendix V. Natural Enemies of Bruchidae of the United States and

Canada ............................................................................................................. 260Glossary of Morphological Terms ...................................................................... 264

References ....................................................................................................... 269

Index ................................................................................................................ 319

Volume 2 (Illustrations)

Volume 2 contains all figures referenced in Volume 1.

Sennius leucostaurosJohnson and Kingsolver ................................................ 206

Sennius medialis(Sharp) ................................................................................ 206

Sennius morosus(Sharp) ................................................................................ 207

Sennius obesulus(Sharp) ............................................................................... 208

Sennius simulans(Schaeffer) .......................................................................... 209

Sennius whiteiJohnson and Kingsolver .......................................................... 210Genus StatorBridwell ........................................................................................ 211

Stator bealiJohnson....................................................................................... 212

Stator bottimeriKingsolver .............................................................................. 213

Stator chihuahuaJohnson and Kingsolver ...................................................... 214

Stator coconinoJohnson and Kingsolver ......................................................... 214

Stator limbatus(Horn) .................................................................................... 215

Stator pruininus(Horn) ................................................................................... 216

Stator pygidialis(Schaeffer). ........................................................................... 218

Stator sordidus(Horn) .................................................................................... 219

Stator subaeneus(Schaeffer) ........................................................................... 219

Stator vachelliaeBottimer ............................................................................... 220

Genus StylantheusBridwell ............................................................................... 221

Stylantheus macrocerus(Horn) ....................................................................... 221

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Acknowledgments

I gratefully acknowledge the assistance andsuggestions of the following individuals

who read or commented on the manu-script; C.D. Johnson, G.S. Pfaffenberger,

D.R. Miller, R.D. Gordon, the late D.R.Whitehead, the Systematic EntomologyLaboratory hymenopterists, and my late

wife, Cynthia, who assembled the host andparasite lists and the index.

Specimens were loaned for this study bythe following institutions and individuals:

American Museum of Natural History, NewYork; Arizona State University, Tempe, AZ;Charles Bellamy, Escondido, CA; Bernice P.Bishop Museum, Honolulu; British Muse-um of Natural History, London; California

Academy of Sciences, San Francisco; Cali-fornia Department of Agriculture, Sacra-mento, CA; Canadian National Collections,Ottawa, ON; Carnegie Museum, Pitts-

burgh; Donald S. Chandler, Durham, NH;Cornell University, Ithaca, NY; Field Mu-seum of Natural History, Chicago; FloridaState Collections, Gainesville, FL; H.F. &

A.T. Howden, Ottawa, ON; Illinois Natural

History Survey, Urbana, IL; Michael Ivie,Bozeman, MT; C.D. Johnson, Flagstaff, AZ;Los Angeles County Museum, Los Angeles;Michigan State University, East Lansing,

MI; Museum of Comparative Zoology, Cam-bridge, MA; Museum National dHistoireNaturelle, Paris; C.W. OBrien, Tallahas-see, FL; Naturhistoriska Riksmuseet,Stockholm, Sweden; Ohio State University,Columbus, OH; Purdue University, WestLafayette, IN; Karl Stephan, Red Oak, OK;

Texas A&M University, College Station, TX;Robert H. Turnbow, Ft. Rucker, AL; Na-tional Museum of Natural History, Wash-ington, DC; University of Arizona, Tucson,

AZ; University of California, Berkeley, CA;

University of California, Davis, CA; Univer-sity of Kansas, Lawrence, KS; University ofMichigan, Ann Arbor, MI; University ofMissouri, Columbia, MO; University Zoo-logical Museum, Copenhagen, Denmark;Zoological Museum, University of Moscow,Russia; Utah State University, Logan, UT.

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Abbreviations

Type Specimen Depositories

BMNH: British Museum (Natural History), London

CASC: California Academy of Sciences, San Francisco

CNCI: Canadian National Collections, Ottawa, ON

FMNH: Field Museum of Natural History, Chicago

ICCM: Carnegie Museum, Pittsburgh

MCZC: Museum of Comparative Zoology, Harvard University, Cambridge, MA

MNHP: Museum National dHistoire Natural, Paris

NHRS: Naturhistoriska Riksmuseet, Stockholm, Sweden

OSUC: Ohio State University Collections, Columbus, OH

PURC: Purdue University Collections, West Lafayette, IN

UCDC: University of California, Davis, CAUSNM: U.S. National Museum of Natural History, Washington, DC

UZMC: University Zoological Museum, Copenhagen, Denmark

ZMUM: Zoological Museum, University of Moscow, Russia

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NC North Carolina

ND North Dakota

NE Nebraska

NF Newfoundland

NH New Hampshire

NJ New Jersey

NM New Mexico

NS Nova Scotia

NT Northwest Territories

NV Nevada

NY New York

OH OhioOK Oklahoma

ON Ontario

OR Oregon

PA Pennsylvania

PE Prince Edward Island

PQ Quebec

RI Rhode Island

SC South Carolina

SD South Dakota

SK Saskatchewan

TN Tennessee

TX Texas

UT Utah

YT Yukon Territory

VA Virginia

VT Vermont

WA WashingtonWI Wisconsin

WV West Virginia

WY Wyoming

State and Province Abbreviations

AB Alberta

AK Alaska

AL Alabama

AR Arkansas

AZ Arizona

BC British Columbia

CA California

CO Colorado

CT Connecticut

DC District of Columbia

DE DelawareFL Florida

GA Georgia

HI Hawaii

IA Iowa

ID Idaho

IL Illinois

IN Indiana

KS Kansas

KY Kentucky

LA Louisiana

MA Massachusetts

MB Manitoba

MD Maryland

ME Maine

MI Michigan

MN Minnesota

MO Missouri

MS Mississippi

MT Montana

NB New Brunswick

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Introduction

The beetle family Bruchidae has beenrecognized for centuries for the ability ofcertain species to destroy stores of comes-tible leguminous seeds. About 30 speciesof bruchids in the world are serious pests,

and at least 9 are cosmopolitan as theresult of commercial activities. Bruchid lar-

vae feed entirely within seeds, making theirdetection and control difficult. The mostsuccessful species are those that attackstored seeds. Because they have adaptedto oviposit and develop in several hosts,they are inadvertently assisted by humansmobility and storage methods. Althoughthe larva is the destructive life stage, de-scriptions of this form are available foronly a few species. For most genera of

bruchids, identification is tedious and dif-ficult because many species are closely re-lated, individuals are small (average length1.06.0 mm), and dissection of genitalia isoften necessary for positive identification.

Bruchids are found naturally on all ma-jor land masses except New Zealand andAntarctica. More intense speciation hasoccurred in the tropical regions than in thetemperate, and fewer species are found intropical rain forests than in more xeric re-

gions. Small isolated islands usually havelittle or no bruchid fauna because estab-lishment of most bruchids depends on theprevious invasion and establishment ofsuitable food plants for oviposition.

No single information source exists for theBruchidae of the United States and Can-ada. The literature is scattered through a

wide range of publications generally un-available to most workers. Because many

bruchids are critically important pests ofstored legumes, their correct identificationis vital to effective control. The U.S. andCanadian literature was combed for dataon classification, host plant associations,geographical distributions, and parasitoidsfor this compilation. The world literature

was scanned for the several species of

cosmopolitan stored-product species in thegeneraAcanthoscelides, Bruchus, Calloso-bruchus, Caryedon,and Zabrotesto provideas broad a coverage as possible since mostof the biological studies on these species

were undertaken outside North America.

This handbook contains distinguishing

characterizations for 156 species in 24genera, plus generic and specific descrip-tions, synonymical names, references topapers on biology, and new or revisedtaxonomic keys for all subfamilies, genera,and species. New information on hosts anddistribution was recorded primarily fromspecimens deposited in the National Col-lection of Insects in Washington, DC, andfrom specimens loaned by many other U.S.(including Hawaiian) and Canadian col-lections. Included, however, were recordsfrom faunal lists such as Kirk (1969, 1970)for South Carolina and Kirk and Bals-

baugh (1975) for South Dakota. Approxi-mately 30,000 specimens were examinedduring 28 years study of the Bruchidae.Each species is illustrated with photo-graphs and line drawings to facilitate iden-tification. A separate key to the HawaiianBruchidae is included in appendix I. Otherappendices include a cross-referencedindex to host plants (III and IV), a list of

parasitoids (V), and a synonymical check-list (II). There is also a glossary of termsused in this manual. All of the elementsnecessary to produce a future annotatedcatalog of U.S. and Canadian Bruchidaeare included.

The geographical scope of the handbookis primarily the 50 United States, Canada,and the associated continental islands.Beyond these limits, only general distribu-tions are provided.

Although bruchids are difficult to identify,information presented in this handbookshould be useful to taxonomists, biologicalcontrol specialists, port identifiers, plantquarantine officers, museum curators,ecologists, students and teachers, andFederal and State entomologists.


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Importance of the Bruchidae

This family has received much attentionduring the past three decades for severalreasons. Demands for more food for theexpanding world population has focusedon the need for better control of stored-product insects, especially those attackingcomestible leguminous seeds in larders ofdeveloping countries. Storage losses can

be ruinous not only from initial damageby bruchid larvae but also from ensuinginvasions of other organisms (Howe 1973;

Yoshida 1990).

Morallo-Rejesus (1990) estimated thatin the Philippines, losses of mung beansinfested by Callosobruchus chinensis(Lin-naeus) ranged from 7.8 percent to 9.8percent at harvest, and without protection

by insecticides or proper storage post- har-vest losses rose to 80 to 100 percent within3 months. Warthen (1989) summarizedthe uses of neem (Azadirachta indicaAdr.

Juss.) as a control agent for bruchids instored grains.

Laboratories in Colombia, England,France, India, Japan, Nigeria, and theUnited States are investigating many as-pects of physiology, morphology, plant hostresistance, and pheromones in relation to

the principal cosmopolitan species of Bru-chidae. A voluminous literature on thesetarget species is being generated. South-gate (in Aitken 1975) summarized avenuesof dispersal of the major economic bruchidspecies.

Another group of bruchids presents apotential threat to the propagation of treestargeted for forest plantings in undevel-oped countries where firewood sourceshave been depleted. Seed stocks can be de-stroyed by bruchid larvae (Johnson 1983c;

Southgate 1983).

Biological Control

The accidental introduction of undesirableplants into certain parts of the world (Aus-tralia, Hawaii, Thailand) has stimulatedsearches for biological agents in the native

country of the plant that might assist incontrolling the spread of the pest species.Since larvae usually destroy the seeds,the feeding habits of Bruchidae reduce thepotential for reseeding. Other programs (forinstance, in Argentina) have attempted tointroduce foreign bruchids to help in con-

trolling a native plant that is invading graz-ing land, but native parasites destroyedthe introduced populations (Erb and Fras1983). Efforts to control the weedy shrubMimosa pigraLinnaeus usingAcanthos-celides quadridentatus(Schaeffer) andA.

puniceusJohnson are under way in Aus-tralia and Thailand (Kassulke et al. 1990;Forno et al. 1989; Cock and Evans 1984).

The effectiveness of bruchids as biocontrolagents needs further exploration. In South-east Asia and the southwest Pacific, experi-mental efforts are underway to control thetwo weedy leguminous plants Cassia toraLinnaeus and Senna obtusifoliaLinnaeus

by the Neotropical bruchid Sennius instabi-lis(Sharp) (Cock and Evans 1984).

Recent studies in the use of resistantvarieties of host plants as a means ofbiological control of harmful Bruchidaehave been conducted by Brewer and Horb-er (1984), Dobie (1981), Dobie et al. (1979),Fernandez and Talekar (1990), Horber

(1978), Kitamura et al. (1990), Minneyet al. (1990), Nwanze and Horber (1976),Osborn et al. (1988), and others.

Medical Importance

Bruchid adults and larvae feed only onplant tissue and are not associated withany disease-causing organisms of warm-

blooded animals. Instances of allergicreactions have been recorded, however.Chittenden (1912a) noted that handlers of

broad beans (Vicia fabaLinnaeus) infestedby Bruchus rufimanusBoheman developedirritating rashes from adults crawling ontheir skin. Wittich (1940) reported twocases in Minnesota of allergic reaction by

women who had been sorting stored beansand peas infested with Zabrotes subfas-


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4

Some concurrent evolutionary trends havebeen observed, however, that allow us tospeculate.

The Bruchidae is apparently a monophy-letic, spermatophagus, chrysomeloid groupspringing from an ancestor common to thechrysomelid subfamily Sagrinae (Crowson

1946; Monros 1955). Similarities in wingvenation, male genitalia, form of the meta-femur, presence of tibial spurs, internalfeeding by the larvae, and other characterspoint to a relatively recent divergence ofthe bruchids followed by development ofspermophagy to a fine degree (Kingsolver1995b; Verma and Saxena 1996).

Kingsolver (1995b) listed the characters hethought were sufficient to give the bru-chids full family status, and his view was

supported by Hawkeswood (1996) andVerma and Saxena (1996). Lawrence andNewton (1982) and Reid (1996), however,placed the bruchids as a subfamily of theChrysomelidae based on cladistic analy-sis. Schmitt (1989) argued that Bruchidaeshould properly be a subfamily within theChrysomelidae but that Bruchidae should

be retained with the -idaefamily endingfor stability and unambiguity of scientificcommunication. Monros (1955) treated theBruchidae as a subfamily of the Chrysome-

lidae.

The groups seemingly nearest to the Sagri-nae are in the bruchid subfamily Pachym-erinae, especially the genera Caryoborusand Caryobruchus.Although resemblanceof the sagrine Carpophagusto these pa-chymerines is striking, Crowson (1946) didnot think that this genus was in any wayrelated to the Bruchidae. Without know-ing what bruchid species he used in thiscomparison, I am of the opinion that Car-

pophagushas more in common with thepachymerine genera than with any otherchrysomeloid genus. Similarities are malegenitalia with a simple, curved, tubularmedian lobe, bases of the lateral lobesfused straplike with only the apices ex-panded; absence of a crop in the digestive

system (Kasap 1978b); swollen metafemurwith ventral denticles; metepisternum withan angular sulcus (also found in someprimitive Cerambycidae); similar wing

venation patterns including the presenceof a wedge cell in some species (Suzuki1969); larvae as internal feeders in plants

(sagrines in stems or crowns, pachymer-ines in seeds). Differences are mostlyexternal: Elytra throughout the familyBruchidae always striate whereas those ofCarpophagusand most other sagrines lackdistinct striae; frons in the pachymer ines

with a median carina, sagrines with anX-shaped sulcus; larval mandibles gouge-shaped in pachymerines, toothed in thosesagrines whose larvae are known. Maulik(1941) presented useful comparative char-acters for the Sagrinae.

Evolutionary Trends

Certain evolutionary trends are apparentwithin the Bruchidae. They indicate thatthe most likely primitive genera are in thePachymerinae and that the Rhaebinae,

which are found only in central Asia, areprobably a relict group. The Amblycerinae,Eubaptinae (found only in South America),and Kytorhininae are successively morespecialized, and the Acanthoscelidinae areprobably the most derived group.

1. Trochantins are present in the foreand middle legs in the Pachymerinae,

Amblycerinae, Eubaptinae (SouthAmerica), Rhaebinae (Asia), and Kyto-rhininae but are modified or lost in theBruchinae except in the mid legs ofMegacerini.

2. Median lobe of male genitalia has anunmodified apex in Pachymerinae andRhaebinae (figure 2) but is fracturedinto a separate ventral valve in all other

subfamilies (figure 3).

3. Ocular sinus is shallow in Pachymeri-nae (figure 1) and inAmblycerusof the

Amblycerinae (figure 4) but deep inmore specialized subfamilies. In somespecies of Meibomeus(figure 5), thesinus nearly divides the eye.


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4. Adult antennal forms range from sim-ple serrate or flattened moniliform inPachymerinae, Rhaebinae, and Eu-

baptinae to clavate, strongly serrate, orpectinate in males in the Kytorhininaeand in some species in Bruchinae (fig-ures 613).

5. Mesepimeron is gradually reduced byventral encroachment of mesepister-num (figures 2527) as the subfamiliesprogress in degree of specialization.

6. Venation of anal area of wing graduallylost the wedge cell and veins assumedan h shape in most of the morederived families.

7. Lateral pronotal carina modified or lostas specialization progressed.

Genetic Studies

Skaife (1925, 1926) was a pioneer in bru-chid genetics. Certain species of Bruchidaeare now being used in several laboratoriesthroughout the world as test organisms forgenetic studies because they are readilycultured in the laboratory and have shortlife cycles. Species used are those that

breed continuously in stored seedsAc-anthoscelides obtectus, Callo- sobruchuschinensisand C. maculatus,and Zabrotes

subfasciatus,for example.

Life Histories

For a general discussion of bruchid biolo-gies, see Skaife (1926), Southgate (1979),and Zacher (1929 and 1930). See also theProceedings of the Second InternationalSymposium on Bruchids and Legumes (Fu-

jii et al. 1990) for other life history studies.Johnson (1990b) contributed a paper tothis symposium.

Larval Feeding

A first larval instar is faced not only withescaping from the egg but also with pen-etrating one or two layers of plant integu-ment before it can reach its food source.During eclosion, larvae of most species

bore directly through the floor layer of

the egg, through the pod or seed integu-ment, through the endocarp, and into thecotyledon, whereas others cut throughthe top of the egg and walk around on thesubstrate searching for a crack or creviceto help them gain purchase to bore intothe seed. Some species are provided with

a suckerlike appendage on the tenth ab-dominal segment to help them adhere tothe surface during their search for a suit-able entry site (Pfaffenberger and Johnson1976). Special adaptations of the neonatelarva include well-developed legs in thosespecies that emerge from the top of the eggand shorter legs in those forms that drilldirectly through the floor of the egg.

The most remarkable character, however,is a toothed dorsal prothoracic plate thatassists the larva in escaping from the eggand gives leverage in penetrating the podor seed. All bruchid species for which thefirst larval instar is known possess thistoothed plate (figure 16). Pfaffenberger and

Johnson (1976) and Kannan (1923) illus-trated this character for several species. Inits first molt, the larva loses these specialadaptations and assumes a scarabaeiformaspect with legs much reduced and withgouge-shaped mandibles. It feeds andmolts through its remaining three instars

inside an excavation scarcely larger thanits own bulk.

In most cases, the larva, immediatelybefore entering the pupal stage, will boreto the surface of the seed or pod, leav-ing a circular cap in the epidermis thatcan easily be cut and pushed off by theadult during emergence. After excavatingthis escape route, the larva returns to thefeeding chamber and pupates. In somespecies, however, pupation occurs partlyor completely outside individual seeds (for

example, Caryedon serratus).Species toolarge to develop in individual seeds mayfeed on several seeds inside an envelope(the pod or capsule, for example) and spina cocoon inside the envelope (examplesinclude some species ofAmblycerusand ofSennius). Glands producing the materialfor the cocoon have not been investigated.


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Adult Nutrition

Although bruchid larvae are well known tosubsist mainly on tissue of the seed coty-ledon, comparatively little is known aboutadult food habits for most species. Mostobservations on adult feeding have beenmade on the species affecting cropsthe

so-called economic species. Adult man-dibles with sharp medial edges and lack ofteeth are not especially adapted for bitingtissue except to complete cutting of theoperculum of the seed prior to emergence.Maxillae in many species are equipped

with well developed, feathery setae that as-sist in raking pollen into the oral cavity.

Most so-called host records for adults,usually floristic associations with what-ever plant happens to be in bloom at the

time the adults are flying about, may havelittle or no relationship with the larval hostplant. Because adults cause little dam-age by their food intake, it has been pre-sumed that either they do not feed or theyfeed sparingly on nectar or pollen. Female

Acanthoscelides obtectushave been ob-served biting the sides of green pods or thesuture of more mature pods of Phaseolusto make an opening through which theydeposit eggs into the interior of the pods.

Whether or not they ingest tissue is not

known (Slingerland 1892). Brindley et al.(1946, 1952) and Pajni and Sood (1975)reported that Bruchus pisorumcan onlyreproduce after ingesting pollen of its larvalhost, the pea plant (Pisum sativum).Adultsof Bruchus pisorumoverwinter in trash orunder bark and emerge from diapause tofeed on pea blossoms; then they are readyto oviposit on green pods.

De Luca (1966) compiled a list of flower-visiting records from the world literature.

No descriptions of actual ingestion weremade, although his introduction to thepaper is a discussion of pollen and nectarnutrition. Johnson (1977d) listed eightspecies of plants from which adultAcan-thoscelides pauperculuswere collected,

but precise observations on actual feed-ing were not made. He suggested thatfeeding on pollen is necessary for mating

and oviposition. Lago and Mann (1987)recorded several species of bruchids oninflorescences of Daucus carotaLinnaeus(wild carrot).

Jarry (1987) identified the pollen of 18different species of plants in the diges-tive tracts ofAcanthoscelides obtectusin

France. Pollen from the principal host,Phaseolus vulgaris,made up only 9 percentof the total, whereas weedy plants in areassurrounding the plots of P. vulgarismadeup the bulk of pollen identified. Grass pol-len was well represented.

Adults ofAcanthoscelides obtectuscanreproduce and oviposit without first tak-ing food, but Leroi (1978) determined that

well-fed, newly emerged individuals cansurvive more than 200 days on honey-pol-

len mixtures. Ovarian production by well-fed females was 50 percent more than that

by starved females. Shinoda and Yoshida(1987a) found that when newly emergedCallosobruchus chinensiswere fed onpowdery mildew (Sphaerotheca fuliginea)orrust (Uromyces azukiola)their mean lon-gevity was three times that of control indi-

viduals and the numbers of eggs produceddoubled. Zacher (1929) thought that adultsshould be able to retain sufficient fat fromthe larval stages to carry them through

adult life.

Larson and Fisher (1938) included a sum-mary of the literature on adult feeding tothat date. Many of the observations weremade on caged bruchids and do not neces-sarily reflect field conditions. Slingerland(1892) claimed that bean weevils (Acan-thoscelides obtectus)confined on live beanplants apparently fed on the parenchymaof the leaves, but he did not actually ob-serve feeding.

Leroi et al. (1984) found that forAcan-thoscelides obtectus,although larvae andadults utilize different types of food (thatis, seed endosperm vs. pollen grains),carbohydrate compositions of the two foodsources are remarkably similar. Glyco-sidase activity in the guts of larvae andadults are likewise similar.


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For an excellent discussion of bruchidphysiology and nutrition, see Johnson andKistler (1987).

Oviposition

Huignard et al. (1990) observed that mostBruchidae are specialists developing on a

limited number of host species and hostselection is accomplished by females ovi-positing on pods or fruits that are onlyavailable during a short period of the year.Neonate (newly hatched) larvae alone can-not search for their food substrate but de-pend on the female to locate suitable hostplants or seeds upon which to oviposit.

Bruchid eggs are usually either cementedto the pod or seed of a suitable larval foodplant, dropped into a pod after the female

cuts a hole in the pod wall, or tucked intosurface cracks or old emergence holes.

The adhesive used originates either in theepithelium of the egg follicles (Snodgrass1935) or in accessory glands (Wigglesworth1947) and not only cements the egg tothe substrate but also covers it. In somespecies, strands of adhesive resemblingguy wires extend from the edge of the eggcovering, apparently to aid in holding theegg to the pod surface as the pod expandsduring growth (Forister and Johnson 1970;

Pfaffenberger and Johnson 1976).

Johnson (1981a) classified egg placementinto three general types, or guilds:

1. Mature pod guild, in which eggs arecemented to the surface of the seedpod. Neonate larvae in this guild must

bore through the pod wall and intothe seed. (Examples:Acanthoscelideschiricahuae, Merobruchusspp., Mi-mosestesspp.)

2. Mature seed guild, in which eggsare cemented to the surface of seedsstill attached to the inside of a partlydehisced pod, so that larvae needonly bore through the seed integu-ment. (Examples: Stator limbatus, S.

pruininus.)

3. Scattered seed guild, in which eggs arelaid on mature seeds that have fallen tothe ground from a fully dehisced pod.(Examples: Stator sordidus, Zabrotesspp.)

Acanthoscelides obtectusfemales bite holesin green Phaseolusspp. pods through

which they insert eggs to fall freely withinthe pod. In later generations, they inserteggs in old emergence holes. Ovipositionfor this species begins early in the seasonon immature pods and may end on maturepods that have dropped to the ground oron naked seeds in storage.

Physiology

No attempt is made here to discuss physi-ological aspects of bruchids, but the follow-ing references will provide an introductionto the literature. Discussions ofAcanthos-celides obtectus, Callosobruchus chinen-sis, C. maculatus, Caryedon serratus,andZabrotes subfasciatusinclude additionalreferences on the subject. For a discus-sion of nutritional ecology of bruchids, see

Johnson and Kistler (1987).

Chromosomes.Takenouchi 1955, 1971a,1971b; Bawa et al. 1974; Smith and Brow-er 1974; Garaud and Lecher 1982; Garaud

1984.Digestion.Gatehouse et al. 1979; Choud-huri and Paul 1983; Gatehouse and Boul-ter 1983; Gatehouse et al. 1989.

Embryology.Brauer 1925; Tantawi et al.1976; Daniel and Smith 1994.

Enzymes.Sharma and Sharma 1979,1981; Gatehouse and Anstee 1983; Puriand Sharma 1984; Sharma and Rai 1984;Garg et al. 1990.

Egg production.Huignard 1970; Sandnerand Pankanin 1974; Hinton 1981; Choud-huri and Paul 1984; Dick and Credland1984; Butare and Bimont 1987.

Genetics.Breitenbacher 1925; Sano-Fujii1986; Messina 1987, 1990.

Lipids.Nwanze et al. 1976; Sidhu et al.1984.


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Nervous system.Ogijewicz 1948.

Overwintering.Brauer 1942; Hodek et al.1981.

Pheromones.Hope et al. 1967; Halstead1973; Rup and Sharma 1978; Tanaka etal. 1982; Qi and Burkholder 1985; Kitch

and Murdock 1986; Sakai et al. 1986.Temperature effects.Larson and Simmons1924a; Menusan 1934b, 1936; Menusanand MacLeod 1938; Sano 1967; Kistler1985.

Morphology of the Bruchidae

Adults

Little attention has been paid to the com-parative morphology of New World bruchid

adults. Descriptions have been writtenwith little thought to consistency of no-menclature of body parts, causing confu-sion when papers of various authors arecompared. Most references to morphologi-cal characters have been incidental in ge-neric or specific descriptions. Such papersas Doyens Skeletal anatomy of Tenebriomolitor(1966), Snodgrass 1908 and 1909papers on the insect thorax, or Hopkins

venerable but valid Contributions towarda monograph of the bark-weevils of the ge-

nus Pissodes(1911) are useful for generalreference but, of course, do not explainmorphological peculiarities of the bruchids.

Approximately 275 taxonomic papers havebeen written about Western Hemispherebruchids during the past 50 years, butonly two compared a structure of a bodypart (male genitalia) in several genera ofthis family (Teran 1967, Kingsolver 1970a).

Although nomenclature used in the twopapers differed, a basis for subsequent

studies was established. In addition tooffering a nomenclature for various sclero-tized parts, Kingsolver included a diagnosisof the genitalic musculature. No one hassurveyed the various genera for femalegenitalia, wing venation, elytral form, headform, legs, or antennae. Lukyanovich and

Ter-Minassian (1957) and Borowiec (1987)

incidentally included illustrations of struc-tures for several genera, including Ameri-can forms, but the field has been almosttotally neglected by American workers.

A moderately large resource of morpho-logical papers has recently emerged fromseveral Old World laboratories. Singh

(1973, 1978a,b, 1981a,c, 1982, 1983), forinstance, published on morphological char-acteristics of several Old World bruchidspecies. Many of the papers on Old World

bruchids are cited at the end of this sec-tion.

Johnson and Kingsolver (1973) introducedthe terms lateral, lateroventral, ventral, anddorsomesalto designate metatibial carinaein the genus Sennius.Here, I am modifyinglateroventralto ventrolateralto be consis-

tent with dorsomesal. These metatibialterms are valid for the majority of generain the Bruchinae; however, the positionsof the tibial carinae are different in thePachymerinae and in the Megacerini and

will be further elaborated in the descriptivesections for those taxa.

For the purposes of this handbook, onlyexternal morphology and male genitaliaand considered. A thorough morphologicalanalysis is sorely needed in order to makecomparisons between various sections ofthe family and to provide sets of charactersfor phylogenetic analyses. Only those bodyparts that can be seen without dissectionor dismemberment of the body (exceptmale and female genitalic parts, wings, andmouthparts) will be discussed and illus-trated here. One species is illustrated as a

basic reference, and important variationsare illustrated.

Because Bruchidae develop in a restrictedspacein most instances the excavated

interior of a single seedand must emergethrough a circular hole excavated by thelarva, evolution has favored a smoothlyrounded body lacking protuberances orspines that would hinder escape from theinterior of the seed. Most bruchids areovate or subelliptical in outline and gener-ally circular or oval in cross section. Legs


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and antennae are capable of being tuckedclose to the body, and the head hangs

vertically and usually rests on the proster-num. In many species, the dorsal profileis arched, which leads to a shortening ofsternal areas of the thorax and consequentreduction of sclerites in this part of the

body. In addition, the elytra are abbreviat-ed, exposing the heavily sclerotized seventhtergum (pygidium).

Head.The bruchid head at rest is eithervertical (hypognathous) or reflexed so thatit rests on the prosternum (opistho- gna-thus) (figure 17). In a frontal view (figures18 and 68), the head is obovate with thesubspherical base inserted into the ante-rior foramen on the front of the prothorax.In lateral aspect, the head tapers towardthe apices of the mandibles. Compoundeyes are usually protuberant but in somespecies follow the contour of the lateralmargin of the head.

Most of the sutures delimiting sclerites ofthe head in primitive beetles are lost inBruchidae, and only general areas can bedefined. The vertexis the area of the headabove the upper limits of the eyes, some-times set off by a transverse sulcus(figure18). Thefronslies between the eyes andsometimes carries a vertical, medianfron-

tal carina. The ventral margin of the fronsis the transversefrontoclypeal suture. Insome genera (for example, CaryobruchusandAmblycerus)thefrontogenal sutureextends from the inner corner of the eyeto the frontoclypeal suture separating thefrons from the cheek portion of the head(the gena) but in most bruchids this sutureis lost and the frons and gena are continu-ous beneath the eye. The dorsal portion ofthe gena carries the antennal socket.

The clypeusis a quadrangular sclerite,sometimes elongated, usually rounded

basally, and truncate apically. The fronto-clypeal suture limits it dorsally, the lateralmargins extend to the mandibles, and theapical boundary is the clypeolabral suture.

The clypeus partly covers the mandiblesand maxillae. The labrumis attachedto the apical margin of the clypeus and

usually carries a transverse row of longsetae.

The compound eye is surrounded by anarrow, setiferous channel, the supra-ocular sulcus(Kingsolver 1980a:230),

which may correspond to the ocular su-ture found in more generalized insects

(Snodgrass 1935:109) or may be second-ary.

The antennal socket is situated on theventral rim of the eye at the base of anemargination, the ocular sinus. The sinus

varies in depth from a slight emargination(as inAmblycerusand Caryobruchus) toan almost complete division of the eye (asin Meibomeus).The antennal socket car-ries on its ventral rim a small projection,the antennifer, forming an articular point

upon which the bulbous base of the an-tennal scapepivots. The scape is usuallylonger than the second segment, thepedi-cel(figures 613); theflagellumis invari-ably composed of nine segments and may

be uniformly serrate, gradually clavate,strongly eccentric, or even pectinate.

On the ventral side of the head, the eyesare prominent on either side of a flat,central submentum(figure 21). The latterarea is continuous with the gena in frontof and behind the eye and is limited be-hind by a deep, transverse sulcus. At theposterior end of the head is a large open-ing, the occipital foramen, through whichthe digestive and nervous systems passinto other parts of the body. On the ventralpart of the rim of the foramen is a quadrateor trapezoidal sclerite, the gula. Along thelateral margins of the gula are theposteriortentorial pits, which are the attachments ofthe internal skeleton of the head. Attachedto the anterior margin of the submentum

is the mentum(figure 21), an emarginate,transverse sclerite to which thepremen-tumis attached. The prementum is the

base for the paired, three-segmented labialpalpiand the ligula, which is divided intopairedparaglossae. On either side of thementum and submentum is a deep emar-gination, the maxillary fossa, into whichthe maxillais inserted. The maxilla (figure19) is composed of five parts: the cardo,


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the stipes, the three-segmented maxillarypalpus, the galea, and the lacinia. Boththe galea and the lacinia are provided withlong curved setae, which, in some species,are plumose. These specialized setae areapparently used to rake pollen into the oralcavity. The inner surface of the labium and

of the labrum is also provided with rows ofsetae to assist in ingesting pollen.

The mandiblesproject beyond the labrum,but their attachment is on each side ofthe head with the dorsal pivots, or articu-lations, at each end of the frontoclypealsuture and the ventral articulations at thelateral corner of the maxillary fossa. Theapex of each mandible is acute, and theinner margin forms a sharp, entire cuttingedge. In some species, the inner marginis fringed with long setae. A membranousflap, theprostheca, and a flattened platenear the base of the mandible, the mola(figure 20), are characteristic of all Bruchi-dae. The face of the mola is covered withshort spines to form a grinding surface.

Prothorax.Theprothorax, the first leg-bearing segment, is composed of twosclerites: theprosternum(figures 22, 23,and 41) is a Y-shaped sclerite bounded

by the front coxal cavities, thepleuroster-nal suture, and the anterior foramen, and

thepronotumis a large, continuous dor-sal sclerite making up the remainder ofthe body of the prothorax (figure 41). Thepronotum consists of a dorsal surface,the disk, and the hypomeron(the lateral,concave area dorsal to the coxa). The diskis limited in some species by a distinctlateral carinabut more often is continuouslaterally with the hypomeron itself, markedonly by the concave surface. The epimeron,an extension of the hypomeron but not aseparate sclerite, projects behind the front

coxae to meet the apex of the intercoxalpiece.

In some species in the Pachymerinae andAmblycerinae, the margin of the pronotaldisk is distinctly sulcate and corners ofthe disk bear one to three setiferous punc-tures.

The prosternum is usually short in front ofthe procoxae because of the ventral com-pression of the body. In Spermophagus,anOld World genus, it is reduced to a mem-

branous bridge. The prosternal processmay be flat as in Caryobruchusand some

Amblycerus(figure 41) or narrow as in

otherAmblycerus(figure 45), or it may bean internal, vertical lamella partly hidden

by the coxae (Acanthoscelides).

The attachments of the fore and middlecoxae generally take two forms in bru-chids. In the generalized form (as inAmbly-cerusand Caryobruchus),the coxa freelyarticulates with the trochantin, which inturn articulates with the body wall (figure23). In the more advanced form (such as

Acanthoscelidesand Sennius), the trochan-tin is replaced by a straplike extension ofthe sternum forming a socket that allowsthe coxa to rotate freely through about 90degrees of arc but limits lateral movementof the leg (figure 24). The triangular tro-chanter is rigidly attached to the basal endof the femur.

Mesothorax and Metathorax.The twowing-bearing segments in Bruchidae arerather rigidly fused together to providestrength. Internal sutural ridges (phragma,or ingrowths from the body wall) provide

attachment points for the massive musclesrequired to operate the mesothoracic ely-tra, the metathoracic flight wings, and thelegs. Only the external sclerites, however,

will be identified here.

Each of the thoracic segments is dividedinto a ventral sternal region, a lateralpleu-ral region, and a dorsal notal region. Exceptfor the apex of the scutellum protruding

between the bases of the elytra (figures 30and 60), the notal sclerites are concealed

beneath the elytra and have not beenincluded in descriptions of bruchids. The

flight wings, concealed beneath the elytra(figures 14 and 15), are folded longitudi-nally and transversely.

The mesothoraxis the shortest of the threethoracic segments. It consists of three vis-ible platesthe mesosternum, the mesepis-


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ternum, and mesepimeron(figure 28). Thevisible mesosternum is a Y-shaped platelying in front of and between the middlecoxae and is separated laterally from the

ventral end of the mesepisternum by thesternopleural suture. The intercoxal processrests on the corresponding extension of the

metasternum. The mesepisternum and mesepimeron are adjoining

vertical plates extending from the elytralmargin to the coxal cavity. In more gener-alized bruchids (figure 25) (Caryobruchus,

Amblycerus),these two plates are approxi-mately the same size, but in more special-ized forms the mesepimeron is reduced insize and shape by the encroachment of themesepisternum (figures 26 and 27).

The coxal cavitylies between the metaster-

num, the mesosternum, and the ventralend of the two pleural sclerites. The attach-ment of the leg is essentially the same asdescribed for the prothorax.

The form of the elytra in beetles is a radicalmodification of the metathoracic wings toform a protective cover for the notal areasof the mesothorax, the metathorax andabdomen, and the flight wings. At rest, theelytra meet on the midline with a tongue-and-groove interlock mechanism. Duringflight, the elytra are rotated laterally on

the mesopleural wing process to allow theflight wings to unfold and operate freely.

Although the elytra in bruchids exhibitmany modifications of surface sculptureand configurations of the striae, especiallyin the basal area, 10 striae are alwayspresent.

Venation of the flight wings throughoutthe family is not detailed here (except infigures 14 and 15). The wings of only afew species have been illustrated, and a

thorough survey of venation in the familyhas not been made. References to papersthat include bruchid wing patterns can befound at the end of this section.

The metathoraxis considerably largerthan the mesothorax to accommodate the

volume of muscles that operate the flightwings and legs. The size and relative posi-

tion of the sternal and pleural sclerites isdistinctly different from that of the me-sothorax. The metasternumis rigidly at-tached to the mesosternum and to thehorizontal metepisternum(figure 28) andcarries the median articulation for themetacoxa as well as the attachment of the

abdomen. It also forms the posterior wall ofthe mesocoxal cavity and the anterior wallof the metacoxal cavity. Thepleurosternalsutureseparates it from the metepister-num. Thepostmesocoxal sulcusfollows theposterior margin of the mesocoxal cav-ity and in some genera bends sharply toparallel the pleurosternal suture (parasu-tural sulcus),but in other, more specializedgenera it ends at or near this suture. Alongthe anterior border of the metacoxal cav-ity extends a narrow, transverse sclerite

variously termed the antecoxal pieceor theprecoxal strap.

The metepisternumis an elongated, hori-zontal sclerite, set between the mesepi-meron and the dorsal one-half of themetacoxal cavity, and carries the dorsalarticulationof the metacoxa. In some moregeneralized genera, the face of this sclerite

bears an angulate sulcusthe metepister-nal sulcus(figure 28)which extends to-

ward and bends to parallel the pleuroster-

nal suture and forms the correspondingparasutural sulcusto that of the metaster-num. This sulcus is not present in morespecialized genera. The metapleural sutureseparates the metepisternum from the nar-row, horizontal metepimeronthat, exceptfor the posterior end, is hidden beneath themargin of the elytra.

The external face of the hind coxais anelliptical or reniform plate rotating on atransverse axis between the ventral coxalarticulation and the dorsal articulation.

The coxa, however, is roughly triangularin cross section and projects into a cavity

between the metasternum and the abdo-men. The coxa can rotate on its axis about30 degrees, and the trochanter can rotatein its fossa near the ventral end of the hindcoxa more than 180 degrees. Extendinglaterad from the trochanteral fossa is a


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Modifications of the visible sterna includea tuft of setae or a pit on the basal ster-num of males in some genera, the terminalsternum in many genera being more deeplyemarginate in males than in females, orthe abdominal segments being telescoped.

The membranous pleural areas of the

abdomen that carry the spiracles for eachsegment are hidden beneath the elytra,except that the spiracle for the 7th tergum(the pygidium) is located at the extremeupper corner of the tergum. The pygidiumin this family is always exposed beyond theapices of the elytra and is always heavilysclerotized; however, in Kytorhinusthe 5thand 6th terga are also sclerotized (figure51). The sculpture, shape, and setationof the pygidium is often a good source ofgeneric and specific characters. The malepygidium is often more convex and inflexedapically than is that of the female.

Genitalia.Zacher (1930) was the firstworker to explore the usefulness of malegenitalia in the classification of bruchids,

but his descriptions of the various partswere meager and poorly illustrated. Muker-ji and Chatterjee (1951) compared genitaliaof species in five genera and identified the

various parts using the nomenclature ofSnodgrass (1935). Teran (1967) illustrated

genitalia of species in nine additional gen-era but used a different set of morphologi-cal terms. Kingsolver (1970a) made a studyof male genitalia from an evolutionarystandpoint, including the identification ofgenitalic muscle strands and a theoreticalexplanation of the mechanism for evertingthe internal sac. His nomenclature, dif-ferent from that of Mukerji and Chatterjeeor that of Teran, followed Sharp and Muir(1912) and has generally been employed byrecent workers in the family. Most of the

taxonomic papers published since 1960use and illustrate male genitalia for speciesdiscrimination. Several papers have re-cently been published on male genitalia ofOld World species of Bruchidae. These arereferenced at the end of this section.

thin suture usually bordered by a polished,impunctate strip. I am here applying theterm fossula to this suture (figure 28).

The trochanteris more or less rigidly at-tached to the proximal end of thefemur(figure 29). The latter segment of the hindleg is laterally compressed and often dor-

soventrally expanded. The ventral marginis often furnished with denticles of varioussizes and numbers (figures 40, 47, 49, 50,53, and 6365), a primary source of sub-family and generic characters and, in someinstances, specific characters. Despitethe obvious resemblance to the saltato-rial, or jumping, hind legs in other fami-lies and orders of insects (such as fleasand fleabeetles), legs of bruchids are notequipped for jumping. The development ofthe internal musculature is opposite thatfor jumpers; that is, the greatest develop-ment of muscle bands is in those that flexthe metatibia against the ventral margin ofthe metafemur. The hind leg thus becomesa grasping organ, probably for holding toedges of leaves or perhaps used in mating.See Furth and Suzuki (1990).

The proximal end of the metatibia hingesbetween two terminal plates of the metafe-mur and has a range of movement of about120 degrees in the same plane as the

metafemur. Terminal spurs are found inonly a few genera, the most common termi-nal armament being a ventral spine (mucro)and several smaller terminal denticles (co-rona) (figures 44, 47, 49, 50, 53, and 54).

Abdomen.The abdomenin Bruchidaeconsists of five externally visible, heavilysclerotized sternal segments, but the basalsternum is anatomically the 3rd segment,the 1st and 2nd sternites being reduced tolightly sclerotized or membranous relicts

forming the vertical anterior wall of theabdomen behind the metacoxae. For con-venience, the visible sternal segments arenumbered from 1 to 5. The terminal seg-ments, including those transformed intosex organs, are radically modified and are

withdrawn into the end of the abdomen.


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Little has been published in the UnitedStates on the female genitalia of Bruchi-dae. See below for a discussion of thesestructures.

Male Genitalia.Male genitalia provide themost consistent source of generic and spe-cific identifying characters in this family.

Moreover, the combination of character-istics of the genital complex is sufficientlydistinctive to warrant separation of thefamily from other Chrysomeloidea.

The male reproductive organs are of bothmesodermal and ectodermal origin. Thetestes and vasa deferentia originate in themesoderm, whereas the ejaculatory duct,accessory glands, and intromittent organsare ectodermal. Ectodermal parts containchitin and will withstand treatment in

sodium hydroxide to remove muscle fibers,whereas endodermal parts will dissolve inthe same solution.

As is true of most Coleoptera, the medianlobe, lateral lobes, internal sac, and associ-ated parts are completely invaginated intothe end of the abdomen and are normallynot visible without dissection. To examinedetails of the various components, caustictreatment is usually necessary to removeextraneous body tissues (see Materials andMethods). Significant characters for spe-cific and sometimes generic identificationare usually found in the parts resistant tothe caustic solution.

Male genitalia of bruchids are modifiedfrom the cucujoid type (Crowson 1955)

with a basic form similar to those found inthe phytophagous families Cerambycidae,Chrysomelidae, and Curculionoidae. Thenearest approach of bruchids to anotherfamily appears to be that of the Pachym-erine genera Caryoborusand Caryobruchus

to the Sagrine genus Carpophagus.

The genital complex comprises three prin-cipal parts (figures 3034):

The tegmen(figure 31) comprises thelateral lobes(parameres), the basal

piece(tegmen ring) surrounding themedian lobe, and the ventral strut.

The median lobe(aedeagus) (figure 30)is a sclerotized tube with an enlarged,spoon-shaped base (cucullus) modi-fied from ventral struts found in relatedfamilies.

The eversible internal sac(figures 30and 34) normally lies retracted inside

the median lobe and is attached to theapical orifice of the median lobe.

These various parts are highly modifiedderivatives of the terminal segments of theabdomen and are enclosed in the elongateintersegmental membranes of the 7th, 8th,and 9th segments.

The apical orifice(figure 30a), which is theattachment for the base of the internal sac,is located near the apex of the median lobe.

The basal orifice(figure 30b) is located at

the point at which the basal piece sur-rounds the median lobe. Although the rigidapex of the median lobe of the bruchidsubfamilies Pachymerinae and Rhaebinaeis similar to that found in the Sagrinae andthe Cerambycidae, the apex in the Am-

bly-cerinae, Kytorhininae, and Bruchinaehas developed into a separate sclerite, theventral valve(figure 30c). Some groups of

bruchids also have a sclerotized or largelymembranous, hoodlike dorsal valve(figure30d) above the apical orifice. The configu-ration of the ventral valve is often one ofthe specific diagnostic characters.

The lateral lobes are attached to the dorsalrim of the basal piece and lie on the dorsalside of the median lobe. They take variousforms, sometimes fused into an apically

bilobed straplike structure, or are vari-ously divided, sometimes nearly to theirattachment with the basal piece. The basalpiece surrounds the middle of the medianlobe and is attached to the ventral rim of

the basal orifice by a sclerotized mem-brane. The tegminal strut extends beneaththe cucullus and may be flat or keeled.Between the rim of the cucullus and thetegminal strut extend thick muscle bands(figure 32) bridging the space between thetwo parts. The membrane between the

basal piece and basal orifice effectively


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seals the cavity formed by the musclebands, and compression of the musclesconverts the structure into a pump to evertthe internal sac during copulation.

Associated sclerotized structures are theY-shaped, ventrally positioned 8th sternite(spiculum gastrale) (figure 32), to which

muscle bands that move the genital com-plex in and out of the body are attached,and the modified 8thtergite, a hood-shaped sclerite lying beneath the pygidiumand dorsad of the lateral and median lobes.

The internal sacis the primary intromit-tent organ (figures 30, 33, and 34). It isattached at the apical orifice near the apexof the median lobe and is essentially anextension of that lobe. For that reason,the terminology applied to the internal

sac seems contradictory but is logical if itis remembered that it is the median lobeextended. When the sac is invaginated

within the median lobe in repose, its baseis nearest the apex of the median lobe andits apex is nearer the base of the medianlobe (figure 30). Inflation of the sac is ac-complished through pumping action of the

basal pump. Various sclerites(spicules,spines, denticles, serrata)(figure 35) at-tached to the sac emerge on the externalsurface of the sac as it is evaginated into

the vaginal tract and undoubtedly act asholdfasts to keep the sac in place duringcopulation. The arrangement of this saccalarmature is, in many instances, the sourceof characters for specific identification.Because of the difficulty of everting the sacof prepared specimens, illustrations of themale genitalia usually show the sac in theinvaginated position.

The vas deferens(figure 32) extendsthrough the muscle strands of the pump

and is attached to the closure valve, ortransfer valve, of the internal sac. Duringinflation of the sac, it follows the sac apexthrough the lumen of the median lobe andfinally through the lumen of the sac.

Leopold (1941) discussed the role of maleaccessory glands in reproduction.

Withdrawal of the sac from the vagi-nal tract begins at its apex. Fine musclestrands attached to saccal sclerites andinserted on the internal walls of the medi-an lobe contract to effectively peel the sacfrom inside the vagina and restore it to itsoriginal position inside the median lobe.

Female Genitalia.Female genitalia gener-ally have not been illustrated nor describeddue to their apparent lack of taxonomicsignificance. This facet of bruchid mor-phology, however, should be more fullyexplored for potential species group orgeneric characters.

Kingsolver and Johnson (1978) publishedillustrations of ovipositors for North Ameri-can species of Mimosestes,and theirs isthe only American paper illustrating fe-

male genitalia for all of the species treated.No one has made a comparative studyof female genitalia for the Bruchidae inthis country. Nilsson and Johnson (1990)included drawings for two species of Caryo-bruchus,and Donahaye (1974) sketchedgenitalic parts for seven species of Bru-chus.Mukerji and Bhuya (1937) detailedthe reproductive system of both Calloso-bruchus maculatusand C. chinensis.Tan-ner (1927) includedAcanthoscelides(asMylabris) obtectusin his survey of female

ovipositors of Coleoptera. Johnson et al.(1989) included various female charactersin New World Stator.Several papers detail-ing ovipositors and internal reproductiveorgans have been published for Old World

bruchids, including those for BruchidiusmackenzieiKingsolver from Australia (fig-ure 36), and for several species of Bruchid-iusby Iablokoff-Khnzorian and Karapetian(1973). References to these papers arelisted at the end of this section.

The female genital complex, as in the male,has its origin in both the mesoderm andthe ectoderm. Products of the mesodermare the ovaries, the lateral oviducts, andperhaps part of the common oviduct. Ofectodermal origin are the vagina, the bursacopulatrix, the spermathecaand spermath-ecal glands, part of the common oviduct,


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and the ovipositor. These are shown infigure 36.

In certain groups of bruchids (such asAc-anthoscelidesand Callosobruchus), a pairof structures first described by Mukerjiand Bhuya (1937:200) are probably glan-dular in function and are positioned one

on each side of the vaginal tube at the en-trance of the bursa copulatrix (figure 36).

These vary from being merely convex evagi-nations into the lumen of the vagina tooviform, membranous sacs nearly occlud-ing the lumen. Their attachment is ringlike

with a central opening through which passtracheae into the gland. These are lackingin Bruchus,but a pair of glands are posi-tioned at the anterior end of the bursa. Thefunction of these structures is not known.

A family-wide survey of their position andstructure may offer phylogenetic clues.

Papers on Adult Morphology

Head morphology.Stickney 1923; Ferris1942; Tandon 1960; Mathur and Dhadial1961, 1963; Singh 1981c, 1982, 1983.

Brain.Satija et al. 1975; Sandhu andNeena 1982.

Eyes.Schmitt et al. 1982.

Thorax.Snodgrass 1909; Hlavac 1972.

Metendosternite.Crowson 1938, 1944.

Wings.Forbes 1922; Kempers 1923; Mar-cu 1939; Chen 1940; Seeliger 1943; Jolivet1957; Suzuki 1969; Wallace and Fox 1975,1980; Singh 1981a.

Alimentary canal.Kasap 1978b; Mannand Crowson 1983a.

Ventral nerve cord.Kasap 1978a; Mannand Crowson 1983b.

Sense organs.Pouzat 1981.

Tracheal system.Pajni 1969.

Reproductive systems.Kannan 1923;Tanner 1927; Zacher 1932; Zia 1936;Mukerji and Bhuya 1937; Mukerji 1949;Mukerji and Chatterjee 1951; Srivas-tava 1953a,b; De Luca 1959; Pajni 1959;

Surtees 1961; Iablokoff-Khnzorian 1966,1967; Teran 1967; Huignard 1968; King-solver 1970a; Arora 1971; Singh 1973;Iablokoff-Khnzorian and Karapetian 1973;Spirina 1974; Ahmed et al. 1976; Leopold1941; Thukral 1976; Pawar and Verma1977; Singh 1978a,b; Cassier and Huig-

nard 1979; Singh 1979; Ahmad and Murad1980a,b; Kasap and Crowson 1980; Kumarand Verma 1980; Monga and Sareen 1980;Hamon et al. 1982; Singh 1982; Crowson1984; Kasap and Crowson 1985; Kasapand Crowson 1988.

Larval Characters

The destructive feeding period in theBruchidae occurs during the larval stages,

yet bruchid larvae are relatively poorly

known and efforts to improve on theiridentification are in their infancy.

G.S. Pfaffenberger, who has contributedmore to the study of bruchid larvae thananyone else in the United States, kindlysupplied the following description of larvalcharacteristics.

Bruchid larvae have four instars, the firstbeing significantly different from the oth-ers (figures 37 and 38). Body sizes of firstinstars of various species vary significantly

but are characterized by the presence of asclerotized X or H-shaped prothoracic plate(figure 16) and a sclerotized abdominalspine immediately above the spiracle of thefirst abdominal segment. These structuresare used to gain leverage to penetrate theintegument of the seed or pod. Append-ages, when present, are highly variable inlength and form.

Integument.Cream colored except pro-notum yellowish brown. Chaetotaxy of the

first instar may consist of noticeably longor short sensilla, but they are distributedin recognizable patterns.

Head.Hypognathous and retractable; yel-lowish brown with mouthparts more darklypigmented.

Antenna.With one to three telescopic seg-ments. A single elongate sensillum chaeti-


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cum and at least one sensillum basiconi-cum located distally (Pfaffenberger 1985b).

Clypeolabrum.Clypeal portion with singlesensillum trichodeum near each lateral

border, subtended medially by sensillumampullaceum. Labral portion rounded dis-tally, single or double arc of sensilla tricho-

dea located distally among mat of variablelength microtrichia; lateral sensillum trich-odeum and medial sensillum ampullaceumlocated laterally near proximal margin oflabrum (Pfaffenberger 1985b: figure 34).

Epipharnyx.Epipharyngeal groove bor-dered laterally by one to three pairs ofdecurved sensilla trichodea.

Mandible.Monocondylic, with gouge-shaped mesal face.

Maxilla.Cardo, stipes, palpifer, palpus,and lacinia present. Membranous stipeswith variable num

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